Proceedings of the California Academy of Sciences, 4th series

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PROCEEDINGS

OF THE

California Academy of Sciences

FOURTH SERIES

Vol. XXXIX

SAN FRANCISCO
PUBLISHED BY THE ACADEMY
1972-1974

COMMJTTEE ON PUBLICATION
GrorcE E. Lrnpsay, Chairman

Diana R. Youne, Editor

No.

No.

No.

No.

No.

No.

alle

lz.

CONTENTS OF VOLUME XXXIX

HERTLEIN, LEo G. Description of a new species of
Chlamys (Mollusca: Pelecypoda) from the Galapagos
Nietegrmd pee 908 So 0 5 th a a ie Pe Oh Bn

SmitH, ALLYN G. Three new land snails from Isla Santa
Cruz (Indefatigable Island), Galapagos -_______________

HERTLEIN, Leo G. Pliocene fossils from Baltra (South
Seymour) Island, Galapagos Islands ——_______

ABE, TOKIHARU, and WILLIAM N. ESCHMEYER. A new
species of the scorpionfish genus Helicolenus from the north
apiCHOccahee tt AONE Pe ood ee Neat oe eee

ESCHMEYER, WILLIAM N., and K. V. Rama Rao. Two new
scorpionfishes (genus Scorpaenodes) from the Indo-west
Pacific, with comments on Scorpaenodes muciparus
(Phe encica) eee iets, Bee 2 AR BE

ZULLO, Victor A., DEA B. BEACH, and JAMEs T. CARLTON.
New barnacle records (Cirripedia, Thoracica) —...__

CAsTEEL, RicHarp W. A key; based on scales, to the
families of native California freshwater fishes

AsHLOCK, PETER D. The Lygaeidae of the Galapagos
islands< (Hemiptera; «Eleteroptera). 2d

Brown, WALTER C., and Marjorie V. C. FALANRUW. A
new lizard of the genus Emoia (Scincidae) from the
ianataiicse USGS 9 aoe Mes ee a

McCoskeEr, JOHN E. Two new genera and two new species
of western Pacific snake-eels (Apodes: Ophichthidae)

HERALD, Ear S., and JoHN E. RANDALL. Five new Indo-
Ge SIC EISINC Sie eres en eee

CHEMSAK, JOHN A., and E. G. Linstey. The genus Mecas
ieConte (Colecptera:Cerambycidae) 2. ==

Pages

47-53

55-64

65-74

75-86

87-103

105-110

111-120

121-140

141-184

No. 13.

No. 15.

No. 19.

Now ZL.

No. 23

SMITH-VANIZ, WILLIAM F., and Jon C. STAIGER. Compar-
ative revision of Scomberoides, Oligoplites, Parona, and
Hypacanthus with comments on the phylogenetic position
of Campogramma (Pisces: Carangidae)

ComPAGNoO, L. J. V. Ctenacis and Gollum, two new genera
of sharks (Selachu? Carcharimidat) ">" eae

MAnpRA, York T., A. L. BRIGGER, and H1GHooHT MANDRA.
Chemical extraction techniques to free fossil silicoflagellates
from marine’ sedimentary rocks ==" =" ee

ESCHMEYER, WILLIAM N., YosuHitsuGu Hrrosakti, and
TOKIHARU ABE. Two new species of the scorpionfish genus
Rhinopias, with comments on related genera and species —___

GarTH, JOHN S. The brachyuran crabs of Easter Island _

ESCHMEYER, WILLIAM N., and Kaza V. Rama Rao. Two
new stonefishes (Pisces: Scorpaenidae) from the Indo-west
Pacific, with a synopsis of the subfamily Synanceiinae _____

Compacno, L. J. V. Gogolia filewoodi, a new genus and
species of shark from New Guinea (Carcharhiniformes:
Triakidae), with a redefinition of the family Triakidae and
a key*to Triakid ‘genera’ 4° eee

Howarp, L. D. Muscular anatomy of the forelimb of the
sea otter (Enhydra lutris) i... Oe ee

BURGHARDT, GLENN FE. A _ new Hawaiian chiton
Rhyssoplax linsleyi (Mollusca: Amphineura: Chitonidae) —

Iwamoto,Tomio. Nezumia (Kuronezumia) bubonis, a new
subgenus and species of grenadier (Macrouridae: Pisces)
from Hawaii and the western North Atlantic —

SCHRADER, HANS-JOACHIM. Revised diatom stratigraphy
of the Experimental Mohole Drilling, Guadalupe Site

Index to: Volume: XOX XIX$ 2) es eee

185-256

257-272

273-284

285-310

311-336

337-382

383-410

411-500

501-506

507-516

517-562
563-580

PROCEEDINGS

OF THE

CALIFORNIA ACADEMY OF SCIENCES

FOURTH SERIES
Vol. XXXIX, No. 1, pp. 1-6; 5 figs. January 21, 1972

DESCRIPTION OF A NEW SPECIES OF CHLAMYS
(MOLLUSCA: PELECYPODA) FROM THE
GALAPAGOS ISLANDS

By
Leo G. Hertlein

INTRODUCTION

The known marine molluscan fauna of the Galapagos Islands in general
is sparser than that of the mainland. Eleven species of scallops (Pectinidae) have
been reported living in the Galapagos Archipelago. These are: Pecten (Flabelli-
pecten) sericeus Hinds, Pecten (Oppenheimopecten) galapagensis Grau, Chlamys
(Chlamys) lowei Hertlein, Chlamys (Argopecten) circularis Sowerby, Chlamys
(Nodipecten) magnifica Sowerby, Cyclopecten (Cyclopecten) exquisitus Grau,
Cyclopecten (Cyclopecten) pernomus Hertlein, Cyclopecten (Delectopecten)
polyleptus Dall, Cyclopecten (Delectopecten) zacae Hertlein, and from very
deep water, Cyclopecten (Cyclopecten) liriope Dall and Cyclopecten (Hyalopec-
ten) neoceanicus Dall. Seven of these are known to occur along the adjacent
mainland.

In 1969, Mr. Anthony D’Attilio, San Diego Society of Natural History, sent
six paired valves (four with the animal) and four single valves representing a
species of Chlamys to Allyn G. Smith, Department of Invertebrate Zoology,
California Academy of Sciences, with a request for identification of the species.
These subsequently were submitted to me for study. A search of the Academy’s
collection as well as of the literature failed to reveal any similar species described
from the eastern Pacific. Upon receipt of this information Mr. D’Attilio re-
quested me to describe the species which appears in the present paper.

[1]

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2 CALIFORNIA ACADEMY OF SCIENCES [PRroc. 4TH SER.

ACKNOWLEDGMENTS

The writer here expresses his thanks to the following persons: to Mr.
Anthony D’Attilio, San Diego Society of Natural History, for permission to
describe the new species and to retain paratypes; to Mr. Allyn G. Smith, Cali-
fornia Academy of Sciences, for advice concerning the specimens; to Mr. Barry
Roth of the same institution for critical reading of the manuscript and for
arrangement of the illustrations; to Dr. Thomas R. Waller, Division of Inverte-
brate Paleontology, United States National Museum, for information concerning
comparative species in the collections of the National Museum; to Mr. Maurice
Giles, Staff Photographer, California Academy of Sciences, who prepared the
photographs from which the illustrations were made.

DESCRIPTION OF NEW SPECIES

Family PECTINIDAE Rafinesque
Genus Chlamys Roding in Bolten

Chlamys (Chlamys) incantata Hertlein, new species.
(Figures 1-5.)

Dracnosis. A species of Chlamys differing from other west American species
in possessing very narrow, compressed, rather low, spinose ribs.

DeEscriIPTION. Shell averaging about 45 mm. in height, ovate, valves gently
and nearly equally inflated, hinge line rather short. Right valve sculptured with
about 25 major ribs which are narrow, compressed, rather low, and occasionally
unequally spaced; on top of each of these ribs is a row of spines which are con-
cave ventrally; submargins with 5 or 6 very fine riblets; interspaces vary in
width but are much wider than the ribs, nearly flat-bottomed, and sculptured
with 1 to 3 fine radial threads, each bearing a row of spines, the ribs and inter-
spaces crossed by fine concentric imbricating lines of growth; auricles unequal,
the anterior one the larger, sculptured with about 5 spiny radial riblets, the
hinge line above the auricle with scaly sculpture, below the auricle a well
developed byssal notch about half the length of the auricle, below this along the
margin there are 4 pectinidial teeth; posterior ear short, slightly concave, sloping
rather steeply downward, sculptured with about 6 radial riblets. Left valve
sculptured similar to the right but lacking a byssal notch. Hinge with one pair
of slight cardinal crura. Interior of valves lightly grooved corresponding to the
external ribbing. Color of the exterior of the valves whitish, the spines rosy,
lending a roseate appearance to the valves, more pronounced on the left one.

Dimensions. Length 40 mm., height 45 mm., length of hinge line 21 mm.,
convexity (both valves together) approximately 13 mm. The largest specimen,
a left valve, is 48 mm. high.

TYPE MATERIAL. Holotype no. 52263, also paratypes, San Diego Society of
Natural History, from off Academy Bay, Santa Cruz (Indefatigable) Island,

VoL, XXXIX] HERTLEIN: A NEW SPECIES OF CHLAMYS 3

Ficure 1. Chlamys (Chlamys) incantata Hertlein, new species. Holotype no. 52263
(San Diego Society of Natural History), from off Academy Bay, Santa Cruz (Indefatigable)
Island, Galapagos Islands, dredged in 200 meters. View of the exterior of the left valve;
height 45 mm. Ficure 2. Chlamys (Chlamys) incantata Hertlein, new species. View of the
exterior of the right valve of the holotype. Ficure 3. Chlamys (Chlamys) incantata Hert-
lein, new species. Paratype (SDSNH), from the same locality as the holotype. View
of the exterior of a left valve; height 48 mm. Ficure 4. Chlamys (Chlamys) incantata
Hertlein, new species. Paratype (SDSNH), from the same locality as the holotype.
View of the exterior of a right valve; height 39.5 mm. Ficure 5. Chlamys (Chlamys)
incantata Hertlein, new species. View of the interior of the right valve shown in figure 2.

Galapagos Islands; dredged in 200 meters; Mrs. Jacqueline DeRoy collector,
July 25, 1969.
Paratypes are deposited in the California Academy of Sciences, the Amer-
ican Museum of Natural History, and in the United States National Museum.
CoMMENTS. The shape, spinose sculpture, and rosy coloration of this new

4 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

species are somewhat similar to those of Chlamys hastata Sowerby (see Arnold,
1906, pl. 42, figs. 1, la, 2, 2a; Grau, 1959, plates 27, 28) from California, but
the ribs are more numerous, lower, more compressed, and not arranged in pairs
on the right valve.

The fewer ribs (25) on the new species serve to separate it from Chlamys
amandi Hertlein (1935, p. 305; Pecten australis Philippi, 1845, p. 56 “Patriae:
Insulae Chonos.”; not Pecten australis Sowerby, 1842) from Chile, which has 30
to 34 ribs which are only slightly scaly toward the submargins.

The sculpture of C. incantata, new species, bears a general resemblance to
that of the species described as Pecten (Chlamys) coccymelus by Dall (1898, p.
741, pl. 34, fig. 1) from strata of middle Miocene age at Plum Point Maryland.
Dall called attention to the similarity of the fossil form to juvenile Pecten
madisonius Say, but believed it to be a distinct species. Mansfield (1936, p. 177)
stated that it “may be an immature specimen representing a varietal form of
P. madisonius,’ and Rowland (1936, p. 1008) suggested that it ‘““may be a case
of arrested development.”

The sculpture of the new species described here bears a general similarity
to that of illustrations of Pecten (Chlamys) nympha Bavay (1906, p. 246, pl.
7, figs. 3 and 4). The type specimen of that species was described as only about
15 mm. high. The type locality was given as “Habitat Caribaeum Mare?”. The
type specimen in the Muséum National d’Histoire Naturelle de Paris, was
originally in a carton which also contained a specimen of Pecten antillarum
Récluz, a typical inhabitant of Caribbean waters. Dr. Thomas R. Waller
(written communication, December 30, 1969), who has given considerable time
to a study of the Pectinidae of the western Atlantic, stated that he feels certain
that Pecten (Chlamys) nympha is a synonym of Chlamys benedicti Verrill and
Bush (in Verrill, 1897, p. 74; not Pecten benedicti Lamarck, 1819) described
from “off Martha’s Vineyard, 1356 fath., dead; West Indies, in 25 to 72 fath.,
living.” Adult specimens of C. benedicti (Weisbord, 1964, pl. 14, figs. 8-11)
are quite distinct from the new species described here from the Galapagos Islands.

The specific name of this new species, “cantata,” is derived from the
vernacular appellation, “Las Islas Encantadas” (the Enchanted Islands), some-
times applied to the Galapagos Islands.

LITERATURE CITED

ARNOLD, R.
1906. The Tertiary and Quaternary Pectens of California. United States Geological
Survey, Professional Paper no. 47, pp. 1-264, pls. 1-53, 2 figs. in text.
Bavay, A.
1906. Sur quelques especes ou variétés nouvelles du genre Pecten. Journal de Conchyli-
ologie, vol. 53, no. 3, pp. 243-247, pl. 7, February 20.
Dati, W. H.
1898. Contributions to the Tertiary Fauna of Florida with especial reference to the
Silex beds of Tampa and the Pliocene beds of the Caloosahatchie River including

Vor. XXXIX] HERTLEIN: A NEW SPECIES OF CHLAMYS 5

in many cases a complete revision of the generic groups treated and of their

American Tertiary species. Transactions of the Wagner Free Institute of Sci-

ence of Philadelphia, vol. 3, pt. IV, pp. I-VIII, 571-947, pls. 23-35, April.
Grau, G.

1959. Pectinidae of the eastern Pacific. Allan Hancock Pacific Expeditions, vol. 23, pp.
I-VII, 1-308, pl. 1-57, September 25.

HERTLEIN, L. G.

1935. The Templeton Crocker Expedition of the California Academy of Sciences, 1932.
No. 25. The Recent Pectinidae. Proceedings of the California Academy of
Sciences, Fourth series, vol. 21, no. 25, pp. 301-328, pls. 18-19, September 26.

MANSFIELD, W. C.

1936. Stratigraphic significance of Miocene, Pliocene, and Pleistocene Pectinidae in the
southeastern United States. Journal of Paleontology, vol. 10, no. 3, pp. 168-
192, 1 text fig., pls. 22-23, April.

Rowranp, H. I.

1936. The Atlantic and Gulf Coast Tertiary Pectinidae of the United States. II. Sys-
tematic Descriptions, continued. American Midland Naturalist, vol. 17, no. 6,
pp. 985-1017, pls. 5-10, November.

VerRRILL, A. E.

1897. A study of the Family Pectinidae, with a revision of the Genera and Subgenera.
Transactions of the Connecticut Academy of Sciences, vol. 10, pt. 1, pp. 41-95,
pls. 16-21, June.

WEIsBorD, N. E.

1964. Late Cenozoic Pelecypods from Northern Venezuela. Bulletins of American

Paleontology, vol. 40, no. 204, pp. 1-564, pls. 1-59, February 18.

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PROCEEDINGS

OF THE

CALIFORNIA ACADEMY OF SCIENCES
FOURTH SERIES

Volume XXXIX, No. 2, pp. 7-24; 25 figs. January 21, 1972

THREE NEW LAND SNAILS FROM ISLA SANTA
CRUZ (INDEFATIGABLE ISLAND), GALAPAGOS

ey
Allyn G. Smith

Associate Curator, Department of Invertebrate Zoology, California Academy of Sciences

From late January to early March, 1964, I had the opportunity to collect land
snails on one of the larger of the Galapagos Islands as a participant in the Gala-
pagos International Scientific Project (GISP). This expedition was sponsored
by the University of California, the California Academy of Sciences through the
Belvedere Scientific Fund, and the Charles Darwin Foundation. Assistance, both
financial and material, was provided by the government of the Republic of
Ecuador, the United States Navy, the National Science Foundation of the United
States*, the Shell Oil Company, and the California Maritime Commission whose
training vessel, the Golden Bear, provided transportation of personnel and equip-
ment to and from the Galapagos.

The major portion of the five weeks in the Galapagos Islands was spent in
collecting on Isla Santa Cruz (Indefatigable). The decision to concentrate on
this island was made for several reasons. First, it is large, high, and well forested,
containing all of the life zones occurring in the Galapagos group; second, the
Darwin Research Laboratory close to the village of Academy Bay made an
ideal headquarters; and third, the well developed trail from Academy Bay into
the highland area, which was pioneered by the members of the California Acad-
emy’s 1932 Expedition sponsored by Templeton Crocker, cut through all life
zones and made good collecting spots accessible (see map, fig. 1). These reasons
fortunately tied in with the fact that the late winter and spring of 1964 proved
to be a good time to find land snails on Isla Santa Cruz. There was considerable

* Through NSF Grant no. GE-2370

[7]

Marine Biological Lebaratory
LIBRA?CYy
JAN 3 1 1972

Weods Hole, Mass.

8 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

DAPHNE MINOR © (Be ISLA SEYMOUR

SKETCH MAP BY: NORTH CHANNELN
David Q. Cavagnaro
ned DAPHNE MAJOR @
Sigvart Horneman
March, 1965
Based on U.S. Hydrographic Chart
Nos. 5921 and 5939.

Punto del Norte

Punta Corrion

BAHIA CONWAY

A & ISLA SANTA CRUZ (INDEFATIGABLE) °
ROCAS
PS GORDON
SS istas
Seoles|e Covagnaro Crater Qc es
CENTRAL HIGHLAND CHAIN ae

+

No

\ } Deep Crater
W237 i
ixe™, “* al - #2 - eae Crocker

* Scalesia
x Chimney.” 34 ** 4p Table Mountain
Sonta)Rosci@). grassland Minow gee bonds ote DCroter
: miconia grassland fi Camote
NAMELESS ee {Le Copa
© ISLAND aE PS ores: \miconla .~“
mora imoras. oo “
i “FARM ZONE =< @ Kastdalen Farmhouse
ponds H “ss... Castro °,@Horneman Farmhouse

El Chato Farmhouse Sow Bello Vista

“\.@ El Rancho Puerto Nunez

Transition c Transition
Bahia Tortuga | Old Trail
z New Trall “2*<.5 Coastal

°s. Barranco
ne Sei smograph
= Coastal Way oT 20g ote
4.3 mm. = 1 km. Punta re
lem. =23.14 km. Tomayo

APPROXIMATE SCALE

~ Punta Nunez

Darwin Research Station

= © Jensen Island
unto

Ploya Estrada
de los

Perros

"~ BAHIA ACADEMY
Bahia Tortuga

1

Ficure 1. Outline map of Isla Santa Cruz (Indefatigable Island), Galapagos Islands,
with some of the principal features sketched in.

rainfall, even in the Arid Zone along the shores of the island, which promoted
snail activity.

The collections made through personal efforts were considerably augmented
by other GISP scientists during their field work on other islands as well as Santa
Cruz. To these appreciation is due. In addition, special thanks go to André
and Jacqueline De Roy, a Belgian couple living at Academy Bay, for much
pertinent information on possible productive collecting areas. Although not an
active conchologist like his wife, M. De Roy became interested in the species of
land snails to be found on his island and has made a number of special trips into
uncollected areas in the highlands of Isla Santa Cruz in the past several years
with signal success.

The following new species discovered as a result of these efforts make a
significant addition to the land snail fauna of the Galapagos. The symbols
“CASG” and “CASIZ” represent separate collections maintained by the Califor-

VoL. XXXIX] SMITH: NEW GALAPAGOS LAND SNAILS 9

nia Academy of Sciences’ Geology and Invertebrate Zoology departments, respec-
tively, the latter consisting mainly of specimens preserved in alcohol.

Naesiotus deroyi A. G. Smith, new species.
(Figures 2-9.)

DescripTION. Shell fairly large for the genus, elongate-conic, yellowish
white at the apex graduating to pure white on the body whorl. Whorls about 7,
gently rounded, the sutures well impressed. Aperture subquadrate, with 2 promi-
nent denticles, the first a strong, rounded boss on the columella, and the second a
less strong, somewhat laterally compressed, subtriangular, parietal denticle posi-
tioned a short distance behind the plane of the aperture. Peristome thickened
slightly but not reflected except on the basal portion, which partly covers a small
shallow umbilicus. The peritreme is completed by a fairly heavy wash of callus.
The columellar axis is simple, solid, and only slightly twisted. (See fig. 4.)

Nuclear whorls 2’, erect, sculptured by many fine, closely spaced, occa-
sionally anastomosing, slightly sinuate and protractive transverse lirae. The
first postnuclear whorl is relatively smooth and marked by hardly noticeable
lines of growth. Beginning with the second postnuclear whorl, a rough irregular
but generally transverse wrinkling appears, becoming very heavy, somewhat
warty, and more or less patternless on the penultimate and especially on the
body whorl. Underlying this strong wrinkled sculpture are fine, closely spaced,
spiral lirae, also beginning on the second postnuclear whorl, but not overriding
the raised areas forming the wrinkles.

The animals, in alcohol, are yellowish white in color with no apparent special
markings on the mantles. Unfortunately, I did not see specimens alive.

Hototype. An adult shell (CASG Type Collection, no. 13730) with animal
in alcohol (CASIZ Type Series, no. 466), the shell measuring: height, 19.8;
maximum diameter, 12.0; height of aperture, 8.8; maximum width of aperture,
7.6 mm. Number of whorls, 6%. The apical angle of the spire is about 50°.
(Seedies: 2,3.)

PaRATYPES. Seventy-two specimens collected with the holotype. These have
been deposited in various museums having recognized type collections, including
the CASIZ Type Series, and in several private collections.

Type Locality. Isla Santa Cruz (Indefatigable Island), Galapagos, on the
northwest side at an elevation of about 264 m. (870 ft.), on a species of thorny
bush (locally called “mora,” the botanical name not available) by André De
Roy, 18 February 1964.

OTHER MATERIAL EXAMINED (ALL FROM ISLA SANTA CRUZ).

(1) Three adult specimens (CASG no. 40129) found in a locality “1-2 hours
walk west of the Horneman Farm,’’—at an elevation of about 200 m. (650 ft.).

10 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

4 2

Ficure 2. Naesiotus deroyi A. G. Smith, new species. Holotype. Height, 19.8 mm. Aper-
tural view. CASG Type Collection, no. 13730. Ficure 3. Same. Back view. FIGURE 4.
Same. Shell with cut-away section to show configuration of columella. Paratype. Height,
18.1 mm. CASG Type Collection, no. 13731. Ficure 5. Naesiotus cf. N. deroyi. Adult

VoL. XXXIX] SMITH: NEW GALAPAGOS LAND SNAILS 11

These were the first ones found by André and Jacqueline De Roy in 1963 in
the same general area as the type lot collected in 1964. Others are in the De Roy
Collection.

(2) Six adult shells, typical (CASG no. 40076), collected on the trail 6 miles
to the west of the village of Bella Vista, on bushes, by André De Roy, 10 June
1964.

(3) One typical but rather small subadult (CASG no. 40302), collected in an
area to the north of the central chain of craters well north and slightly west of
the village of Santa Rosa, on the ground in a dense forest of Scalesia pedunculata,
by André De Roy, 26 November 1966.

(4) One juvenile (CASG no. 40122), collected 5 miles northwest of Bella
Vista, on “mora,” at about 265 m. (870 ft.) elevation by André De Roy, 18
February 1964.

(5) Three rather small adults (CASG no. 40225), collected on the north side
of the hill nearest to Santa Rosa Spring, on grass in a grassy glade, 200 m. (650
ft.) elevation, by André De Roy, 29 June 1965.

(6) Fourteen adults and subadults (CASG no. 40022), found 2 km. north-
east of the village of Santa Rosa, on the ground, by André and Jacqueline De
Roy, 28 June 1965. This series is not typical.

(7) Seven specimens (mostly subadults) preserved in alcohol (CASIZ Coll.),
taken about 2 km. northeast of Santa Rosa, in open Scalesia forest hanging on
the leaves of various plants and bushes, by André De Roy, 27 November 1966.
This and the preceding lot are similar.

Remarks. This is one of the largest and most strikingly sculptured of the
Galapagos species of Naesiotus. No species closely similar has been described.
While an average specimen is about 20 mm. high, with 7 whorls or slightly less,
an unusually large one may be as much as 23 mm. in height, with the whorls
numbering 7% to 72. It appears to be a terminal species in the group of
Galapagos snails that have developed a heavily wrinkled sculpture on nearly all
of the postnuclear whorls rather than on the last portion of the body whorl, which

<

shell from lot no. 6 with weaker wrinkled sculpture. Hypotype. Height, 17.4 mm. CASG
Type Collection, no. 13733. Apertural view. FIGURE 6. Same. Back view. FIGURE 7.
Naesiotus deroyi. Adult shell from lot no. 5. Hypotype. Height, 17.5 mm. CASG Type
Collection, no. 13734. Apertural view. Ficure 8. Same. Back view. FicurE 9. Galapagos
Naesiotus related to N. deroyi. Left to right: N. rabzdensis (Dall, 1917) from Isla Rabida
(Jervis Island), hypotype, height, 13.5 mm. (CASG Type Collection, no. 13735); N.
sculpturatus (Pfeiffer, 1846) from Isla Santiago (James Island), hypotype, height, 13.6
mm. (CASG Type Collection, no. 13736); N. darwini (Pfeiffer, 1846) from the same
island, hypotype, height, 15.9 mm. (CASG Type Collection, no. 13737) ; N. deroyi, holotype,
fig. 2.

12 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

is the type of sculpture on such species as Naesiotus lycodus (Dall, 1917), N.
ochsneri (Dall, 1917), and several others. So far, NV. deroyi has been collected
in the highland area of Isla Santa Cruz, only toward the northwest side of the
island; it seems to prefer the moister forested area characterizing the Scalesia
Zone. The nearest relative is Naesiotus darwini (Pfeiffer, 1846) from James
Island, which is smaller, chunkier, and has a finer and much less rude type of
wrinkled sculpture. Other species in the group include \. sculpturatus (Pfeiffer,
1846) from James Island, and NV. rabidensis (Dall, 1917) from Jervis. (See fig.
9.)

Snails from lots 6 and 7 listed under “other material examined” come from
the same general locality as the others. However, they are smaller in size than
those from the type locality, averaging 17.1 mm. in height, with a range of 15.7—
17.6 mm. Color is a light yellow-brown and all shells have a narrow, darker
brown band encircling the periphery of the whorls; many have the entire base
tinged with the same brown color. The wrinkled sculpture is much less strong,
some shells having smooth patches with no wrinkles at all. (See figs. 5, 6.) As
in some other Galapagos snail species, this particular population evidently
represents an evolutionary trend of recent origin. Whether a subspecies is in
the process of development is difficult to say until the range limits of both forms
can be determined and other related factors studied. At present, it seems suffi-
cient to call attention to the occurrence of another race closely allied to V. deroyi.

It is with considerable satisfaction that I take this opportunity to name a
striking new species of Galapagos land snail for M. André De Roy, who collected
the first specimens as well as the type and other lots.

Naesiotus cavagnaroi A. G. Smith, new species.
(Figures 10-18.)

DescripTION. Shell fairly large for the genus, smooth, broadly elongate-
conic, with a fairly heavy texture and a tumid body whorl. Normal color pattern
is reddish brown to chocolate brown with a narrow yellowish band coloring the
sutures of the postnuclear whorls and encircling the body whorl slightly below
its periphery. Whorls about 6’, rounded, the sutures impressed. Aperture ovate,
white inside, with 2 well developed denticles, the first an elongate, rounded
swelling on the columella, the second one smaller, arcuate, and parietal, set well
inside the retractive plane of the aperture and forming a U-shaped bay with the
columellar denticle. Peritreme rather sharp, thinned down toward its edge, not
reflected, the outside edge yellowish in color. The peritreme is completed by a
heavy layer of callus, especially in older shells. There is a small shallow umbil-
icus partly covered by the basal reflection of the peristome. The columella is
simple and only slightly twisted.

Nuclear whorls about two, dimpled, appearing smooth to the naked eye but

VoL. XXXIX] SMITH: NEW GALAPAGOS LAND SNAILS 13

under magnification revealing a sculpture of extremely fine, closely spaced,
transverse lirations that are slightly sinuate and beaded at their summits as a
result of being cut by excessively fine spiral striae. Postnuclear whorls sculptured
by lines of growth and very fine, closely spaced, spiral striations, the latter best
seen under considerable magnification.

The surface of most shells exhibits a rather dull finish overall, with a tendency
in some toward a more shining exterior. Occasional shells are yellowish in color
with no suggestion of a revolving band. Animals in alcohol are tuberculate dor-
sally and light grey in color, there being no appreciable color difference between
those occupying normally colored or xanthic shells.

Hototype. An adult shell preserved without the animal has been deposited
in the CASG Type Collection, no. 13738. It measures: height, 22.7; maximum
diameter, 13.9; height of aperture, 10.4; maximum width of aperture, 9.1 mm.
Number of whorls, 6%. Apical angle, about 70°. (See figs. 10, 11.)

ParaTyPeEs. A total of 52 specimens collected with the holotype. Of this
total 30 have adult shells, 17 are subadults, and 6 are juveniles. Of this same
total, 10 have xanthic shells (6 adults, 3 subadults, and 1 juvenile). About half
of the type lot were collected alive; animals with their shells of a few of both
color forms have been preserved in alcohol. A distribution of these paratypes
will be made in a manner similar to that indicated for the previously described
species (Naesiotus deroyt).

Type Locatity. Isla Santa Cruz (Indefatigable Island), Galapagos, about
7 km. northeast of the farming village of Santa Rosa in the vicinity of a series
of small volcanic craters in a Scalesia forest; collected under lava rocks and
dead wood by André De Roy, 27 November 1966.

OTHER MATERIAL EXAMINED.

(1) Three dead, bleached, adult shells (CASG no. 40158) collected along the
trail near the summit of Mt. Crocker at an elevation of 650-870 m. (2130-
2850 ft.) by Robert L. Pyle, 25 February 1964.

(2) Seven dead, bleached, adult shells (CASG no. 40237) collected near an
isolated rocky crater (subsequently designated as Cavagnaro Crater), in a
Scalesia forest on the north slope of the island at an elevation of about 500 m.
(1600 ft.) by David Q. Cavagnaro, 10 April 1964.

(3) One dead, bleached adult shell (CASG no. 40512) found about 0.5 mile
below Santa Rosa on the trail from Bella Vista by André and Jacqueline De Roy,
1 March 1965.

(4) One dead but fresh adult shell, 1 subadult and 1 juvenile (CASG no.
40229) collected in the Scalesia forest near Cavagnaro Crater, elevation about
625 m. (2050 ft.), by André and Jacqueline De Roy, 2 March 1965.

(5) Sixty-two dead adult shells (CASG no. 40221) collected on the ground

14 CALIFORNIA ACADEMY OF SCIENCES [Proc, 4TH SER.

12 15 18

Ficure 10. Naesiotus cavagnaroi A. G. Smith, new species. Holotype. Height, 22.7 mm.
CASG Type Collection, no. 13738. Apertural view. Ficurre 11. Same. Back view. FIGURE

VoL. XXXIX] SMITH: NEW GALAPAGOS LAND SNAILS 15

in the vicinity of Cavagnaro Crater, elevation about 690 m. (2200 ft.), by André
and Jacqueline De Roy, 2 March 1965.

(6) One dead, bleached, adult shell (CASG no. 40228) collected on the
ground near Santa Rosa school by André De Roy, 2 November 1965.

(7) One dead, bleached, adult shell (CASG no. 40234) collected 2 miles
from the Santa Rosa school on the trail from Bella Vista, by André De Roy, 2
November 1965.

(8) Seven dead, bleached, adult shells (CASG no. 40295) collected in the
vicinity of Cavagnaro Crater, by André De Roy, 23 November 1966.

(9) Three bleached “bones” (CASG no. 40300) collected in a small lava
cave to the west of Chimney Mountain, by André De Roy, 25 November 1966.

(10) Thirty-eight adult shells, all xanthic (CASG no. 27537), collected “at
the very top of the island” (7.e., summit of Mt. Crocker) by Templeton Crocker,
10 or 11 May 1932.

(11) Forty-seven adult shells, all xanthic but darker in color than the pre-
ceding lot (CASG no. 27538), collected “‘on the trip to top of Mt.,” by Templeton
Crocker, 10 or 11 May 1932.

(12) Two adults and 1 juvenile with color pattern reversed taken alive half
way between Chimney Mountain and Santa Rosa Spring under low bushes in
an open area, elevation about 600 m. (1950 ft.), by André and Tui De Roy,
27 June 1965.

(13) Thirty-three mostly adult shells taken alive (CASG no. 43333; 15 in
alcohol are CASIZ Collection) 2 miles west of Mt. Crocker at the foot of a volcanic
crater on the ground among small trees, by André and Jacqueline De Roy, 10
May 1970.

Remarks. This smooth brown snail with a yellowish revolving band has no
close relatives of comparable size either on Isla Santa Cruz (Indefatigable
Island) or on any of the other Galapagos Islands. Its range on Indefatigable is
limited to the north slope of the island extending down from the summit of the

<_

12. Same. Shell with cut-away section to show configuration of columella. Paratype.
Height, 22.0 mm. CASG Type Collection, no. 13739. Ficure 13. Same. Xanthic color-
phase, adult, from type lot. Paratype. Height, 21.2 mm. CASG Type Collection, no. 13740.
Ficure 14, Same. Xanthic shell, adult, with over-all greenish-brown color tone, from lot
no. 11. Hypotype. Height, 23.1 mm. CASG Type Collection, no. 13741. Apertural view.
Ficure 15. Same. Back view. Ficure 16. Same. Adult shell with color pattern reversed,
from lot no. 12. Hypotype. Height, 18.8 mm. CASG Type Collection, no. 13742. Ficure 17.
Same, Adult shell with normal color pattern reversed, from lot no. 13. Hypotype. Height,
26.7 mm. CASG Type Collection, no. 13743. Ficure 18. Naestotus duncanus (Dall, 1893).
Adult dead shell from Isla Pinzén (Duncan Island). Hypotype. Height, 18.7 mm, CASG
Type Collection, no. 13744.

16 CALIFORNIA ACADEMY OF SCIENCES [PRroc. 4TH SER.

main volcanic crater, Mt. Crocker, generally in the Scalesia Zone, at elevations
above 500 m. (1600 ft.). It is a ground snail, frequenting moist habitats under
lava rocks, Scalesia dead-falls, or equivalent cover.

Naesiotus cavagnaroi is remarkably consistent in size and general configura-
tion. Measurements of 15 adult shells from the type locality average 21.9 mm.
in length and 13.4 mm. in maximum diameter, with about 6’ whorls. The cor-
responding range measurements for this series are 21.0—23.5 and 12.4—14.6, with
the number of whorls ranging from 5*4 to 6%, the latter number being the most
frequent. Young shells are prominently keeled at the base of the developing
whorls.

The occurrence of xanthic shells along with normally colored ones is of
particular interest. (See figs. 13-15.) No similar situation has been observed
in any other Galapagos species of Naesiotus. According to André De Roy, who
collected the type series, the ratio of xanthic to normally colored shells is about
one to five, which is confirmed by the count of specimens sent by him for identifi-
cation and study. There is no difference in the color of the animals and all
xanthic shells seen fall within the range of measurements given above. Lot nos.
10 and 11 in the preceding list are all the bandless xanthic color form. They
were collected by Templeton Crocker during the Academy’s 1932 Galapagos
Expedition at the time he and his party pioneered the first ascent of the island’s
main crater, later called Mt. Crocker in his honor. The series of shells he col-
lected at the crater rim are of the same yellowish color as xanthic shells from
the type locality of the species. The second lot, collected at a lower elevation on
the way to the top, are darker in color with a greenish brown cast. Both of these
lots evidently represent pure xanthic populations of the species; the exact
localities where they were found are presently unknown and a search for them
should be made. .

The three specimens in lot no. 12 in the list of specimens examined have a
color pattern that is the reverse of the normal one, being a light beige-brown
overall with a dark brown encircling band. The two adult shells have red-brown
nuclear whorls but in size and sculpture they are normal for the species. (See
fig. 16.) Until recently this reverse color pattern was thought to be of rare
occurrence, but an apparently pure colony of such shells was collected by the De
Roys in the highland area in May 1970 (lot no. 13, preceding). Of the 33 shells
sent in for study, 21 have the same reverse color pattern just mentioned except
that the encircling brown band is bordered below by a whitish band of somewhat
variable width (see fig. 17); 7 have weak or indistinct banding and thus ap-
proach the xanthic form; and 5 are similar to the xanthic shells occurring in the
type lot. Compared with the type series, shells in this lot average slightly higher,
several being a little longer spired with deeper sutures. The largest shell mea-
sures 26.7 X 14.5 mm. in height and maximum diameter. Mme. De Roy’s com-

VoL. XXXIX] SMITH: NEW GALAPAGOS LAND SNAILS 17

ments on this lot (personal communication, 1 August 1970) are of such interest
that they are repeated, as follows:

Surprisingly, we found them living in an area where André had collected before, only
that he searched a couple of hundred yards on either side. In fact, the area where
this colony of snails is living is restricted (?) to a brushy patch surrounded by grass-
covered hills and may be no more than 3 or 4 acres. Snails there are quite abundant
but this could change before long, as barbed-wire fence already divides the place in
two lots and the land will be converted into pastures.

Although the reverse color phase may possibly turn out to be a different species
when the relation between it and the color phase considered to be the normal one
is better known, present indications are that there is only a single species in-
volved, in spite of the fact that there are four color forms.

Two described Galapagos species have some superficial resemblance to
Naesiotus cavagnaroi. One is N. duncanus (Dall, 1893) from Duncan Island
(Isla Pinzon), shown in figure 18, which has the same general shape and number
of whorls although it is considerably smaller in size, has a less well developed
parietal denticle, and lacks the columellar thickening or flange present on J.
cavagnaroi. Unfortunately, NV. duncanus has not been reported as having been
collected alive and may, in fact, be extinct. The other similar species is the rare
N. jervisensis (Dall, 1917), which again is much smaller than NV. cavagnaroi,
has a less well developed parietal denticle and columellar flange, but has the
same type of fine spiral sculpture: it differs, however, in having the last whorl
irregularly corrugated and in being unicolored and unbanded.

The species is named for David Q. Cavagnaro, California Academy ento-
mologist and a member of the GISP, who collected some of the first specimens
and also a number of other land snails of considerable scientific value on Isla
Santa Cruz and on several other Galapagos islands he was able to visit.

Naesiotus scalesiana A. G. Smith, new species.
(Figures 19-25.)

DESCRIPTION. Shell of medium size for the genus, elongate to globose-conic,
the apex rose-colored with the remaining shell whitish or tinged with beige,
yellowish white, or very light red-brown. Whorls 5% to 6%, almost flat-sided,
the sutures only moderately impressed. Aperture subovate, usually without
denticles; the peristome thin and unflared, hardly reflected at the base of the
columella, leaving open a small but permeable umbilicus, not joined across the
parietal wall of the body whorl with any appreciably thickened wash of callus.

Nuclear whorls nearly 3, erect, sculptured with extremely fine, closely spaced,
nearly straight transverse riblets or lirae, extending all the way across them.
Postnuclear whorls somewhat shining, the first one or two with coarse, irregular
lines of growth, later ones with additional, irregulary placed, rather deep indenta-

18 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

24

Ficure 19. Naesiotus scalesiana A. G. Smith, new species. Holotype. Height, 14.1 mm.
CASG Type Collection, no. 13745. Apertural view. Ficure 20. Same. Back view. FIGURE
21. Same. Shell with cut-away section to show configuration of columella. Paratype. Height,
13.6 mm. CASG Type Collection, no. 13746. Ficure 22. Same. Enlarged view of portion

VoL. XXXIX] SMITH: NEW GALAPAGOS LAND SNAILS 19

tions and elongate impressions, which are most prominent on the body whorl.

Animals in alcohol generally yellowish white, occasionally tinged with light
gray on the dorsal surface, the foot usually being of a slightly lighter color tone.
Rarely, an animal is quite dark-colored. There appears to be no correlation
between the color of the animal and the color of the shell.

Hototype. An adult shell measuring: height, 14.1; maximum diameter,
7.5; height of aperture, 6.4; width of aperture, 3.7 mm. Number of whorls, 6%.
Apical angle, about 45°. CASG Type Collection, no. 13745.

PaRATYPES. Ninety-two additional specimens preserved dry, deposited in
various museums maintaining type collections including the CASG Type Collec-
tion, and 4 preserved in alcohol in the CASIZ Type Series, no. 470.

Type LocaLity. Isla Santa Cruz (Indefatigable Island), Galapagos Islands,
in the Scalesia Zone, Horneman Farm area, at an elevation of about 244 m. (800
ft.), on Scalesia trees and tall bushes, collected by A. G. Smith, 19 February
1964.

RANGE AND ECOLOGY. A total of 36 separate lots, in addition to the type lot,
have been collected and are available for study. All of these have contributed
to the present consideration of the species. They represent over 500 specimens,
mostly preserved dry, although a representative series of animals were drowned
and preserved in alcohol in an expanded condition.

Naestotus scalesiana seems to be the most prevalent species throughout the
Scalesia Zone on Isla Cruz. Its occurrence most often on the trunks and branches
of Scalesia trees makes the name given to it an appropriate one. This habitat is
shared with NV. lycodus (Dall, 1917), also pink-tipped but with a much more
wrinkled shell, which is almost as common at the type locality although it does
not climb as high up as NV. scalesiana. The latter species has been collected also,
though sparingly, at the upper edge of the Transition Zone and in the lower part
of the Miconia Zone but it does not seem to thrive outside the areas occupied
by the forests of Scalesia pedunculata wherever they occur under conditions of
heavier rainfall and hence of moister conditions than exist in the lower or the
higher, somewhat dryer life zones.

Along the “old” or original trail from Academy Bay village to the highland

<

of holotype to show sculptural detail. Ficure 23. Same. Group of five adult shells from
Scalesia Zone below Bella Vista village, Isla Santa Cruz. Allyn G. Smith and Ira L. Wiggins,
collectors, 19 February 1964. Hypotypes. Height (left-hand shell), 13.3 mm. CASG Type
Collection, nos. 13747, 13747a, 13747b, 13747c, 13747d. From CASIZ Color Slide, no. 955.
Ficure 24. Same. Group of five adult shells from type lot. Holotype at left (height, 14.1
mm.) ; the rest paratypes, CASG Type Collection, nos. 13748, 13748a, 13748b, 13748c. From
CASIZ Color Slide no. 1937.

20 CALIFORNIA ACADEMY OF SCIENCES [PRroc. 4TH SER.

area on Isla Santa Cruz, Dr. Ira L. Wiggins and I first encountered NV. scalesiana
in 1964 in the Transition Zone at an elevation of about 120 m. (400 ft.) (CASG
no. 40083, 2 adult specimens). It became quite common at the lower end of the
Scalesia Zone on young Scalesia trees, along with N. lycodus, and continued
prevalent in the vicinity of the village of Bella Vista, the areas of the Horneman
and Kastdalen Farms, and for a short distance into the Miconia Zone (CASG
no. 40070, 3 adult specimens). Elsewhere in the Scalesia Zone M. André De
Roy and others collected it toward the eastern side of the island in the Table
Mountain area (CASG no. 40232), on the northern side in the general vicinity
of Cavagnaro Crater (CASG no. 40221), around the village of Santa Rosa, and
to the south of Santa Rosa near the end of the trail into the Tortoise Reserve
at El Chato.

The Scalesia snail lives in more of an arboreal habitat than any other Gala-
pagos land snail with which I am familiar. On one collecting trip, Dr. Wiggins
and I decided to sacrifice one large Scalesia branch to see just how far up the
snail might go. We found two living adults (CASG no. 40231) at the very tip
of the branch, about 35 feet from the ground, among the small sunflower-like
Scalesia blossoms in flower at the time. It does not confine itself entirely to
Scalesia plants and trees, however, but has been collected living on Darwin-
iothamnus, a bush of medium height, and below Santa Rosa village in thickets
of thorny shrubs called “mora” by the native Ecuadoreans in company with
another new species of Galapagos land snail, Naesiotus deroyi, described earlier
in this report.

Unfortunately, nothing is known at present of the breeding habits of the
species; its egg clusters were not found in 1964 and have not been reported
subsequently.

INDIVIDUAL VARIATION. The type lot of VV. scalesiana was selected for general
consistency in size and color of the shells, and also for the accessibility of the
type locality at the Horneman Farm, making it relatively easy for topotypical
material to be collected. Color in the type lot ranges from almost pure white
to an overall suffusion of beige or light brown; only one adult shell in the series
has darker postnuclear whorls, with a body whorl having a distinctly brown
and white banded pattern. (See fig. 24.) The rose-colored tip is consistent
within the type lot and for most of the other shells at hand.

Variation in size within the type lot is shown in the following table of mea-
surements of 25 adult paratypes, selected more or less at random:

Item Height (mm.) Max. Diam. (mm.) Apical Angle (°) No. Whorls
Tallest shell i522 7.6 49 614
Shortest shell 115 6.2 51 5%

Avg. of 25 1333) Til 50 5%,

VoL. XXXIX] SMITH: NEW GALAPAGOS LAND SNAILS 21

Ficure 25. Naesiotus scalesiana. Shell with living animal. Collected by Robert L. Usinger
and Earl G. Linsley, Bella Vista village, 4 February 1964. Author’s field photo taken at
Academy Bay. From CASIZ Color Slide, no. 440.

Shells from other localities on the island have measurements that are consistent
with the preceding figures. A few adult shells from the vicinity of Santa Rosa
have slightly more tumid whorls and lack the usual rose-colored apex.

Most shells have simple apertures with no parietal or columellar denticles
or denticular swellings; this is true of all specimens in the type lot and of others
collected in the Horneman Farm area. A few shells from the general vicinity
of Santa Rosa do have small to subobsolete parietal denticles, and in some but
not all of these there are perceptible swellings on the columella where a columellar
tooth would normally occur, were one present. Such denticular processes are
not a character of this species although they are normal in the apertures of other
species of Naesiotus from Isla Santa Cruz and other Galapagos Islands. Whether
the rare occurrence of apertural denticulation represents an evolutionary trend
toward development of prominent teeth in the aperture, or away from this, is
a question that cannot be answered from the present sparse knowledge of the
biology of the species and its probable ancestral relationships.

A well rounded body whorl is characteristic of NV. scalesiana, although occa-

22 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH Ser.

sionally an adult shell shows a barely perceptible angulation at its base near
the periphery. As might be expected, this feature is seen more often on juvenile
and subadult shells.

The indented sculpture on the body whorl is peculiar; it cannot be said to
be malleate in terms of small, rounded hammer marks. Rather, it consists of
pits in combination with elongated, sometimes anastomosing furrows that have
the appearance of impressions made by a blunt point or an elongated, blunt edge.
This type of sculpture varies somewhat in intensity between populations from
which specimens have been collected. (See figs. 22, 25.) On the whole, however,
the shells of NV. scalesiana exhibit characters that are quite consistent throughout
its range.

CoMMENTS. It seems unusual that such an abundantly represented species
of Galapagos land snail has escaped notice for so long a time. W. H. Dall did
not recognize it as having been collected during earlier Galapagos expeditions;
apparently neither Baur nor Wolf saw it. Reibisch (1892) reported nothing like
it, his only large snail from Indefatigable Island (Isla Santa Cruz) being
Naesiotus wolfi, an entirely different species which he described as new. It was
not found by Snodgrass and Heller during the Hopkins-Stanford Expedition of
1897—98; nor was it collected by W. H. Ochsner during the California Academy’s
1905-06 Expedition, which spent quite a little time at Academy Bay. The
California Academy’s 1932 Expedition did not find it even though in the first
ascent to the rim of the main crater the party led by Templeton Crocker traversed
the Scalesia Zone on the way to the top as well as on the way back. The United
States National Museum has no specimens like it in its large representative col-
lection of Galapagos land snails; however, it does possess a single adult specimen
from an unidentified island, collected or obtained by Hugh Cuming, under the
name Bulimulus calvus (Sowerby, 1833) (USNM no. 104,864) that is quite
close to Naesiotus scalesiana. The first authentic collecting record known to
me is a single shell found by Dr. Robert I. Bowman, “5 miles North of Academy
Bay,” March 2, 1953, at an elevation of 775 feet (CASG no. 34649). All shells
collected subsequently were taken during the Galapagos International Scientific
Project of 1964 and since.

Naesiotus scalesiana has no demonstrable relationship with any of the other
15 species of Naestotus from Isla Santa Cruz, including the two previously
described as new in this report. I have not seen shells like it from any of the
other islands in the Galapagos Archipelago. Closest relatives would appear to
be certain species of Naesiotus from the mainland of South America (Weyrauch,
1956, 1967) but lack of appropriate mainland material at the present writing
places any detailed comparison beyond the scope of this discussion.

VoL. XXXIX] SMITH: NEW GALAPAGOS LAND SNAILS 23

ACKNOWLEDGMENTS

Appreciation and thanks to André and Jacqueline De Roy of Academy Bay,
Isla Santa Cruz, and to several scientists of the Galapagos International Scientific
Project of 1964 have already been expressed. In addition, David Q. Cavagnaro,
California Academy entomologist with the GISP, also should have special men-
tion for his land-snail collecting efforts on top of a full-time entomological pro-
gram.

I am greatly indebted to the California Academy’s scientific photographer,
Mr. Maurice Giles, for his excellent work in providing acceptable snail illustra-
tions from actual specimens, and for his black-and-white reproductions of three
of the author’s 35 mm. color transparencies.

LITERATURE CITED

DALL, WILLIAM HEALEY
1893. Preliminary notice of new species of land-snails from the Galapagos Islands,
collected by Dr. G. Baur. Nautilus, vol. 7, no. 1, pp. 52-56. May.
1896. Insular landshell faunas, especially as illustrated by the data obtained by Dr.
G. Baur in the Galapagos Islands. Proceedings of the Academy of Natural
Sciences of Philadelphia, 1896, pp. 395-459, pls. 15-17.
1900. Additions to the insular land-shell faunas of the Pacific Coast, especially of the
Galapagos and Cocos islands. Proceedings of the Academy of Natural Sciences
of Philadelphia, 1900, pp. 88-105, pl. 8.
1917. Preliminary descriptions of new species of Pulmonata of the Galapagos Islands.
Proceedings of the California Academy of Sciences, ser. 4, vol. 2, pt. 1, no.
11, pp. 375-382. San Francisco. December 31.
1920. On the relations of the sectional groups of Bulimulus of the subgenus Naesiotus
Albers. Journal of the Washington Academy of Sciences, vol. 10, no. 5, pp.
117-122. Washington, D. C. March 4.
DALL, WILLIAM HEALEY, AND WASHINGTON HENRY OCHSNER
1928. Landshells of the Galapagos Islands. Proceedings of the California Academy of
Sciences, ser. 4, vol. 17, no. 5, pp. 141-185, pls. 8-9. San Francisco. June 22.
Pirspry, Henry AUGUSTUS
1897-98. American Bulimulidae: Bulimulus, Neopetraeus, Oxychona, and South Amer-
ican Drymaeus. Manual of Conchology, ser. 2, Pulmonata, vol. 11, pp. 1-339,
pls. 1-51. Philadelphia.
REIBISCH, PAUL
1892. Die concholiologische Fauna der Galapagos Inseln. Sitzungsberichte und
Abhandlungen Naturwissenschaftlichen Gesellschaft Isis in Dresden, 1892
(January—June), pp. 13-32, 1 text fig. (map), pls. 1-2. Dresden.
STEARNS, ROBERT EDWARDS CARTER
1893. Report on the mollusk fauna of the Galapagos Islands with descriptions of new
species. Scientific Results of Explorations by the U. S. Fish Commission
Steamer Albatross, no. 30. Proceedings of the United States National Museum,
vol. 15, no. 942, pp. 353-450, pls. 50-52. Washington, D. C. August.
WEYRAUCH, WOLFGANG
1956. The genus Naesiotus, with descriptions of new species and notes on other Peruvian

CALIFORNIA ACADEMY OF SCIENCES [Proc, 4TH SER.

24

Bulimulidae. Proceedings of the Academy of Natural Sciences of Philadelphia,

vol. 108, pp. 1-17, pl. 1. June 22.

1967a. Treinta y ocho nuevos gastropodos terrestres de Peru. Acta Zoologica Lilloana,
tomo 21, pp. 343-455, pls. 1-9. Tucuman, Argentina.

1967b. Descripciones y notas sobre gastropodos terrestres de Venezuela, Columbia,
Ecuador, Brasil y Peru. Acta Zoologica Lilloana, tomo 21, pp. 457-499, pls.

1-4. Tucuman, Argentina.

PROCEEDINGS

OF THE

CALIFORNIA ACADEMY OF SCIENCES
FOURTH SERIES

Volume XXXIX, No. 3, pp. 25-46; 39 figs. April 6, 1972

PLIOCENE FOSSILS FROM BALTRA (SOUTH
SEYMOUR) ISLAND, GALAPAGOS ISLANDS

By
Leo G. Hertlein

California Academy of Sciences, San Francisco, California 94118

INTRODUCTION
The present paper deals chiefly with invertebrate fossils which I collected
while a member of the first Expedition of the Velero I// to the Galapagos Islands
in 1931-1932. A general account of the itinerary of this expedition appeared in
a report by Fraser (1943, pp. 50, 260, 262, 272-273). A report dealing with
fossils of Pleistocene age collected on this expedition was published by Hertlein
and Strong (1939) and in a later paper the same authors (1955) reported on the
Recent shells. Fossils believed to be of Pliocene age, chiefly mollusks, were
collected during January, 1932. The pressure of other duties delayed completion
of the present paper.
In addition to the collection which I assembled, Joseph R. Slevin collected
a few fossils of Pliocene age in 1927 when he visited the island on Captain G.
Allan Hancock’s Oaxaca. A few other specimens received through the courtesy
of Dr. A. Myra Keen of Stanford University, are included in the present paper.
The author is grateful for having had the opportunity to accompany the
expedition offered by the late G. Allan Hancock, owner of the Velero J/I. The
author acknowledges aid received from two other individuals, now deceased:
A. M. Strong, who identified the micro-gastropods and E. H. Quayle who con-
tributed information concerning the corals. Mr. Barry Roth, Department of
Geology, California Academy of Sciences, furnished information concerning
the identification of the species in the family Marginellidae, contributed helpful
comments concerning certain other species, and also aided in 1 the preparation
Marine Biological Labor..tory

LIBRAFRY

[25]

APR 14 1972
Woods Hole, Viass. :

26 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

the plates. The identifications of the corals listed in this paper were furnished
by Dr. J. W. Durham, Department of Paleontology, University of California,
Berkeley. The author is grateful to Carmen Angermeyer and to Jacqueline
De Roy, residents of Academy Bay, Santa Cruz Island, Galapagos Islands, who
presented many specimens of Recent shells from those islands to the California
Academy of Sciences. These specimens have been very useful for comparison
with fossil forms from that archipelago.

Photographs used to illustrate the fossils were prepared by Mr. Maurice
Giles, Photographer, California Academy of Sciences. Margaret M. Hanna
kindly retouched some of the photographs. The manuscript was typed by Enid
Cook.

One new subspecies, Diplodonta subquadrata baltrana, is described in the
present paper.

GENERAL REMARKS

Darwin visited the Galapagos Islands during the voyage of the Beagle in
1835. He mentioned (1844, p. 115; see ed. 3, 1896, p. 130) sea shells belonging
to modern genera, apparently on Chatham Island, ‘embedded several hundred
feet above the sea, in the tuff of two craters, distant from each other.” Later
Wolf (1895, pp. 250, 254, 265) reported the occurrence of fossil shells in the
islands in palagonite tuff at a height of 100 meters above sea level.

W. H. Ochsner, during an expedition of the California Academy of Sciences
to the Galapagos Islands on the schooner Academy, spent most of one year in
the islands and discovered fossils on Isabela (Albemarle), Santa Cruz (Inde-
fatigable), and Baltra (South Seymour) islands. On this expedition Ochsner
collected the fossils of Pliocene age on Baltra (South Seymour) Island mentioned
by Dall (1924) and later reported upon by Dall and Ochsner (1927, see espe-
cially pp. 94-99). They listed 28 species (and four others identified only as to
genus) from beds believed to be of Pliocene age, 24 species from the “upper zone”
and 4 in the “lower zone”. Chesterman (1963, p. 344) reported on rocks of
the Galapagos Islands and mentioned fossiliferous limestone and “fossiliferous
tuffaceous sandstone” on Baltra (South Seymour), collected by Ochsner. He
also described a specimen of andesite from the south end of the island.

Baltra Island is composed chiefly of lava and other pyroclastic material with
very minor amounts of intercalated limestone and fossiliferous tuffaceous beds.

This island experienced faulting in comparatively recent time, the lines of
fracture trending approximately east-northeast. This faulting resulted in alter-
nating raised and depressed blocks. A depressed block separates Baltra (South
Seymour) from Santa Cruz (Indefatigable) Island and a similar depressed block
separates Baltra from Seymour (North Seymour) Island. Lewis (1956, p 290)
remarked on the relatively recent faulting as did Williams (1966, pp. 55-70)

Vot. XXXIX] HERTLEIN: PLIOCENE FOSSILS FROM BALTRA ISLAND 27

who discussed the general geology of the Galapagos Islands. More recently,
McBirney and Williams (1969, pp. 17-20, fig. 7) discussed the geology and
petrology of Baltra, as well as that of other islands in the archipelago.

The fossil-bearing bed on the south end of the bay on the west side of the
island is about 3 meters (10 feet) thick and dips about 5° or 6° south. Large
angular blocks of lava in the lower portion of the bed offer evidence of deposition
very near the shore.

The majority of the species in the present list were taken from the tuffaceous
ashy bed. A number of small specimens were obtained by passing some of the
white ashy material through a sieve.

COLLECTING STATIONS

Locality 1305 (CAS), cliff on southwest side of Baltra (South Seymour)
Island, Galapagos Islands. Joseph R. Slevin collector, December 16, 1927.
Pliocene.

Locality 27249 (CAS), white and yellowish tuffaceous strata interbedded
with lava on the south side of the bay on the west side of Baltra (South Seymour)
Island, Galapagos Islands. Leo G. Hertlein collector, January 16-18, 1932.
Pliocene.

Locality 27251 (CAS), strata about 1.5 meters (5 feet) thick on the northern
side of the bay, about the middle of the west side of the island, near the top of
the plateau, Baltra (South Seymour) Island, Galapagos Islands. Leo G. Hertlein
collector, January 14-15, 1932. Pliocene.

Locality 31838 (CAS), Baltra (South Seymour) Island, Galapagos Islands.
Fossils received by Dr. A. Myra Keen who presented them to the California
Academy of Sciences, March 22, 1943.

LIST OF FOSSILS FROM BALTRA (SOUTH SEYMOUR) ISLAND*

An asterisk * indicates that the species is extinct.

COELENTERATA

Pavona gigantea Verrill
Psammocora (Stephanaria), species indeterminate

ECHINOIDEA
Encope micropora galapagana H. L. Clark, locs. 13051; 27249 (CAS)
Eucidaris thouarsii Valenciennes, loc. 1305 (CAS)

PELECYPODA
Anatina (Raéta) undulata Gould, loc. 31838 (CAS)
* Anodontia spherica Dall and Ochsner, locs. 1305; 27251 (CAS)
Anomia peruviana d’Orbigny, loc. 27249 (CAS)

1 Reported from this locality as Encope micropora L. Agassiz by Grant and Hertlein (Publ. Univ. Calif.
Los Angeles Math. Phys. Sci., vol. 2, p. 98, 1938).

28 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Arca pacifica Sowerby, loc. 1305 (CAS)
Arca (Arcopsis) solida Broderip and Sowerby, loc. 27249 (CAS)
Atrina cf. A. tuberculosa Sowerby, loc. 27249 (CAS)
Barbatia reeveana d’Orbigny, loc. 27249 (CAS)
Cardium elenense Sowerby, loc. 27249 (CAS)
Chama species, loc. 1305 (CAS)
Chione undatella Sowerby, loc. 27249; (cf.) 31838 (CAS)
Chione species, loc. 1305 (CAS)
Chlamys (Argopecten) circularis Sowerby, loc. 27249 (CAS)
Chlamys (Nodipecten) magnifica Sowerby, loc. 27249 (CAS)
Ctena galapagana Dall, locs. 1305; 27249 (CAS)
Cuminga ctf. C. lamellosa Sowerby, loc. 27249 (CAS)
Diplodonta subquadrata baltrana Hertlein, new subspecies, loc. 27249 (CAS)
Florimetis cognata Pilsbry and Vanatta, loc. 27249 (CAS)
Glycymeris maculata Broderip, loc. 27249 (CAS)
Megapitaria cf. M. aurantiaca Sowerby, loc. 27249 (CAS)
Megapitaria squalida Sowerby, loc. 31838 (CAS)
Modiolus capax Conrad, loc. 27249 (CAS)
Nioche cf. N. zorritensis Olsson, loc. 27249 (CAS)
Ostrea megodon Hanley, loc. 27249 (CAS)
Ostrea palmula Carpenter, loc. 1305 (CAS)
Ostrea species, loc. 27249 (CAS)

* Pecten (Pecten) slevini Dall and Ochsner, loc. 27249 (CAS)
Protothaca (Tropithaca) grata Say, locs. 27249; 31838 (CAS)

* Protothaca (Tropithaca) cf. P. (T.) seymourensis Dall and Ochsner, loc. 27249 (CAS)
Tagelus cf. T. dombeii Lamarck, loc. 27251 (CAS) (cast)
Tagelus species, locs. 27249; 31838 (CAS)

GASTROPODA

Acanthina grandis Gray, loc. 1305 (CAS)

Acteocina infrequens C. B. Adams, loc. 27249 (CAS)

Alaba supralirata Carpenter, loc. 27249 (CAS)

Alvania cf. A. galapagensis Bartsch, loc. 27249(CAS)
Alvania ct. A. halia Bartsch, loc. 27249 (CAS)

Alvania cf. A. hoodensis Bartsch, loc. 27249 (CAS)
Alvania cf. A. nemo Bartsch, loc. 27249 (CAS)
Amphithalmus trosti Strong and Hertlein, loc. 27249 (CAS)
Anachis cf. A. tabogaensis Bartsch, loc. 27249 (CAS)

Balcis cf. B. berryi Bartsch, loc. 27249 (CAS)

Bulla punctulata A. Adams in Sowerby, locs. 27249; 31838 (CAS)
Cancellaria cf. C. ovata Sowerby, loc. 27249 (CAS)
Cantharus janellii Kiener, locs. 1305; 27249 (CAS)

* Cantharus cf. C. scissus Olsson, loc. 27249 (CAS)
Cerithiopsis curtata Bartsch, loc. 27249 (CAS)
Cerithiopsis cf. C. galapagensis Bartsch, loc. 27249 (CAS)
Cerithium adustum Kiener, locs. 1305; 27249 (CAS)
Cheila equestris Linnaeus, loc. 27249 (CAS)

Conus fergusoni Sowerby, loc. 27249 (CAS)
Conus cf. C. lucidus Wood, loc. 27249 (CAS)

VoL. XXXIX] HERTLEIN: PLIOCENE FOSSILS FROM BALTRA ISLAND 29

Cylichna ct. C. defuncta Baker and Hanna, loc. 27249 (CAS)
Cymatium lineatum Broderip, loc. 1305 (CAS)

Cypraea nigropunctata Gray, locs. 1305; 27249 (CAS)
Cytharella camarina Dall, loc. 27249 (CAS)

Diodora alta C. B. Adams, loc. 27249 (CAS)

Engina pyrostoma Sowerby, loc. 27249 (CAS)

Erato (Hespererato) marginata galapagensis Schilder, loc. 27249 (CAS)
Eulimostraca cf. E. galapagensis Bartsch, loc. 27249 (CAS)
Fissurella cf. F. macrotrema Sowerby, loc. 27249 (CAS)
Fissurella virescens Sowerby, loc. 27249 (CAS)

Fusinus dupetitthouarsii Valenciennes, loc. 27249 (CAS)
Gastrocopta munita Reibisch, loc. 27249 (CAS) [a land snail]
Granula cf. G. minor C. B. Adams, loc. 27249 (CAS)
Granula cf. G. polita Carpenter, loc. 27249 (CAS)

Granula species, loc. 27249 (CAS)

Granulina margaritula Carpenter, loc. 27249 (CAS)

Hipponix pilosus Deshayes, locs. 1305; 27249 (CAS)
Hipponix pilosus Deshayes, loc. 1305; 27249 (CAS)

Tselica cf. I. kochi Strong and Hertlein, loc. 27249 (CAS)
Malea ringens Swainson, loc. 1305 (CAS)

* Mangelia cf. M. hancocki Hertlein and Strong, loc. 27249 (CAS)

Mitra gausapata Reeve, loc. 1305 (CAS)

Mitra lens Mawe, loc. 1305 (CAS)

Modulus cerodes A. Adams, loc. 27249 (CAS)
Nassarius nodicinctus A. Adams, loc. 27249 (CAS)

* Nerita oligopleura Dall and Ochsner, loc. 27249 (CAS)
Odostomia (Menestho) aequisculpta Carpenter, loc. 27249 (CAS)
Odostomia (Miralda) galapagensis Dall and Bartsch, loc. 27249 (CAS)
Odostomia (Chrysallida) rinella Dall and Bartsch, loc. 27249 (CAS)
Oliva species, locs. 27249; 31838 (CAS)

Pedipes angulatus C. B. Adams, loc. 27249 (CAS)
Persicula imbricata Hinds, loc. 27249 (CAS)
Persicula cf. P. phrygia Dall, loc. 27249 (CAS)
Persicula species, loc. 27249 (CAS)

Polinices dubius Récluz, loc. 27249 (CAS)

Polinices uber Valenciennes, loc. 27249 (CAS)
Pyramidella (Triptychus) ct. P. (T.) olssoni Bartsch, loc. 27249 (CAS)
Pyramidella (Voluspa) species, loc. 27249 (CAS)
Pyrene castanea Sowerby, locs. 1305; 31838 (CAS)
Pyrene fuscata Sowerby, loc. 27249 (CAS)

Pyrene haemastoma Sowerby, loc. 27249 (CAS)
Rissoina cf. R. firmata C. B. Adams, loc. 27249 (CAS)
Rissoina signae Bartsch, loc. 27249 (CAS)

Strombina gibberula Sowerby, loc. 27249 (CAS)
Tectarius galapagensis Stearns, loc. 27249 (CAS)

* Tegula forbesi Dall and Ochsner, loc. 27249 (CAS)

Tegula snodgrassi Pilsbry and Vanatta, locs. 1305; 27249 (CAS)
Triphora cf. T. galapagensis Bartsch, loc. 27249 (CAS)
Triphora cf. T. panamensis Bartsch, loc. 27249 (CAS)

30 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Trivia pacifica Gray, loc. 27249 (CAS)
Trivia radians Lamarck, loc. 27249 (CAS)
Turbo agonistes Dall and Ochsner, loc. 27249 (CAS)

* Turbo vermiculosus Dall and Ochsner, locs. 27249; 31838 (CAS)
Turbonilla (Chemnitzia) houseri Dall and Bartsch, loc. 27249 (CAS)
Turritella broderipiana marmorata Kiener, loc. 27249 (CAS)
Vermicularia eburnea Reeve, loc. 27249 (CAS)

Volvarina taeniolata Morch, loc. 27249 (CAS)

REMARKS ON THE AGE AND RELATIONSHIPS OF THE FAUNA

This list contains 102 identified species, 1 coral, 2 echinoids, 26 pelecypods,
and 73 gastropods. Of these, 7, and probably 8, are extinct. Twenty-eight
species are only provisionally identified but are compared with known species.
In addition to the 102 species, 8 forms are identified only as to genus.

The percentage of extinct species in the present faunal assemblage is approxi-
mately 7.8 percent. However, a greater number of extinct species evidently occur
at the present locality as Dall and Ochsner described additional extinct species
from apparently the same locality. They also described a number of extinct
species from what they believed to be approximately correlative strata on Santa
Cruz (Indefatigable) Island. Dall and Ochsner considered the age of the fauna
which they reported from those two islands to be of probable Pliocene age. Five
of the eight extinct species in the present faunal list have not been reported from
fossil assemblages of Pleistocene age from either Isabela (Albemarle) or from
San Salvador (James) Island. Furthermore, the occurrence of the fossils in ashy
beds (which in some places are decidedly indurated), interbedded with volcanic
flows, lends support to the viewpoint that these beds are of Pliocene rather than
of Pleistocene age. McBirney and Williams (1969, p. 19) mentioned two lavas
on Baltra Island reported to be 1.47 million years old.

Another factor contributing to the belief that the present fauna is of Plio-
cene age is the fact that Anodontia spherica Dall and Ochsner (reported from
strata of Pliocene age in the Galapagos Islands) was reported by Pilsbry and
Olsson (1941, p. 57) to occur abundantly and to be a characteristic species of
their “Zone H” in strata referred to Pliocene age at Punta Blanca, Ecuador.
Ten species in the present faunal list were included among those reported from
that locality.

Most of the Recent species in this assemblage are known living in Galapagos
waters. Those not known from there are chiefly elements of the Panamic fauna,
but further collecting may reveal their presence in the Galapagos Archipelago.
A consideration of the habitat of the Recent species in this fauna leads one to
infer that the fossil forms lived under conditions similar to those now prevailing
in the Galapagos Islands, namely, warm, shallow water.

Vot. XXXIX] HERTLEIN: PLIOCENE FOSSILS FROM BALTRA ISLAND 31

An interesting occurrence among the fossil forms is that of a land snail,
Gastrocopta munita Reibisch. This species now lives on Baltra (South Seymour)
Island (Hertlein, 1932b, p. 69) as well as on most of the other larger islands in
the Archipelago.

SYSTEMATIC PALEONTOLOGY
PELECYPODA

Chlamys (Nodipecten) magnifica Sowerby.
(Figures 5, 15, 25.)

Pecten magnificus SOWERBY, variety a, Proc. Zool. Soc. London for 1835, p. 109, issued
October 9, 1835. “Hab ad Insulas Gallapagos.” “A single specimen of var. a was found
in a coral sand at a depth of six fathoms.” SowerBy, Thes. Conch., vol. 1, p. 65, pl. 15
(Pecten, pl. 5), fig. 114, 1842. “East Columbia.” [lapsus calami for West “Columbia.”
Locality doubtful.] Rrrve, Conch. Icon., vol. 8, Pecten, species 9, pl. 2, fig. 9, 1852.
“Tsle of Plata, West Columbia (in coral sand at a depth of from six to seventeen
fathoms); Cuming.” [Locality doubtful.] Kopert, Syst. Conch—Cab. von Martini
und Chemnitz, Spondylus und Pecten, Bd. 7, Abt. 2, p. 164, pl. 46, fig. 1, 1888. (Descrip-
tion and illustration from Reeve.)

Pecten (Lyropecten) magnificus Sowerby, GRANT and GALE, Mem. San Diego Soc. Nat. Hist.,
vol. 1, p. 182, pl. 9, fig. 1; pl. 10, fig. 6, 1931. Galapagos Islands.

Nodipecten magnificus Sowerby, Grau, Allan Hancock Pac. Exped., vol. 23, p. 132, pl. 44,
1959. “Galapagos Islands. (Ecuador doubtful.)”

Lyropecten (Nodipecten) magnificus Sowerby, Otsson, Mollusks of the Tropical Eastern
Pacific (Paleo. Res. Inst.: Ithaca, New York), p. 161, pl. 22, fig. 1, 1961. (Illustration
from Reeve, 1852).

Two left valves, one nearly complete, the other only one half of a valve were
collected at locality 27249 (CAS). The larger specimen is 126 mm. long and
122.3 mm. high. There are 13 or 14 radial ribs. The entire valve is covered
with radial striae of which about 5 or 6 occur in each interspace. The ribs are
decidedly nodose. The nodes are hollow and develop in the areas of “ledges”
(Moore, 1934, p. 216) between concentric constrictions of the valve. The
nodosity on the east American species, Chlamys (Nodipecten) nodosa Linnaeus,
increases from north to south in its range into warmer waters.

A small left valve of C. magnifica 49 mm. high, was reported by Hertlein
and Strong (1939, p. 369) from a raised beach on San Salvador (James) Island.

Two Recent right valves in the collections of the California Academy of
Sciences, the larger one 127 mm. high, were collected by Ochsner on the beach
at Baltra (South Seymour) Island. These are devoid of nodes. The color is
bright red.

One right valve 59.8 mm. long and 60 mm. high, in the Academy’s collection,
received from Jacqueline De Roy, was dredged in 9 to 18 meters (5 to 10 fathoms)
off Santa Fé (Barrington) Island.

32 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Ficure 1. Anodontia spherica Dall and Ochsner. Hypotype, left valve, no. 13653 (Calif.
Acad. Sci. Dept. Geol. Type Coll.), from locality 802 (CAS), about 1% miles northeast of
the settlement of Vilamil, Isabela (Albemarle) Island, Galapagos Islands, elevated beach
deposit, about 12 meters (40 feet) above sea level. Pleistocene. Length 65 mm. Ficure 2.
Anodontia spherica Dall and Ochsner. Hypotype, right valve, no. 13654 (Calif Acad. Sci.
Dept. Geol. Type Coll.), from locality 803 (CAS), from virtually the same locality as the

Vor. XXXIX] HERTLEIN: PLIOCENE FOSSILS FROM BALTRA ISLAND 33

Anodontia spherica Dall and Ochsner.
(Figures 1, 2, 6, 7.)

Lucina spherica DALL AND OcusNER, Proc. Calif. Acad. Sci., Fourth Ser., vol. 17, no. 4, p.
121, pl. 3, fig. 8; pl. 4, figs. 2, 7, June 22, 1928; “from upper horizon (zone D) on east
shore of Indefatigable Island, Galapages Greuv. Probably Pliocene.”

Loripinus (Pegophysema) spherica Dall and Ochsner, Pitspry AND Otsson, Proc. Acad.
Nat. Sci. Philadelphia, vol. 93, p. 57, 1941. Pliocene of Punta Blanca, Ecuador.

Anodontia (Lissosphaira) spherica Dall and Ochsner, Orsson, Mollusks of the tropical
eastern Pacific (Paleo. Research Inst.: Ithaca, New York), p. 222, pl. 30, fig. 2, 1961.
(Figured specimen from Cabo Blanca, Ecuador, Pliocene. Reported as Recent from
“Lower California to Ecuador. Columbia: Isla del Gallo.”) Also earlier records.
Specimens assigned to this species in the present collection [locs. 1305 and

27251 (CAS) | are casts. Olsson reported this species living from Baja California

to Ecuador but it has not been reported from the Panamic Province by other

authors. I have not seen Recent specimens.

The deeply depressed posterior dorsal area on Anodontia spherica easily
serves to separate it from A. edentuloides from the Gulf of California. The
presence of well differentiated, deeply impressed, dorsal areas was the basis for
the proposal of the subgenus Lissophazra Olsson.

Large specimens of A. spherica are 74 mm. long. Berry (1968, p. 71) men-
tioned specimens of 4. edentuloides which are 75 mm. long.

Marks (1951, pp. 67-70), discussed the shell characters of Anodontia and
Pegophysema, and some of the species referred to those groups. Apparently, the
species cited by Marks (p. 69) as “C. densata” Dall and Ochsner is referable
to A. spherica.

A large globose species, 82 mm. long, Anodota sphericula Basedow, was
reported by Ludbrook (1959, p. 227, pl. 3, figs. 1, 2, 3; pl. 5, figs. 1, 4) from
strata of Pliocene age in Australia. The posterior dorsal portion of the shell
of that species lacks a depressed area: it is a typical Anodontia believed to be
related to A. philippiana Reeve in the western Pacific.

Diplodonta subquadrata baltrana Hertlein, new subspecies.
(Figures 8 and 11.)

DescripTIon. Shell subquadrate, somewhat anteriorly attenuated, thin,
beaks very low, anterior dorsal margin only slightly sloping; sculptured only

co

specimen shown in figure 1. Length 58 mm. Ficure 3. Trivia radians Lamarck. Hypotype
no. 13668 (Calif. Acad. Sci. Dept. Geol. Type Coll.), from locality 27249 (CAS), Baltra
(South Seymour) Island, Galapagos Islands. Late Pliocene. Length 18 mm. FicureE 4.
Trivia radians Lamarck. Ventral view of the specimen shown in figure 3. FicurE 5. Chlamys
(Nodipecten) magnifica Sowerby. Hypotype, right valve, no. 13652 (Calif. Acad. Sci. Dept.
Geol. Type Coll. ), from locality 23167 (CAS), Baltra (South Seymour) Island, Galapagos
Islands. Recent. W. H. Ochsner Collector. Height 85 mm.

34 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SEr.

13

Ficure 6. Anodontia spherica Dall and Ochsner. Hypotype, right valve, no 13655 (Calif.
Acad. Sci. Dept. Geol. Type Coll.), from locality 803 (CAS), about 1% miles northeast of
the settlement of Vilamil, Isabela (Albemarle) Island, Galapagos Islands, elevated beach
deposit, about 12 meters (40 feet) above sea level. Pleistocene. Length 54.5 mm. FIGURE 7.
Anodontia spherica Dall and Ochsner. View of the interior of the specimen shown in figure

Vor. XXXIX] HERTLEIN: PLIOCENE FOSSILS FROM BALTRA ISLAND 35

with fine concentric lines of growth. Dimensions: length 22.6 mm., height 20
mm., convexity (both valves) 14.6 mm.

Holotype, no. 13656, and paratype, a left valve, no. 13657 (California Acad-
emy of Sciences Department of Geology Type Collection), from locality 27249
(CAS), white and yellowish tuffaceous strata interbedded with lava, on the
south side of the bay on the west side of Baltra (South Seymour) Island, Gala-
pagos Islands. Late Pliocene. L. G. Hertlein, collector.

CoMMENTS. Specimens of this new subspecies, in general features, resemble
fossil forms of Diplodonta subquadrata Carpenter (see illustrations by Durham,
1950, pl. 19, figures 4, 4a, 1950) from Santa Inez Bay, Baja California, Mexico.
The present fossils from Baltra Island differ from Carpenter’s species in the
narrower, somewhat attenuated anterior end and in the more nearly horizontal
anterior dorsal margin. The general shape of the type specimen of the new
subspecies resembles that of the Recent specimen of D. subquadrata from the
Galapagos Islands illustrated by Hertlein and Strong (1947, plate 1, figure 11),
more than it does that of the fossils illustrated by Durham.

The paratype, 21.2 mm. long, is less attenuated anteriorly than the holotype.
An imperfectly preserved right valve also was collected at the type locality.

Protothaca (Tropithaca) cf. P. (T.) seymourensis Dall and Ochsner.
(Figures 38, 39.)

Twenty-one single valves of a venerid varying from 11 mm. to 39.4 mm.
in length are present in the collection from locality 27249 (CAS), Baltra (South
Seymour) Island. These vary greatly in shape and sculpture.

<

6. Ficure 8. Diplodonta subquadrata baltrana Hertlein, new subspecies. Holotype no.
13656 (Calif. Acad. Sci. Dept. Geol. Type Coll.), from locality 27249 (CAS), Baltra (South
Seymour) Island, Galapagos Islands. Late Pliocene. Length 22.5 mm. Ficure 9. Nioche cf.
N. zorritensis Olsson. Hypotype, left valve, no. 13658 (Calif. Acad. Sci. Dept. Geol. Type
Coll.), from locality 27249 (CAS), Baltra (South Seymour) Island, Galapagos Islands. Late
Pliocene. Length 39.5 mm. Ficure 10. Nioche cf. N. zorritensis Olsson. View of the interior
of the specimen shown in figure 9. Ficure 11. Diplodonta subquadrata baltrana Hertlein,
new subspecies. Dorsal view of the specimen shown in figure 8. Ficure 12. Nerita
oligopleura Dall and Ochsner. Hypotype no. 13666 (Calif. Acad. Sci. Dept. Geol. Type Coll.),
from locality 27249 (CAS), Baltra (South Seymour) Island, Galapagos Islands. Late
Pliocene. Height 14.5 mm. View showing fine threadlets between major concentric cords.
Ficure 13. Nerita oligopleura Dall and Ochsner. Hypotype no. 13667 (Calif. Acad. Sci. Dept.
Geol. Type Coll.), from the same locality as the specimen illustrated in figure 12. Apertural
view. Height 17.5 mm. Ficure 14. Nerita oligopleura Dall and Ochsner. Dorsal view of the
specimen shown in figure 13. Ficure 15. Chlamys (Nodipecten) magnifica Sowerby.
Hypotype, left valve, no. 13676 (Calif. Acad. Sci. Dept. Geol. Type Coll.), from locality
27249 (CAS), Baltra (South Seymour) Island, Galapagos Islands. Late Pliocene. Height
128 mm.

36 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Ficure 16. Turbo agonistes Dall and Ochsner. Hypotype no. 13671 (Calif. Acad. Sci.
Dept. Geol. Type Coll.), from locality 38977 (CAS), Baranco, Punta Nunez, Santa Cruz
(Indefatigable) Island, Galapagos Islands. Recent. Jacqueline DeRoy collector. Height
21 mm. Ficure 17. Turbo agonistes Dall and Ochsner. Dorsal view of the specimens shown
in figure 16. Figure 18. Turbo agonistes Dall and Ochsner. Hypotype no. 13672 (Calif.

Vor. XXXIX] HERTLEIN: PLIOCENE FOSSILS FROM BALTRA ISLAND 37

One small specimen in the present series is quite similar in outline and sculp-
ture to the early stage of growth of the type specimen of Chione seymourensts
Dall and Ochsner from the same area.

The type specimen of Chione seymourensis Dall and Ochsner (1928, p. 123,
pl. 3, figs. 1 and 5) was described “from upper horizon, Seymour Island, Gala-
pagos Group. Probably Pliocene.” It is a right valve, ovately rectangular in
outline (see figs. 36, 37), lacks an escutcheon, and the radial sculpture is some-
what reduced in strength as a result of erosion. Dall and Ochsner mentioned a
similarity between their species and Chione pertincta Dall, a Recent species
living in the Galapagos Islands. Compared with C. pertincta, the type specimen
of C. seymourensis is in general, more elongated, thinner, and it has finer radial
sculpture.

Compared with Protothaca grata Say, the type specimen of C. seymourensis
is more elongate in outline, the radial sculpture is coarser, the lunule is narrower,
and the median cardinal tooth in the right valve is larger.

The variation in shape and sculpture in the present series of specimens is
similar to that in a series of Recent specimens of Protothaca grata Say (the type
species of the subgenus Tropithaca Olsson), and most of them are here referred
to Say’s species.

GASTROPODA

Tegula forbesi Dall and Ochsner.
(Figures 24, 29.)
Tegula forbesi DALL AND OcHSNER, Proc. Calif. Acad. Sci., Fourth Ser., vol. 17, no. 4, p.

116, pl. 2, fig. 13, June 22, 1928; “from upper horizon on Seymour Island, Galapagos
Group. Probably Pliocene.”

<

Acad. Sci. Dept. Geol. Type Coll.), from locality 27249 (CAS), Baltra (South Seymour)
Island, Galapagos Islands. Late Pliocene. Height 28.5 mm. Ficure 19. Turbo agonistes
Dall and Ochsner. Dorsal view of the specimen shown in figure 18. Ficure 20. Turbo
agonistes Dall and Ochsner. View of the exterior of the operculum of the specimen shown
in figures 16 and 17. Ficure 21. Turbo agonistes Dall and Ochsner. View of the interior
side of the operculum shown in figure 20. Ficure 22. Turbo agonistes Dall and Ochsner.
Hypotype no. 13673 (Calif. Acad. Sci. Dept. Geol. Type Coll.), from the same locality as
the specimen shown in Figures 18 and 19. Height 33 mm. Dorsal view of an unusually
large specimen. Ficure 23. Turbo agonistes Dall and Ochsner. Apertural view of the speci-
men shown in figure 22. Ficure 24. Tegula forbesi Dall and Ochsner. Hypotype no. 13660
(Calif. Acad. Sci. Dept. Geol. Type Coll.), from locality 27249 (CAS), Baltra (South
Seymour) Galapagos Islands. Late Pliocene. Height 26 mm. FuicurE 25. Chlamys
(Nodipecten) magnifica Sowerby. Hypotype no. 13652 (Calif. Acad. Sci. Dept. Geol. Type
Coll.), from locality 23167 (CAS), Baltra (South Seymour) Island, Galapagos Islands. W.
H. Ochsner Collector. Recent. Height 85 mm. View of the interior of the specimen shown
in figure 5.

38 CALIFORNIA ACADEMY OF SCIENCES [Proc. 47TH SER.

Ficure 26. Turritella broderipiana marmorata Kiener. Hypotype no. 13669 (Calif. Acad.
Sci. Dept. Geol. Type Coll.), from locality 27249 (CAS), Baltra (South Seymour) Island,
Galapagos Islands. Late Pliocene. Height (incomplete) 53 mm. Ficure 27. Turritella
broderipiana marmorata Kiener. Dorsal view of the specimen shown in figure 26. FIGURE 28.
Turritella broderipiana marmorata Kiener. Hypotype no. 13670 (Calif. Acad. Sci. Dept.

Vor. XXXIX] HERTLEIN: PLIOCENE FOSSILS FROM BALTRA ISLAND 39

A few specimens of this species were collected at locality 27249 (CAS). The
largest specimen is about 31 mm. in height, the lest whorl is 48 mm. in maximum
width. The concentric sculpture is much finer than that of Tegula aureotincta
Forbes from California. This sculpture is more pronounced and regular on the
upper surface of the whorls, and on the base much coarser, than that on specimens
of T. rugosa A. Adams from the Gulf of California.

Turbo agonistes Dall and Ochsner.

(Figures 16, 17, 18, 19, 20, 21, 22, 23.)

Turbo agonistes DALL AND OCHSNER, Proc. Calif. Acad. Sci., Fourth Ser., vol. 17, no. 4, p.
115, pl. 2, figs. 12, 16, June 22, 1928; “on east shore of Indefatigable Island, Galapagos
Group. Probably Pliocene.”

A number of fossil specimens of this species were collected at locality 27249

(CAS). The shell of this species is quite different from any other described form.
Recently this species was found living in Galapagos waters. Four specimens

from off Santa Cruz (Indefatigable) Island collected by Jacqueline De Roy; and

one taken by Carmen Angermeyer 1’ miles west of Baltra (South Seymour)

Island, were presented to the Academy. Recent shells are attractively colored,

purplish on the base of the body whorl and orange or greenish above.

<

Geol. Type Coll.), from the same locality as the specimen shown in figures 26 and 27.
Height (incomplete) 40 mm. Ficure 29. Tegula forbesi Dall and Ochsner. Hypotype no.
13660 (Calif. Acad. Sci. Dept. Geol. Type Coll.), from locality 27249 (CAS), Baltra (South
Seymour) Island, Galapagos Islands. Late Pliocene. Width 24 mm. Basal view of the
specimen shown in figure 24. Ficure 30. Polinices dubius Récluz. Hypotype no. 13662
(Calif. Acad. Sci. Dept. Geol. Type Coll.), from locality 27249 (CAS), Baltra (South
Seymour) Island, Galapagos Islands. Late Pliocene. Height 37.5 mm. Ficure 31. Polinices
dubius Récluz. Apertural view of the specimen shown in figure 30. Ficure 32. Polinices
dubius Récluz. Hypotype no. 13663 (Calif. Acad. Sci. Dept. Geol. Type Coll.), from locality
804 (CAS), Baltra (South Seymour) Island, Galapagos Islands. Late Pliocene. Height 40.5
mm. Ficure 33. Polinices dubius Récluz. Hypotype no. 13664 (Calif. Acad. Sci. Dept. Geol.
Type Coll.), from the same locality as the specimen shown in figure 32. Height 39.4 mm.
Apertural view. Ficure 34. Polinices dubius Récluz. Apertural view of the specimen shown
in figure 33, showing notch in callus of inner lip. Frcure 35. Turbo vermiculosus Dall and
Ochsner. Hypotype no. 13661 (Calif. Acad. Sci. Dept. Geol. Type Coll.), from locality
27249 (CAS), Baltra (South Seymour) Island, Galapagos Islands. Late Pliocene. Height
34.6 mm. Ficure 36. Chione seymourensis Dall and Ochsner. Holotype, right valve, no.
2970 (Calif. Acad. Sci. Dept. Geol. Type Coll.), “from upper horizon, Seymour Island, Gala-
pagos Group Probably Pliocene.” Length 34 mm. Ficure 37. Chione seymourensis Dall
and Ochsner. View of the hinge of the specimen shown in figure 36. Ficure 38. Protothaca
(Tropithaca) cf. P. (T.) seymourensis Dall and Ochsner. Hypotype, right valve, no. 13659
(Calif. Acad. Sci. Dept. Geol. Type Coll.), from locality 27249 (CAS), Baltra (South
Seymour) Island, Galapagos Islands. Late Pliocene. Length 21 mm. Ficure 39. Protothaca
(Tropithaca) cf. P. (T.) seymourensis Dall and Ochsner. View of the hinge of the specimen
shown in figure 38.

40 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Turbo vermiculosus Dall and Ochsner.
(Figure 35.)

Turbo vermiculosus DALL AND OcusNER, Proc. Calif. Acad. Sci., Fourth Ser., vol. 17, no. 4,
p. 115, pl. 2, fig. 15, June 22, 1928; “frem upper horizon, Seymour Island, Galapagos
Group. Probably Pliocene.”

Three specimens of this species, none perfectly preserved, are present from
locality 27249 (CAS), Baltra (South Seymour) Island. The largest specimen
(figure 35) is 34.8 mm. high, the maximum diameter 23 mm.

This species has an unusually high spire for the genus. The surface micro-
sculpture “‘minutely vermiculately granulose and punctate,’ as mentioned by
Dall and Ochsner, is quite different from any described species of Turbo from
the western Americas.

The shape and sculpture of the present species somewhat resemble that of
Turbo caboblanquensis Weisbord (1962, p. 84, pl. 6, figs. 4, 5) described from
strata of Pliocene age in Venezuela but the shell of that species is umbilicate.

Polinices dubius Récluz.

(Figures 30, 31, 32, 33, 34.)

Natica dubia RéEctuz, Proc. Zool. Soc. London for 1843, p. 209, issued June, 1844 “Hab.
Chile? H. Cuming.” Sowersy, Thes. Conch., vol. 5, p. 86, pl. 458 (Natica, pl. 5), fig.
56, 1883. “Chili.” Rrrvr, Conch. Icon., vol. 9, Natica, species 41, pl. 10, figs. 41a, 41b,
1855. “Hab. Chili?” Tryon, Man. Conch., vol. 8, p. 47, pl. 16, fig. 50, 1886. “Chili, Peru.”

Polynices dubia Récluz, STEARNS, Proc. U. S. Nat. Mus., vol. 16, no. 942, pp. 401, 446, 1893.
“Indefatigable Island.”

One fairly well preserved specimen and four rather imperfect ones were
collected at locality 27249 (CAS). These specimens agree exactly with the
specimens identified by Dall and Ochsner under the name ‘“‘Neverita cf. reclusiana
Deshayes” (see figs. 32, 33, 34), from locality 804 (CAS). That locality is
approximately equivalent to locality 27249 (CAS).

One of the specimens from locality 804 (CAS) retains most of the umbilical
pad. The shape of the shell as well as the details of the callus agree closely with
illustrations of Polinices dubius, a species reported by Stearns from the Recent
fauna of the Galapagos Islands. Ranson (1959, p. 68) reported this species
from an elevated beach at Guadalupito, Peru.

The general shape of the Galapagos fossils is similar to that of the fossil
species described as ‘‘Natica solida” by Sowerby (1846, p. 255; see ed. 3, 1896,
p. 612, pl. 3, figs. 40, 41.) from “Navidad, Chile; Santa Cruz,ePataconiae?
| Miocene: Herm, 1969, p. 87] but the callus on the Galapagos species is much
more extensive and with an indentation on the umbilical margin. Sowerby’s
species was renamed Natica darwinti by Hutton (1886a, p. 334; see also 1886b,
p. 214) because of an earlier usage of the name WNatica solida by Blainville
(1825, p. 251).

Vor. XXXIX] HERTLEIN: PLIOCENE FOSSILS FROM BALTRA ISLAND 41

Nerita oligopleura Dall and Ochsner.
(Figures 12, 13, 14.)

Nerita oligopleura DALL AND OCHSNER, Proc. Calif. Acad. Sci., Fourth Ser., vol. 17, no. 4, p.
114, pl. 2, fig. 11; pl. 6, fig. 15, June 22; “upper horizon, Seymour Island, Galapagos
Group. Probably Pliocene.”

About 30 specimens of this species were collected at locality 27249 (CAS).
The smallest one is 6 mm. high, the largest one 20.5 mm. high and 21 mm. wide.

There are four coarse spiral ridges on the shell and, on well preserved speci-
mens, two small spiral threads are visible in the interspaces between the ribs.

The shell of this species is quite different from any described west American
species, as pointed out by Dall and Ochsner. The Galapagos fossil species bears
a general resemblance to Nerita asperata Dujardin from the Helvetian, Middle
Miocene, of France, illustrated by Cossmann and Peyrot (1917, pl. 7, figures
83, 84). However, close relationship with that species is not postulated.

Turritella broderipiana marmorata Kiener.
(Figures 26, 27, 28.)

Turritella marmorata KIENER, Spéc. Gén. et Icon. Coq. Viv., Famille Turbinacées, Twrritella,
p. 23, pl. 8, fig. 1 (two figs.), 1843-1844. “Habite.”

Over a dozen specimens of a Turritella, mostly incomplete, were collected
at locality 27249 (CAS). These agree closely with Kiener’s illustrations of
Turritella marmorata.

Kiener’s figures depict a rather slender, elongated shell, the whorls nearly
flat-sided, sculptured with fine, concentric threads, cream-colored with narrow
wavy longitudinal brownish-violet lines or flecks.

Reeve (1849, species 6, plate 2, figures 6a, 6b) placed T. marmorata in the
synonymy of 7. broderipiana under which name two figures were shown. One
(6a) represents a shell with fairly wide whorls, somewhat wider posteriorly below
the suture, resulting in a slightly sinuous outline of the outer lip of the last whorl.
The form shown in this illustration is similar to T. gonostoma Valenciennes.
The other figure (6b) represents a slender, elongated shell with nearly flat-sided
whorls comparable to Kiener’s illustrations of 7. marmorata.

Turritella broderipiana d’Orbigny (1840, p. 388), was originally described
but not illustrated from “les environs de Payta, ou elle a été péchée sur les fonds
de sable.” The type specimen illustrated by Keen (1966, p. 3, pl. 1, fig. 21)
reveals a slender, elongated shell with slightly concave whorls. Earlier, Keen
(1958, p. 290) considered T. marmorata to be a variant of the extremely variable
T. gonostoma Valenciennes (1832, p. 275; Kiener, 1843-1844, pp. 21-22, pl.
10, fig. 1). That species was originally described but not illustrated from
“Habitat ad oras Americae australis in portum Acapulco Mexicanorum.” The
posterior portion of the whorls on 7. gonostoma is often the widest; the color

42 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

of rather dense gray-black mottled markings in contrast to the much narrower
flammules of 7. broderipiana and T. b. marmorata.

A Recent specimen in the collections of the California Academy of Sciences,
closely resembling Kiener’s illustrations of 7. marmorata, collected by D. L.
Frizzell at Cabo Blanco, Peru, is 116 mm. long (incomplete), the body whorl
22.9 mm. in diameter. Other specimens collected at the same locality and at
Paita Bay, and at the mouth of the Brazos River at San Ramon south of Sechura,
Peru, have similar color markings but the shells are much less tapering with
correspondingly wider apical angle. One such shell collected by J. G. Marks at
San Pedro, Ecuador, is 133 mm. long (incomplete), the diameter of the last
whorl 38 mm. Evidently the slender and the broad shells either represent ele-
ments of a very variable series, or two distinct forms are living in the region of
Ecuador and Peru.

The fossils from the Galapagos Islands closely resemble the slender form
with flat-sided whorls shown in Kiener’s illustrations of 7. marmorata. In this
character they differ from the type specimen of 7. broderipiana (with slightly
concave whorls) shown in Keen’s (1966) illustration. On that basis, in the
present paper, the form described by Kiener is treated as a subspecies of T.
broderipiana.

Pilsbry and Olsson (1941, p. 43) mentioned that Turritella alturana Spieker,
described from strata of Miocene age in Peru is ‘‘so close to the recent species
| 7. broderipiana| that they cannot be consistently separated.”

The subgenus Broderiptella Olsson was proposed to include the 7. broderipiana
group represented in the northern South American region at least since middle
Miocene time. This group of turritellas was discussed by Merriam (1941, pp.
50-51) and by Woodring (1957, pp. 110-112).

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1849. Conchologia Iconica, vol. 5, Turritella, plates I-XI, and explanations, May to
June.
SLEVIN, J. R.

1959. The Galapagos Islands. A history of their exploration. Occasional Papers, Cali-

fornia Academy of Sciences, no. XXV, pp. I-X, 1-150, figs. 1-31, December 22.
SoweErBY, G. B., (Ist of the name)

1846. Descriptions of Tertiary fossil shells from South America. [Jn] Darwin, C. A.,
Geological observations on South America. Appendix to Part II. [See Darwin.]

1896. Edition 3, pp. 605-623, pls. 2-4.

STEARNS, R. E. C.

1893. Scientific Results of Explorations by the U. S. Fish Commission Steamer
Albatross. XXV.—Report on the mollusks of the Galapagos Islands, with
descriptions of new species. Proceedings of the United States National Museum,
vol. XVI, no. 942, pp. 353-450, pls. 51, 52 (map), September 29.

VALENCIENNES, A.

1832. Coquilles univalves de l’Amérique Equinoxiale, recueillies pendant le voyage de

MM. de Humboldt et Bonpland. [Jz] Humboldt, F. H. A., von, and A. J. A.

46

CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Bonpland, Voyage aux régions equinoxiales du nouveau continent. Paris, pt.

2, Recueil d’observations de Zoologie et d’Anatomie comparée, vol. 2, pp. 262—
SO), joll, S75

WEISBORD, N. E.

1962.

Late Cenozoic Gastropods from northern Venezuela. Bulletins of American
Paleontology, vol. 42, no. 193, pp. 1-672, pls. 1-48, March 5.

Wi.iAMs, H.

1966.

Wotr, T.
1895.

Geology of the Galapagos Islands. [Jn] The Galapagos. Proceedings of the
Symposium of the Galapagos International Scientific Project. R. I. Bowman,
editor, University of California Press, pp. I-X VII, 1-318, illustrated. [Williams,
Geology, pp. 65-70].

Die Galapagos-Inseln. Verhandlung der Gesellschaft fiir Erdkunde zu Berlin,
Bd. 22, pp. 246-265, pl. 3 [map].

Wooprinc, W. P.

1957.

Geology and Paleontology of Canal Zone and adjoining Parts of Panama. Geology
and description of Tertiary mollusks (Gastropods: Trochidae to Turritellidae).

United States Geological Survey, Professional Paper 306—-A, pp. I-IV, 1-145,
pls. 1-23, figs. 14 in text.

PROCEEDINGS

OF THE

CALIFORNIA ACADEMY OF SCIENCES
FOURTH SERIES

Vol. XX XIX, No. 4, pp. 47-53; 2 figs.; 1 table April 6, 1972

A NEW SPECIES OF THE SCORPIONFISH
GENUS HELICOLENUS FROM THE
NORTH PACIFIC OCEAN

By
Tokiharu Abe

Tokai Regional Fisheries Research Laboratory,

Ministry of Agriculture and Forestry, Katchidoki 5-5-1, Kyobashi P. O.,
Tokyo 104, and Zoological Institute, Faculty of Science,
University of Tokyo, Hongo, Tokyo 113, Japan
and

William N. Eschmeyer

California Academy of Sciences, Golden Gate Park, San Francisco,
California, U.S.A. 94118

Apstract: Helicolenus avius is described as a new species in the fish family Scor-
paenidae, subfamily Sebastinae. The type locality is the northwestern Pacific Ocean
between Japan and Midway Island at the southern end of the Emperor Seamount
Chain, 32°40/N., 172°17’E. to 35°05’N., 171°46’E., at a depth between 450 and 600
meters. A description and 2 figures are provided. Remarks on the genus Helicolenus
are given.

INTRODUCTION
In the summer of 1970, around 21 and 22 August, the Japanese vessel Daini-
Oriento Maru of the Tokusui Company, Ltd., was conducting night-time trawling
operations for Beryx spendens at the Emperor Seamount Chain between Japan
and the Hawaiian Islands. At several stations a total of more than 250 kg. of
specimens of a new scorpionfish of the genus Helicolenus was captured. The genus
Helicolenus is virtually worldwide in distribution, containing temperate and tropi-

1 Contribution no. B552 from the Tokai Regional Fisheries Research Laboratory, Tokyo. Supported in
part by U. S. National Science Foundation Grant GB-—15811.

[47]

48 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

cal offshore species. According to Matsubara (1943), the genus Helicolenus,
along with Sebastiscus, Hozukius, and Sebastes, constitutes the subfamily Sebas-
tinae. Recently, Eschmeyer and Hureau (1971) suggested that the genus
Sebastes may be a terminal North Pacific-evolving offshoot characterized by a
reduced rather than an incipient suborbital stay. They suggested that the genus
Sebastes may have evolved from a tropical ancestor, particularly one with a com-
plete suborbital stay and one sharing the characters found in the worldwide tropi-
cal and temperate genus Helicolenus and the Oriental genera Sebastiscus and
Hozukius. The discovery of another species of the genus Helicolenus may help
in understanding the evolution of the subfamily Sebastinae and the family Scor-
paenidae.

The new species is quite distinct from the other species referable to the genus
Helicolenus, particularly in having reduced head spination, smaller scales, longer
gill rakers, better developed toothed protuberances (dentigerous knobs) on the
anterior ends of the premaxillaries, and a forked caudal fin.

As a final introductory comment, we wish to mention that Helicolenus dac-
tylopterus frequently is the main ingredient in French Bouillabaisse, or the
Mediterranean fisherman’s stew. Perhaps this new species has similar qualities.
Although some specimens were discarded, 245.8 kg. of the present new species was
frozen quickly on board the collecting vessel and kept at —20°C. Mr. Tsujisaki,
who collected the specimens, has informed us that the fish kept at this tempera-
ture remain in good condition for sale for three months. Chemical analysis of
flesh from one specimen measuring 21 cm. in standard length was made by Mr.
Masa-aki Takeuchi (Tokai Regional Fisheries Research Laboratory) and showed
the following: body weight 212.5 grams, edible part 27 percent, water of flesh
72.6 percent, crude protein 19.2 percent, crude fat 5.6 percent, crude ash 1.3
percent, and calories per 100 grams 126.2. Species of Helicolenus frequently are
one of the dominant fishes in their habitat, and the new species may prove to be
of commercial value.

METHODS

Methods of measuring follow Eschmeyer (1969) and are similar to methods
used for other teleostean fishes with a few exceptions. Measurements originating
from the anterior end of the upper jaw (head length, snout length, standard
length, jaw length) are taken from the anterior end of the premaxillaries, includ-
ing the dentigerous knobs; pectoral fin length is measured from the base of the
first ray to the end of the longest ray, with the fin pointing back; caudal fin
length is measured from the posterior end of the hypural plate to the most distal
ray when the upper and lower caudal fin lobes are squeezed together. Termi-
nology of head spines for the genus Helicolenus is the same as used by Matsubara
(1943) and by Eschmeyer (1969).

Abbreviations of depositories of specimens are as follows: ABE—personal

VoL. XXXIX] ABE & ESCHMEYER: A NEW SCORPIONFISH 49

collection of the senior author; BMNH—British Museum of Natural History;
CAS—California Academy of Sciences; USNM—United States National Mu-
seum; ZIUT—Zoological Institute, Faculty of Science, University of Tokyo.

ACKNOWLEDGMENTS

The present writers take pleasure in expressing here their sincere thanks to
Messrs. Hisateru Tsujisaki and Shoji Tamayama, Division of Offshore Fishing,
Tokyo Branch, Tokusui Company, Ltd., Tokyo, for their cooperation with the sen-
ior author during the past three years, and especially for providing the specimens
used in the present study; and to Mr. Masa-ake Takeuchi for making the chemi-
cal analysis of the flesh of one specimen of the new species. Comments on the
manuscript were made by Dr. Lo-chai Chen and Mrs. Lillian Dempster. Mrs.
Dempster also helped with literature work and the selection of the scientific
name. Miss Pearl Sonoda aided in curatorial ways.

Helicolenus avius Abe and Eschmeyer, new species.
(Figures 1-2.)

MATERIAL EXAMINED. All specimens were collected in the northwestern Pa-
cific Ocean between Japan and Midway Island, at the southern end of Emperor
Seamount Chain, between 32°40’N., 172°17’E., and 35°05S’N., 171°46’E., at a
depth between 450 and 600 meters (probably most from 475 m.), with a bottom
trawl, vessel Daini—Oriento Maru, around August 21-22, 1970. Holotype: ZIUT
52457 (211 mm.S.L.). Paratypes: ABE 15256 (1, about 200 mm. T.L. [used for
description of color in life only|); BMNH 1971 12.14.1 (1, 173 mm. S.L.); CAS
13614 (2, 172-200 mm.S.L.) and CAS 13615 (1, 178 mm. S.L., cleared and
stained) ; USNM 206327 (2, 176-181 mm. S.L.).

DESCRIPTION. (Measurements and counts summarized in table 1; body shape
and coloration in figures 1-2.) Dorsal fin with 12 spines and usually 13% (13’2-
14%) soft rays. Anal fin with 3 spines and 6’ soft rays. Pectoral fin with 18-20
rays, most frequently 19; 3rd through 11th or 12th rays branched in available
specimens. Pelvic fin with 1 spine and 5 soft rays. Gill rakers total 31-34, 9 or
10 on the upper arch and 2224 on the the lower arch, increasing in size toward
angle of gill arch; longest raker about '2 of orbit diameter. Vertebrae 25.
Airbladder absent. Head spines mostly rudimentary or absent; preorbital
(lachrymal) bone with two weak spinous points over maxillary, sometimes vir-
tually absent; five preopercular spines present, second longest, all broad; supple-
mental preopercular spine absent; nasal spines present; opercular bone with two
spines; preocular spine well developed; supraocular spine weak or absent; post-
ocular and tympanic spines small, mostly covered by skin or developed as ridges
only; upper posttemporal spine poorly developed or absent; supracleithral spine
broad; nuchal spine sometimes present; other spines, including the parietal, sphen-

50 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

TABLE 1. Counts and measurements for the type specimens of Helicolenus avius.
(Measurements are in millimeters; see standard lengths in Material Examined section for
depositories and catalog numbers.)

Standard length 172 173 176 178 181 200 211
Dorsal rays 12-+13% 124-13% 124-134% 12+-13%4 12-+413% 12 eeeeiae
Anal rays 3+6% 346% 346% 346% 346% 346% 346%
Pectoral rays 19, 19 19, 19 20, 20 20, 19 18, 18 19, 19 19,19
Pelvic rays 1+5 1+5 1+5 1+5 1+5 1+5 1+5
Vertebrae 25 25

Gill rakers 10423 9424 9:44:22 9423 9-223) “iosepamemtoses
Head length 61.5 58.7 65.7 62.5 64.2 73.4 76.7
Body depth 48.8 49.1 53.4 50.3 50.7 56.9 57.4
Orbit diameter 18.0 17.3 17.9 18.5 17.8 2233 22.0
Snout length 15.6 13.6 16.0 14.7 16.4 16.8 17.9
Interorbital width 9.2 9.1 9.5 9.2 10.8 11.9 1222
Jaw length 30.0 29.6 28.3 30.1 30.8 34.7 3025
Predorsal fin length 61.3 57.8 61.5 59.3 62.8 68.7 (Bis:
Length 3rd dorsal spine 17.5 18.9 20.2 — — 22.8 PAL
Length 1st anal spine 8.0 6.9 6.3 7.8 8.0 8.9 8.7
Length 2nd anal spine 16.9 15.9 17.0 16.2 i7e5 tee 19.0
Length 3rd anal spine 17.2 16.2 18.1 15.6 18.5 18.2 18.6
Length pectoral fin 49.1 49.2 51.0 50.3 53.7 57.7 57.9
Length pelvic fin 31.4 29.0 S10 28.7 Sil 33.0 Basil
Length caudal fin 39.8 42.3 43.2 42.0 44.2 46.9 51.5

otic, coronal, pterotic, lower posttemporal, and suborbital spine absent or present
only as scarcely developed ridge. Scales on sides of body small, ctenoid; vertical
scale rows from supracleithral spine to end of hypural about 100, difficult to
count; pored lateral line scales about 50-55, plus 5 or more on the caudal fin;
most of head with cycloid scales, including the maxillary, cheek, and interorbital
area; snout unscaled; scales on belly, pectoral fin, and bases of vertical fins mostly
cycloid. Premaxillary, dentary, vomer, and palatine toothed; longest teeth on
the dentigerous knob of the premaxillary; all teeth short, conical, not arranged in
definite rows.

Color pattern of preserved specimens as in figures 1-2. Most conspicuous
feature the dark spots above and below the lateral line anteriorly and on the
back and dorsal fin. Buccal cavity black posteriorly, pallid anteriorly. Peri-
toneum black. (Color in life, according to Mr. Tsujisaki, was unchanged when
the frozen specimens were handed to the senior author. A color slide was made
on 12 October, 1970, of one specimen (ABE cat. no. 15256) by the senior author
soon after receipt of the specimens, and the following is taken from the slide.
The coloration was also illustrated in Abe (1970) ). Body mostly red above
and white ventrally. Dorsal surface of body and dorsal fin mottled with dark
brown spots on a red background; spots arranged as in figure 1 of a preserved
specimen. Pectoral and caudal fins red with yellowish tips.

VoL. XXXIX] ABE & ESCHMEYER: A NEW SCORPIONFISH 51

Ficure 1. Lateral view of Helicolenus avius, CAS 13614, paratype, 172 mm. in standard
length.

Name. The specific name avius is the Latin word for “‘out of the way, remote
or solitary.”’ This name is in reference to the type locality, an isolated seamount.
A Japanese name, ‘“‘okikasago,” meaning Scbastes-like or Helicolenus hilgendor fi-
like fish, was given to this species (Abe, 1970).

DIsTRIBUTION. This species is known only from the type locality. We expect
that it might occur at other seamounts in the northwestern Pacific and partic-
ularly along the Hawaiian Island chain.

COMPARISON AND REMARKS. The subfamily Sebastinae has been defined by
Matsubara (1943). Two of the four genera, Sebastes (including Sebastodes)
and Sebastiscus, have an incomplete suborbital stay which does not attach to
the preopercle. The other two genera of the subfamily, Helicolenus and Hozu-
kius, have a complete suborbital stay. Species of Helicolenus lack an airbladder
and have 25 vertebrae; while the single species of Hozukius has an airbladder
and 26 vertebrae; the species show cranial differences also.

At most there are five or six closely related known valid species or sub-
species belonging to the genus Helicolenus. Helicolenus dactylopterus (Dela-
roche, 1809) has a wide distribution in the Atlantic Ocean extending barely into
the Indian Ocean in South Africa; synonyms include H. maculatus (Cuvier,
1829), H. imperialis (Cuvier, 1829), H. maderensis Goode and Bean, 1896,
H. thelmae Fowler, 1937, H. uruguayensis Fowler, 1943, and as a subspecies
H. d. lahillei Norman, 1937 (see Eschmeyer, 1969). Helicolenus mouchezt
(Sauvage, 1875) is a senior synonym of H. tristanensis Sivertsen, 1945, and is
known from the south Atlantic at Tristan da Cunha and in the southern Indian
Ocean at Saint Paul and Amsterdam islands (see Eschmeyer and Hureau,

unr
bo

Ficure 2. Lateral and dorsal views of
paratype, 172 mm. in standard length.

the

head of

CALIFORNIA ACADEMY OF SCIENCES

Helicolenus

[Proc. 4TH SER.

avius,

CAS 13614,

VoL. XXXIX] ABE & ESCHMEYVER: A NEW SCORPIONFISH 53

1971). The nominal Oriental species is H. hilgendorfi (Steindachner and Déoder-
lein, 1884), although this species has been treated as H. dactylopterus by some
authors. Helicolenus lengerichi Norman, 1937, is known from the southeastern
Pacific. The Australian-New Zealand species is H. papillosus (Schneider, 1801)
with the following as synonyms, H. percoides (Richardson, 1842), H. cottoides
(Forster, 1849), H. barathri (Hector, 1875) and H. maccullochi (Phillipps,
1927) (see Whitley, 1968, p. 83). Helicolenus microphthalmus Norman, 1935,
was shown to belong to the genus Sebastiscus (Wheeler and Eschmeyer, 1968),
and H. rufescens Gilbert, 1905, from Hawaii belongs in the subfamily Scor-
paeninae (Eschmeyer, 1969). Helicolenus avius agrees with these six species
or subspecies in such features as counts of fin rays, general body shape, absence
of an airbladder, condition of the suborbital stay, and 25 vertebrae. Helicolenus
avius differs from the other species of Helicolenus in the following features.
In H. avius the caudal fin is forked while the caudal fin in the others is “‘square-
cut” or only slightly emarginate; the gill rakers are longer in H. avius; the
tubed lateral-line scales number more than 50 in H. avius while they are fewer
than 35 in the other species; the body scales are smaller in H. avius, about 100
vertical scale rows versus 80 or fewer; H. avius shows reduced spination, with
some spines which are normally found in the other species either poorly devel-
oped in H. avius, present as slight ridges, or absent; and H. avius, has the pro-
tuberance or dentigerous knob at the end of each premaxillary bone much better
developed.

LITERATURE CITED

ABE, TOKIHARU

1970. Shingao no sakana, 1970 nen ban [New faces of fishes, edition for the year 1970].
Publications of the Shigezo Ito Institute of Ichthyology, Tokyo. 1970, no. 2,
8 pp. (In Japanese).

ESCHMEYER, WILLIAM N.

1969. A systematic review of the scorpionfishes of the Atlantic Ocean (Pisces:
Scorpaenidae). Occasional Papers of the California Academy of Sciences,
no. 79, 130 pp., 13 figs.

EscHMEYER, WILLIAM N., and JEAN CLAUDE HUREAU

1971. Sebastes mouchezi, a senior synonym of Helicolenus tristanensis, with comments
on Sebastes capensis and zoogeographical considerations. Copeia, 1971, no. 3,
pp. 576-579.

MartsusBara, KiyoMATsuU

1943. Studies on the scorpaenoid fishes of Japan. The Transactions of the Sigen-

kagaku Kenkyusyo, no. 1, 486 pp., 4 pls.
WHEELER, ALWYNE, and WILLIAM N. ESCHMEYER

1968. The identity of the “British” scorpionfish Helicolenus microphthalmus. Jour-

nal of the Linnean Society (Zoology), vol. 47 (no. 312), pp. 309-314, 1 fig.
WHITLEY, GILBERT P.

1968. A check-list of the fishes recorded from the New Zealand region. The Australian

Zoologist, vol. 15, part 1, pp. 1-102.

PROCEEDINGS

OF THE

CALIFORNIA ACADEMY OF SCIENCES
FOURTH SERIES

Vol. XX XIX, No. 5, pp. 55-64; 1 fig.; 1 table. April 6, 1972

TWO NEW SCORPIONFISHES (GENUS
SCORPAENODES) FROM THE INDO-WEST
PACIFIC, WITH COMMENTS ON
SCORPAENODES MUCIPARUS (ALCOCK)

By

William N. Eschmeyer

California Academy of Sciences, San Francisco, California 94118
and

K. V. Rama Rao
Zoological Survey of India, Calcutta-16, India

ABSTRACT: Two new scorpionfishes of the genus Scorpaenodes are described from
the Indian Ocean. Additional comments are given on Scorpaenodes muciparus (Al-
cock). These three species are compared with each other and with S. tribulosus
Eschmeyer, the fourth species of the genus Scorpaenodes known from offshore areas
in the Indo-West Pacific.

INTRODUCTION

Eschmeyer (1969a) identified Indian Ocean specimens collected by the
United States International Indian Ocean Expedition as Scorpaenodes muciparus
(Alcock, 1889), offering a description and figures. Recently, we had the oppor-
tunity to examine two specimens of S. muciparus in the collection of the Zoological
Survey of India, and one of us (W.N.E.) was also able to examine the specimen
collected by the Siboga which was identified as S. muciparus by Weber (1913)
and also by de Beaufort (Weber and de Beaufort, 1962). It was then apparent
that the specimens identified by Eschmeyer (1969a) as S. muciparus were not
specimens of that species but represented a new species, a species which one of
us (K.V.R.R.) was in the process of describing as part of current studies on the

Marine Biological Laboratory
LIBRA Pv

APR141972

\VAlan nt | aT ee ee a {

[55]

56 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

scorpionfishes of India. Examination of scorpionfishes in other museums has
resulted in location of yet another closely related species, the description of which
we include in this paper. These three species, along with S. tribulosus (Esch-
meyer, 1969a), live in offshore level-bottom areas, while other members of the
genus are shallow-water rock or reef inhabitants.

ACKNOWLEDGMENTS

The senior author acknowledges the support of National Science Foundation
Grant GB-15811, which permitted him to examine scorpionfishes in most of the
major fish collections. The following museums contained specimens used in this
study, and we wish to thank the curators and staffs listed for courtesies extended
during his visits to their museums: BMNH, British Museum of Natural History
(N. B. Marshall, G. Palmer, A. Wheeler, P. Whitehead) ; CAS, California Academy
of Sciences (L. Dempster, P. Sonoda); NMS, National Museum of Singapore
(E. Alfred); USNM, United States National Museum (N. Gamblin, V.
Springer); ZMA, Zoological Museum, Amsterdam (H. Nijssen); ZMC, Zoologi-
cal Museum, Copenhagen (J. Nielsen) ; ZSI, Zoological Survey of India (A. G. K.
Menon).

Lillian Dempster, Pearl Sonoda, Kathy Smith, Terry Arambula, and Maurice
Giles of the California Academy of Sciences, assisted in the study. We are grate-
ful for comments on the manuscript made by Lillian Dempster and A. G. K.
Menon.

METHODS

Counts, measurements, and terminology of head spines are as used by Esch-
meyer (1969b, pp. 4-6).

SPECIES ACCOUNTS

The limits of the genus Scorpaenodes have been discussed recently (Esch-
meyer, 1969a). Within the subfamily Scorpaeninae, the genus is characterized
by the following combination of characters: normally thirteen dorsal spines,
spinous procurrent caudal fin rays, and no palatine teeth. The genus contains
species which are primarily tropical, shallow-water forms, and species of the genus
are found in all warm oceans.

The four species of Scorpaenodes treated in the present paper are the only
ones of the genus occurring in offshore Indo-Pacific waters below a depth of about
20 fathoms. These four species may be separated as follows:

1. Vertical scale rows (above lateral line from first lateral line scale to end of hypural)
OMOT oI) O1 CG ae ee 2 nee oe ee Se S. muciparus
1. “Vertical ‘scale rows about 50 or fewer 2.20 Eee 2

2. Soft dorsal rays 8%, scales strongly ctenoid with long ctenii, underside of head covered
with ctenoid scales —___ ES GV ANE en Ye ee S. tribulosus

Vor. XXXIX ESCHMEYVER & RAO: TWO NEW SCORPIONFISHES 57

2. Soft dorsal rays usually 914, scales ctenoid, underside of head naked or with cycloid
SOPUBS en eed EE Se ee ee ae Se ee en ee a 3

Remarks. A new species, Scorpaenodes steinitzi, was described recently by
Klausewitz and Frgiland (1970) from the Red Sea. Seventy-five additional speci-
mens of this species are available to us. Two of the major differentiating char-
acters used by these authors were that the dorsal and anal soft ray counts were 1
higher in S. steimitzi than in the other species, but this results from the fact that
Klausewitz and Fr¢iland counted the last double ray as 2 rays, while recent
authors count this as 1 or 1'2 rays. Despite this, Scorpaenodes steinitzi appears
to be a valid species which is distinguishable from other species most easily by its
coloration. We mention this because readers may wonder if S. steinitzi might not
be one of the species treated here by us. Scorpaenodes steinitzi is a pale-colored
species with some brown pigment on the sides and it usually has a dark spot near
the posterior end of the spinous dorsal fin, but S. steinitzi does not have the
lateralis canal head pores well developed and it is a shallow-living species. In
S. steinitzi the spots on the caudal fin when present are arranged in rows while
they are more scattered in S. smithi; and the dark pigment on the sides in S.
steinitzi is not arranged in definite bars as in S. smithi, S. investigatoris, and S.
muciparus.

Scorpaenodes investigatoris Eschmeyer and Rama Rao, new species.

Scorpaenodes muciparus, ESCHMEYER, 1969a, pp. 4-8, figs. la, b, table 1 (description; these
specimens now referred to this new species, not S. muciparus (Alcock, 1889).

Scorpaenodes varipinnis, SMiTH, 1957, p. 65 (in part; only the 50 mm. specimen taken from
the stomach of a specimen of Pristipomoides microlepis Bleeker collected in 110 fathoms
off Memba and tentatively referred to S. varipinnis).

REMARKS. The specimens wrongly identified by Eschmeyer (1969a) as
S. muciparus (Alcock) are designated as the types of this new species along with
two specimens from the collection of the Zoological Survey of India. The holo-
type (CAS 24264) and one paratype (CAS 24265) were figured by Eschmeyer
(1969a, fig. la, b).

TYPE MATERIAL. Holotype. CAS 24264 (1, 71.2 mm. S.L.), West Pakistan,
24°13’ N., 65°52’ E., in 93 fathoms, Anton Bruun Cruise 4B, station 267B, 9
December 1963.

Paratypes. CAS 24266 (1, 87.4 mm. S.L.), West Pakistan, 25°02’ N.,
56°52’ E., in 159 fathoms, Anton Bruun Cruise 4B, station 264A, 2 December
1963; CAS 24265 (2, 45.8-48.9 mm. S.L.), off western India, 22°32’ N., 68°07’

58 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

E., in 31.5 fathoms, Anton Bruun Cruise 4B, station 221A, 18 November 1963;
USNM 204030 (1, 72.4 mm. S.L.), off western India, 17°25’ N., 71°39’ E., in
52.5 fathoms, Anton Bruun Cruise 4B, station 202A, 13 November 1963; ZSI
916/1 (1, 86 mm. S.L.) and ZSI 917/1 (1, 67 mm. S.L.), Arabian Sea, off Rat-
nagiri Coast, 17°27’ N., 71°41’ E., 56-58 fathoms, bottom of fine sand, Agassiz
trawl haul, /nvestigator station 242, 11 October 1898. (Smith’s specimen men-
tioned above in the Synonymy is regarded only as an additional specimen and
not a type.)

DeEscrIPTION. See Eschmeyer (1969a, pp. 4-8, figs. la and 1b, table 1) for
a complete description of this species. The two ZSI specimens have the following
counts: Dorsal rays XIII, 9% anal rays III, 5’2-6'2; pectoral rays 19. The
vertical scale rows in the two specimens number 45-48, and the gill rakers num-
ber 5—6 plus 12-13. Except for the specimen with 6’ soft anal rays, the counts
fall within those given by Eschmeyer (1969a, table 1).

DISTRIBUTION. Scorpaenodes investigatoris is known from the Gulf of Oman
and in the Arabian Sea off western India and West Pakistan. The specimen re-
ported by Smith (1957, p. 65) was taken off Memba, Mozambique. Depths of
capture range from 31 to 159 fathoms.

CoMPARISONS. This species very closely resembles Scorpaenodes smithi in
scalation, counts, and coloration. Scorpaenodes investigatoris and S. smithi may
be easily separated from the other two offshore species of Scorpaenodes: S.
muciparus has over 70 vertical scale rows as opposed to about 50 in S. investigatoris
and S. smithi, and S. tribulosus has strongly ctenoid scales on the underside of the
head while the other species have the underside of the head mostly naked or with
cycloid scales. The descriptions of S. investigatoris and S. smithi are almost
identical. The two characters which may be used to separate them are
pectoral rays and the coloration of the rear of the buccal cavity. In S.
investigatoris the pectoral rays are usually 19 (18-20) while they usually
number 17 or 18 (16-19) in S. smithi. In S. investigatoris the rear of the mouth,
including the pharyngeal bones and the areas adjacent to them, is dusky in color-
ation while this area is pallid in S. smithi. No other clear-cut differences were
noted. Scorpaenodes investigatoris appears to live at deeper depths than S. smith.

Etymo.Locy. The name is based on the Royal Marine Survey Steamer
Investigator, which made important collections in the Indian Seas during the years
1884-1926; the name “‘investigatoris” was used by one of us (K.V.R.R.) for this
species in his unpublished thesis on Indian Ocean scorpionfishes.

Scorpaenodes muciparus (Alcock).

Sebastes muciparus ALCOCK, 1889, pp. 298-299, fig. 3 on pl. 22 (original description; type
locality Bay of Bengal, 26 miles N. by E. of Gopalpur, in 45 fathoms). Atcock, 1898,

pl. 18, fig. 5 (good illustration). WerBER, 1913, p. 491 (brief description; one from
Siboga station 306, Solor Straits, 8°27’S., 122°54’W., in 247 meters).

Vor. XXXIX ESCHMEYER & RAO: TWO NEW SCORPIONFISHES 59

Scorpaenodes muciparus, de Beaufort in WEBER and DE BEAUFORT, 1962, pp. 34-35 (redescrip-
tion of Siboga specimen described by Weber).

MATERIAL EXAMINED. ZSI 1179/1 (1, 80 mm. S.L.) Gulf of Martaban,
14°46/N., 95°52’E., in 61 fathoms, on soft, green muddy and sandy bottom,
Agassiz trawl haul, /nvestigator station 328, 7 March 1904; ZSI 12432 (1, 73 mm.
S.L.) Bay of Bengal, Ganjam coast, 19°24’N., 85°E., in 23 fathoms, muddy
bottom, Blake trawl haul, /nvestigator station 42, 7 March 1889; ZMA 110.246
(1, 146 mm. S.L.) Indonesia, Solor Straits, 8°27’S., 122°54’E., in 135 fathoms,
Siboga station 306, 8 Feb. 1900.

REMARKS. The reader is referred to the references listed above for a more
complete description of this species. The holotype of this species, which was
originally in the ZSI collection, has been lost. If a neotype is ever needed, we
suggest that the specimen listed above from /nvestigator station 42 (ZSI 12432)
would make the best neotype, as it is from the same general locality as the type
and was identified as S. muciparus by Alcock. Scorpaenodes muciparus is charac-
terized by a dorsal ray count of 13 spines and 9’ soft rays, pectoral rays of 18-19,
and vertical scale rows of over 70. The other three species treated in this paper
have a scale count of 50 or fewer. De Beaufort (in Weber and de Beaufort, 1962,
p. 35) gives a scale count of 45—58 for the Siboga specimen. Re-examination shows
that the specimen has a vertical scale row count of about 70-73 when the scale
rows are counted above the lateral line from the supracleithral spine to the end of
the hypural plate.

Scorpaenodes tribulosus Eschmeyer.

Scorpaenodes tribulosus ESCHMEYER, 1969a, pp. 8-10, fig. 1c, table 1 (original description;
type locality East Africa, off Kenya, 02°42’S., 40°53’E., in 77 fathoms and off the
Somali Republic, 11°24’N., 51°35’E., in 40-96 fathoms).

REMARKS. The species is still known only from the western Indian Ocean,
from the holotype (CAS 24267, off Kenya) and one paratype (USNM 204031,
off the Somali Republic) as given above. The reader is referred to Eschmeyer
(1969a) for a description of this species. This species is characterized by a dorsal
ray count of 13 spines and 8% soft rays, pectoral rays 19, and vertical scale rows
under 50. Scorpaenodes tribulosus is most easily distinguished from the other
species by the presence of strongly ctenoid scales on the underside of the head;
the other three species have this area covered with cycloid scales or mostly naked.

Scorpaenodes smithi Eschmeyer and Rama Rao, new species.
(Figure 1.)

TYPE MATERIAL. Holotvpe. BMNH 1929.6.12.6 (52.2 mm. S.L.), Andaman
Sea, off western Malaya, 5°45’N., 98°20’E., in 40 fathoms, Madras- Penang Cable
Survey, Cable ship Patrol, 16 March 1929.

[Proc. 4TH Serr.

CALIFORNIA ACADEMY OF SCIENCES

60

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Vor. XXXIX ESCHMEYVER & RAO: TWO NEW SCORPIONFISHES 61

Paratypes. BMNH1929.6.12.7-15 (12, 21.5-47.0 mm. S.L.), and CAS 13616
(4, 23.2-52.1 mm. S.L.) and ZSI F6254/2 (1, 46.0 mm. S.L.), formerly in BMNH
1929.6.12.6—-15, taken with the holotype; BMNH 1932.2.15.15-20 (6, 24.5-48.3
mm. S.L.), Andaman Sea, west of Malaya, 6°01’20’N., 99°03’05”E., in 46
fathoms, mud bottom, Madras-Penang Cable Survey, no date; BMNH
1972.1.17.1-9 (9, 22.0-51.3 mm. S.L.) and USNM 206501 (4, 22.4—45.6
mm. S.L.) west of Malaya, 80 miles south of Penang, in 24 fathoms, en-
tangled in coelenterates found on cable, cable ship Patrol, May 1923; BMNH
orot.4 (1, 35.6 mm. S.L.), Arafura Sea, 11°59’30’S., 126°38’E., in
60 fathoms, E. and A. Telegraph Company, collected in or before 1933;
Mie P7982—-83 (2, 56.5 mm. S.L.), Gulf of Tonkin, 20°26’N., 108°09’E.,
in 28 fathoms, collected by Schgnau in or before 1895; ZMC P7978-81
(4, 46.2-60.8 mm. S.L.), China Seas, collected by FE. Svenson, in or
before 1893; ZMC P79100-112 (13, 34.0-62.4 mm. S.L.), southwest of Hong
Kong, 26°10’N., 121°00’E., in 44 fathoms, collected by H. Christiansen, 31 May
1912; ZMC P79113-132 (20, 28.7-46.8 mm. S.L.), off western Malaya,
5°09’08’”N., 99°48’10”E., in 32 fathoms, sandy mud bottom, collected by Store
Nordiske, 24 Dec. 1935; ZMC P7991-92 (2, 32.1-58.8 mm. S.L.), Formosa
Strait, ““Namoa” Island, collected by Capt. Svenson, in or before 1912; NMS
uncataloged (10, 15.5-54.4 mm. S.L.), near Singapore 5°59’06”N., 99°08’33”E.,
in 40 fathoms, collected by Tweedie. A total of 89 specimens.

DEscriPTION. Measurements and counts summarized in table 1; body shape
and coloration in figure 1.

Dorsal fin rays normally XIII, 9'%, rarely 8’ or 10’. Anal fin rays normally
III, 5%. Pectoral rays usually 17 or 18 (16-19), rays 2 or 3 through 7 to 9
branched in larger specimens; smaller specimens with fewer branched rays; the
branching of pectoral rays begins at about 25-30 mm. S.L. Spines on head
well developed. Preorbital bone with second and third lobes (Matsubara’s, 1943,
terminology) each as a broad spinous point extending over the maxillary, third
lobe usually with additional spinous points; suborbital ridge with one row of
spinous points, usually one and sometimes several spines on the lateral face of the
preorbital bone in line with three or more spinous points on the suborbital bones,
the posterior one frequently double. Tympanic spines present; coronal spines
usually absent, sometimes present only on one side, rarely present on both sides.
Interorbital ridges rarely ending in spines. Small spines (postfrontal spines of
Smith, 1957) near midline between tympanic spines sometimes present. Supple-
mental preopercular spine present. First preopercular spine long, second usually
small, third present, fourth usually present, points ventrally, and fifth points
down (sometimes virtually absent). Preocular, supraocular, and postocular
spines well developed. Small spine below parietal spine usually absent. Other
spines present include the nasal, parietal, nuchal, sphenotic, pterotic, upper (some-

62 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH Ser.
TABLE 1. Counts’ and measurements of type specimens of Scorpaenodes smithi.
(Measurements are in millimeters; numbers in parentheses are percentages of standard length.)

BMNH
1929.6.12.6 CAS 13616 P7982-3

Standard

length Sf S2el 48.7 44.2 23.2 56.5 56.5
Dorsal fin

rays + XIII+8% XIII+9% XIII+9'4 XII+9% XIII+9% XIII+9% XIII+8%
Anal fin

rays ! TI+5% Wi+5% WI+5% IWI+5% IiI+5% I+6% Ii+5%
Pectoral fin

rays ! Vaats 1st; isseis | 1ssise es-ES 18-P1g) sels-oe
Head length 22.6(43 ) 23.3(45) 21.2(44) 19.1(43) 10.7(46) 24.3(43) 22.0(39)
Snout length 6.1(12) 6.5(12) 6.2(13) SAID) BAU) 6.5(12) 6.2(11)
Orbit diameter 6.8(13) 6.8(13) 6.7(14) 5.9(13) 3.2(14) 6.2(11) 6.7(12)
Interorbital

width 3.3(06) 3.0(06) 2.8(06) 2.9(06) 1.7(07) 3.8(07) 3.2(06)
Upper jaw

length 11.9(23) 1320 C25) leo (24) O)7(22)) Sash(7S))) 12°42) 122222)
Predorsal-fin

length 20.7(40) 21.4(41) 20.0(41) 18.3(41) 10.3(44) 21.0(37) 20.4(36)
Body depth 2222)\(43)) 21.5(41) 19.6(40) 17.0(38) 9.9(43) 22.7(40) 22.0(39)
Pectoral fin

length 17 (SS) 19.1(37) 16.5(34) 15.8(36) 7.4(32) 19.2(34) 21.5(38)
Pelvic fin

length 13.9(27) 135/(26)) 12:6 (26) 112526) 5.7(24) 14.2(25) 14.8(26)
Caudal fin

length 14.8(28) TAPAS) elo O28) aeliZesiC2.8)) 7.0(30) 14.8(26) 15.3(27)

ZSI F6254/2 BMNH 1932.2.15.15-20

Standard length 46.0 48.3 46.6 48.1 47.8 32.2 24.5
Dorsal fin

rays XIII+9'% XII+10% XIII+9'4 XIII+9% XII+9% XIII-+9% XIII-+9%
Anal fin rays I+5% Wl-+5% I1-+5% I1+5% 145% II+5% IiI+5%
Pectoral fin

rays 17+18 17-+-17 18+18 17+17 17+17 18+18 18+18
Head length 19.8(43) 21.5(44) 20.3(44) 19.8(41) 19.7(41) 14.2(44) 11.4(46)
Snout length 5.2(11) 5.7(12). 5.2011) 5,5(11) .S4(11). 4eCie ieee

VoL. XXXIX ESCHMEYER & RAO: TWO NEW SCORPIONFISHES 63

(Table 1 Continued)

ZSI F6254/2 BMNH 1932.2.15.15-20

Orbit diameter 5.7(12) 6.0(12) fl) 5.9(12) 6.0(12) 4.1(13) 3.6(15)

Interorbital

width 2.7(06) 3.2(07) 2.8( 06) 3.0(06) 3.2(07) 2.2(07) 1.8(07)
Upper jaw

length 10.0(22) PROCS) V1OSC2) 10422) e022) 7.5(23) 6.2(25)
Predorsal fin

length 18.8(41) 19.1(40) 18.6(40) 19.4(40) 19.0(40) 14.2(44) 11.2(46)
Body depth 19.0(41) 18.8(39) 18.8(40) 19.8(41) 20.0(42) 13.2(41) 10.6(43)
Pectoral fin

length 14.6(32) 14.5(30) 14.4(31) 14.0(29) 14.4(30) 9.0( 28) 7.7(31)
Pelvic fin

length 12.2(26) IO AS) SAAT) UAE@aD) aC) 8.0(25) 7.0(28)
Caudal fin

length 13.0(28) 124 (26) 2.827) 12292) 126 (26) 8.6(27) 7.1(29)

1 Counts of 63 additional specimens give the following totals (12 small specimens not counted): Dorsal fin
rays XIII+916(70 specimens), XIII+814(2), XIII+9(1), XII+1014(1), XIII+1014(2), and XIII4+-5%(1,
abnormal); Anal fin rays III+514(75 specimens), III+614(1), I4614(1, abnormal); Pectoral fin rays
16+17(1 specimen), 17+17(22), 17+18(6), 18+17(7), 184+18(37), 184+19(1), 19+18(2), 19+19(1).

times absent) and lower posttemporal, two opercular, cleithral, and supracleithral
(double). Fourth suborbital bone isolated, bearing a few small spines (Gins-
burg’s, 1953, postorbital spines). Tentacles and other fleshy appendages associ-
ated with most head spines. Supraocular tentacle about % to ' of orbit diameter,
sometimes reduced, with branches distally. A few tabs on upper part of eye,
two usually larger than the others. Scales on flanks ctenoid; vertical scale rows
about 40 to 50, usually in middle forties; lateral-line scales usually 23 plus 1 or
2 on caudal fin. Bases of fins with a few scales. Scales on chest and pectoral
fin base mostly cycloid. Dorsal parts of head scaled; underside of head unscaled
or with cycloid scales; snout unscaled. Gill rakers including rudiments 15-17;
upper arch with 5—6 short spiny rakers; lower arch with 9-10 rakers on cerato-
branchial and usually 2 rudiments on hypobranchial. Small slit present behind
fourth gill arch. Head pores well developed. Color pattern as in figure 1. Most
conspicuous feature is the black spot on the posterior part of the spinous dorsal
fin. Body usually with four broad bands, first under anterior dorsal spines, second
under posterior spinous dorsal fin, third between soft dorsal fin and anal fin,
fourth at base of caudal fin. All fins with brown or black spots. Three brown bars
radiating from eye; first extends forward across preorbital bone, second below
eye, and third extends down and back from posterior ventral part of eye.
DISTRIBUTION. Scorpaenodes smithi seems to be widespread in the area of

64 CALIFORNIA ACADEMY OF SCIENCES [Proc. 47H Serr.

the South China Sea and through the Straits of Malacca into the Andaman Sea.
A wider distribution in the Indo-Australian Archipelago is expected. Depths
of capture range from 24 to 60 fathoms, and the bottom type appears to be mud

or sand.

COMPARISONS. See the Comparisons section under the account of Scorpaenodes
investigatoris.

EtyMoLocy. We name this species in honor of the late Professor J. L. B.
Smith for his noteworthy contributions to the knowledge of Indian Ocean scor-
pionfishes.

LITERATURE CITED

Atcock, ALFRED
1889. Natural history notes from H. M.’s Indian Marine surveying steamer /nvesti-
gator, Commander Alfred Carpenter, R.N., D.S.O., commanding. No. 12.
Descriptions of some new and rare species of fishes from the Bay of Bengal,
obtained during the season of 1888-89. Journal of the Asiatic Society of
Bengal, vol. 58 (pt. 2, no. 17), pp. 296-305, pl. 22.
1898. Illustrations of the zoology of the Royal Indian marine surveying steamer
Investigator, . . . Fishes—part 5. Calcutta, pls. 18-24.
ESCHMEYER, WILLIAM N.
1969a. A new scorpionfish of the genus Scorpaenodes and S. muciparus (Alcock) from
the Indian Ocean, with comments on the limits of the genus. Occasional Papers
of the California Academy of Sciences, no. 76, 11 pp., 1 table, 1 fig.
1969b. A systematic review of the scorpionfishes of the Atlantic Ocean (Pisces: Scor-
paenidae). Occasional Papers of the California Academy of Sciences, no. 79,
130 pp., 13 figs., 15 tables.
GInsBurG, ISAAC
1953. Western Atlantic scorpionfishes. Smithsonian Miscellaneous Collections, vol. 121,
no. 8, 103 pp., 6 text figs.
KLAUSEWITz, WOLFGANG, and @ysTEIN FROILAND
1970. Scorpaenodes steinitzi n. sp. von Eilat, Golf von Aqaba. Senkenbergiana
Biologica, vol. 51, no. 5/6, pp. 317-321, 2 figs.
MATSUBARA, KrvYOMATSU
1943. Studies on the scorpaenoid fishes of Japan. The Transactions of the Sigenkagaku
Kenkyusyo, no. 1, 486 pp., 4 pls.
SMiTH, J. L. B.
1957. The fishes of the family Scorpaenidae in the western Indian Ocean. Part 1.
The subfamily Scorpaeninae. Ichthyological Bulletin, Rhodes University, no. 4,
pp. 49-69, pls. 1-4.
WEBER, MAx
1913. Die fische der Siboga-Expedition, Siboga Expedition, vol. 57, 710 pp., pls. 1-12,
text figs. 1-123.
WEBER, Max, and L. F. pE BEAUFORT.
1962. The fishes of the Indo-Australian Archipelago, XI. Scleroparei, Hypostomides,
Pediculati, Plectognathi, Opisthomi, Discocephali, Xenopterygii. E. J. Brill,
Leiden, 481 pp., 100 text figs.

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Marine Blological Laborate

“se tote

PROCEEDINGS | Wocds Hole, Mass.

OF THE

CALIFORNIA ACADEMY OF SCIENCES
FOURTH SERIES

Vol. XXXIX, No. 6, pp. 65-74, 22 figs. September 5, 1972

NEW BARNACLE RECORDS (CIRRIPEDIA,
THORACICA)
By

Victor A. Zullo, Dea B. Beach, and James T. Carlton
California Academy of Sciences, San Francisco, California 94118

Balanus amphitrite amphitrite Darwin, 1854.
(Figures 1-4.)

In discussing the introduction of Balanus improvisus Darwin to the Pacific
Coast of North America, Carlton and Zullo (1969) noted that although this
species was relatively common in collections dating back to 1853, no specimens
of B. amphitrite amphitrite were found in collections made prior to 1939, when
it was first observed by F. L. Rogers in San Francisco Bay (Henry, 1942). The
establishment of B. improvisus on the Pacific Coast appears to be tied to the
commercial importation of the North American East Coast oyster, Crassostrea
virginica (Gmelin). Both barnacles are frequent epizooites of this oyster, but
as B. amphitrite amphitrite is a relatively recent addition to the fauna of the
East Coast as well, Carlton and Zullo concluded that its absence in early
Pacific Coast collections reflected a similar absence on the East Coast. How-
ever, further inquiry into the history of the Pacific Coast oyster industry has
revealed that all importations of Crassostrea virginica were from areas north
of Cape Hatteras, North Carolina (the northern range limit of B. amphitrite
amphitrite), thus virtually eliminating this oyster as a vehicle of introduction
for B. amphitrite amphitrite.

While processing miscellaneous collections of barnacles for the Academy of
Natural Sciences of Philadelphia (ANSP), two lots of B. amphitrite amphitrite
were found that had been obtained from the California coast prior to 1939. This
material, collected by Charles R. Orcutt and originally identified by Henry A.
Pilsbry, consists of four specimens with opercular valves from La Jolla, Cali-
fornia, and of a single, large (20 millimeter basal diameter) specimen from San
Diego Bay, California. Both lots bear the number “158/428”, but no date of

[65]

66 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Ficures 1-4. Balanus amphitrite amphitrite Darwin. Figure 1, scutum, height 4 mm.;

figure 2, shell, greatest diameter 20 mm.; figures 3-4, terga, height 4 mm. (Figures 1, 3,
4, La Jolla, California; figure 2, San Diego Bay, California). Ficures 5-6. Opercular
valves of Balanus improvisus Darwin, CAS locality 41227, Delta Mendota Canal, California,

height of scutum and tergum 3.8 mm.

collection. Dr. Robert Robertson and Nancy Rulen of the Philadelphia
Academy were kind enough to search their records, and were able to provide
the following data:
ANSP no. 2257. La Jolla, California, C. R. Orcutt collector, donated
by the U. S. National Museum and cataloged December 6, 1921.
ANSP no. 2426. San Diego, California, C. R. Orcutt collector, De-
cember 9, 1927.

The apparent establishment of B. amphitrite amphitrite in southern Cali-
fornia by at least the early 1920’s and in the late 1930’s in northern California
is likely attributable to transport by ships. Its occurrence may eventually be
placed considerably earlier than this, for it had already arrived in Hawaii by
the early 1900’s (Pilsbry, 1907, p. 190). Balanus amphitrite amphitrite is
restricted to certain estuarine parts of the San Francisco Bay complex where
mean annual temperatures are highest, but even in these areas Newman (1967)
has noted that the temperature regime is clearly suboptimal for this subspecies.
If the establishment of B. amphitrite amphitrite in San Francisco Bay was

VoL. XXXIX] ZULLO, BEACH, & CARLTON: BARNACLE RECORDS 67

indeed later than in southern California, the delay may be explained by the
paucity of suitable environments for successful colonization.

Balanus eburneus Gould, 1841.
(Figures 8-15.)

Balanus eburneus, whose natural range includes the Atlantic coasts of North
America and northern South America, the Caribbean Sea, and the Gulf of
Mexico, has been introduced by shipping to various parts of the world. Known
introductions, emphasizing those in the Pacific Ocean, were summarized by
Matsui e¢ al. (1964) and Utinomi (1966). Definite Pacific localities include
Balboa, in the Bay of Panama, eastern Pacific, the Hawaiian Islands, and the
Japan Sea coast of central Japan. Weltner (1897) reported B. eburneus on
Fasciolaria with B. amphitrite from Manila in the Philippines, but this identi-
fication has not been corroborated.

A sample in the collection of the Department of Invertebrate Zoology of the
California Academy of Sciences (CAS) taken by Mr. Vern Brock in July, 1967,
from between Ansala and Vera islands in Eniwetok Atoll in the Marshall Islands
was found to contain several individuals of B. eburneus in association with B.
amphitrite amphitrite. These specimens represent the first record of B. eburneus
from islands of the central West Pacific, and no doubt reflect the heavy traffic
in American shipping to that area since the Second World War. The occurrence
of this Atlantic American species at Eniwetok suggests that other highly
frequented ports in the central Pacific islands might also harbor successfully
introduced populations of B. eburneus.

This species was also discovered recently at another, albeit more likely,
locality in the western Atlantic. Mrs. Joleen Gordon, in the course of an
ecologic study of Bermuda barnacles during the summer of 1967, found B.
eburneus to be a common member of that fauna in association with B. amphitrite
amphitrite and Chthamalus stellatus thompsoni Henry.

The barnacle fauna of the Bermudas was investigated most recently by
Henry (1958) on the basis of collections made by T. A. and Anne Stephenson
in the early 1950’s. Balanus amphitrite hawaiiensis Broch (= B. amphitrite
amphitrite) and Chthamalus stellatus thompsoni were reported there for the
first time, but Balanus eburneus was not found. Judging from the abundance
of B. eburneus and its association with the above-mentioned subspecies as seen
in 1967, it is possible that this species has been introduced to the Bermuda
fauna since the date of the Stephensons’ survey.

Balanus improvisus Darwin, 1854.
(Figures 5-7.)

A living population of B. improvisus was discovered in December, 1962, on
the concrete lining of a section of the Delta-Mendota fresh water irrigation

68 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Ss
&
°
=
.
S
&
—_
(D ‘.
Se ,
Antioch ~
Z
e
Oe*s
o OES
> Rae oO
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Ficure 7. Relationship of Delta-Mendota Canal system to the San Francisco Bay-

Delta area. Numbers indicate Tracy Pumping Plant (1) and collection site of Balanus
improvisus (2).

canal in central California. The barnacles, collected by G Dallas Hanna and
Allyn G. Smith of the California Academy of Sciences (CAS locality 41227),
were taken at mile post 20.62, or more than 20 miles downstream from the
Tracy Pumping Plant, which is about 9 miles northwest of Tracy, San Joaquin
County, California (figure 7). This section of the canal was discussed previously
by Hanna (1966, p. 40, figs. 30-31) with respect to the phenomenal infestation
of the Asiatic fresh water clam Corbicula manilensis (Philippi). The occurrence
of barnacles in the canal was noted by Prokopovich (1968, p. 53, photo-
graph 48).

Irrigation water is pumped into the Delta-Mendota Canal by the Tracy
Plant via a 2.3-mile-long canal from the Old River Channel of the San Joaquin
River. This intake is well over 25 miles from the established salt water barrier
of the Sacramento-San Joaquin estuary in the vicinity of Antioch, California.

Vor. XXXIX] ZULLO, BEACH, & CARLTON: BARNACLE RECORDS 69

Continuing water quality analyses made at the pumping plant indicate that
total dissolved solids do not exceed 500 parts per million (0.5 parts per
thousand) at the canal intake.

Although a few species of barnacles are known to live in river estuaries
where salinities are negligible for most of the year, there has been only one
other report of an apparent occurrence in totally fresh water (Shatoury, 1958;
B. amphitrite in a cement irrigation holding tank 64 kilometers south of the
mouth of the Nile Delta). On the basis of observational and experimental data,
it has been assumed that euryhaline barnacles, such as B. improvisus and B.
eburneus, can exist in fresh water as adults provided that higher salinities
prevail in the area during larval spawning and settlement. These outlying
populations are apparently unable to reproduce, and must rely, therefore, on
larval recruitment from breeding populations in higher salinities downstream.
The larvae can be transported upstream in the encroaching salt wedge during
the dry season, provided that this season (usually summer months) corresponds
to spawning season.

Laboratory studies of the salinity tolerances of stage I and II nauplii of
B. balanoides (Linnaeus), B. balanus (Linnaeus), and B. crenatus Bruguiére
were made by Barnes (1953), and Crisp and Costlow (1963) investigated the
effects of varying salinities on the developing eggs and early naupliar stages of
B. amphitrite amphitrite Darwin, B. eburneus, and Chelonibia patula Ranzani.
The results from these two studies agree rather closely. Cytolysis of eggs was
observed in all three species examined by Crisp and Costlow at salinities of 10
parts per thousand or less, regardless of temperature. Early stage nauplii of
both studies died in salinities of 5 parts per thousand or less after only slight
exposure, and could not withstand prolonged exposure to salinities below 10
parts per thousand. There was, however, some indication that larvae hatched
from embryos developed at lowered salinities were better adapted to brackish
conditions.

The survival ability of B. improvisus larvae in lowered salinities may be
greater than that of other estuarine barnacles. Mohammad (1962), in a field
and laboratory study of the distribution of barnacle larvae in Newport River,
North Carolina, indicated that B. improvisus larvae could tolerate salinity as
low as O parts per thousand, whereas the lower limit for those of B. eburneus
was about 8 parts per thousand. If larval salinity tolerance were the limiting
factor in the distribution of barnacles in estuaries, then B. improvisus should
be expected in fresh water. However, this is not the case under natural con-
ditions, suggesting that other factors are also limiting.

The only readily apparent physical difference between the two fresh water
localities cited here and natural estuarine habitats is direction of current flow.
Under natural conditions, larvae must counteract river current once outside
the encroaching salt wedge, and Crisp and Costlow (1963) have noted that

70 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH Ser.

Ficures 8-10. Opercular valves of Balanus eburneus Gould, CAS locality 42617, St.
Georges Island, Bermuda, height of scuta and tergum 7.5 mm. Ficures 11-15. Balanus
eburneus Gould, CAS locality 42618, Eniwetok, Marshall Islands. Figures 11 and 13,
scutum, height 4 mm.; figures 12 and 14, terga, height 4 mm.; figure 15, labrum, 100.

larval swimming ability is greatly decreased in lowered salinity. In pumped
irrigation systems, however, weak swimming larvae can be swept up from
river channels and carried into normally unattainable environments. Whether

Vor. XXXIX] ZULLO, BEACH, & CARLTON: BARNACLE RECORDS 71

ot s
' ‘ae &:
" :
“a :
19 —_—

Ficures 16-20. Creusia domingensis Des Moulins, CAS locality 42616, Bermuda.
Figures 16-17, terga, height 2.5 mm., and figures 18-19, scutum, height 2.5 mm.; figure
20, shell in Porites astreoides, greatest diameter of orifice 0.9 mm. FicurEs 21-22. Shell of
Hexacreusia durhami (Zullo), CAS locality 27229, Panama, greatest diameter 3.5 mm.

Figure 21, uncoated shell showing radii; figure 22, shell coated with ammonium chloride
to show external sculpture (apex of rostrum is damaged).

or not flow reversal is an adequate explanation for these unusual occurrences,
the possibility remains that barnacles may pose serious fouling problems in
irrigation systems.

Creusia domingensis Des Moulins, 1866.
(Figures 16-20.)

A coral-inhabiting barnacle of the genus Creusia Leach was also found by
Mrs. Gordon in Bermuda in 1967. The specimens are in two small heads of the

72 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

coral Porites astreoides (Lamarck). The larger coral fragment, measuring
about seven centimeters in greatest diameter, contains eleven barnacles, and the
smaller, measuring about four centimeters, harbors three individuals.

These barnacles were identified through the kindness of Mr. Arnold Ross
of the Natural History Museum, San Diego, who is presently preparing a
monographic study of coral-inhabiting balanids. He indicated that they
probably represent Creusia domingensis; a species originally described in P.
astreoides from Port-au-Prince, Haiti (Des Moulins, 1866), but which has
gone unnoticed since that time. Further examination of specimens of P.
astreoides in the collections of the California Academy of Sciences (CAS) and
of the University of California Museum of Paleontology, Berkeley (UCMP)
disclosed additional localities for C. domingensis from the vicinity of Biscayne
Bay, on the east coast of Florida (CAS locality no. 42265) and from the Dry
Tortugas, off southern Florida in the Gulf of Mexico (UCMP locality nos.
A-8146, A-8149, B-3717).

Previously, coral barnacles have been reported from the eastern Gulf of
Mexico and the West Indies in the Tropical Atlantic region. The discovery of
C. domingensis in the Bermudas and in southern Florida thus significantly
increases the known range both of this little known species and of coralophilous
balanids as a group.

Hexacreusia durhami (Zullo, 1961).

(Figures 21-22.

This six plated coral barnacle was originally described as a species of
Balanus Da Costa and placed in a new subgenus, Hexacreusia Zullo (Zullo,
1961). It was found in late Pliocene, Pleistocene, and Recent specimens of the
coral Porites californica Verrill from various localities in the Gulf of California,
and was later reported in extant specimens of the same coral from the Tres
Marias Islands, Mexico, just south of the Gulf of California by Ross (1962).

Although Hexacreusia is related to Balanus, and especially to the subgenus
Armatobalanus Hoek, it is more closely allied to the genera Creusia Leach and
Pyrgoma Gray, with which it shares a cup-formed basis, “creusioid’”’ rather
than “balanoid” opercular valves, and an obligate coralophilous habitat. For
these reasons, Hexacreusia is removed from the genus Balanus and raised to
generic rank.

It was noted earlier (Zullo, 1967) that certain specimens included by
Darwin (1854) in his original description of the Australasian species Balanus
allium were, in fact, Hexacreusia durhami. These barnacles were in a coral
from Hugh Cuming’s collection purported to be from Australia, but which
is most probably Porites californica from the Pacific Coast of tropical America.
As Cuming had spent some time in Central America, but had never traveled

Vor. XXXIX] ZULLO, BEACH, & CARLTON: BARNACLE RECORDS ae

north of the Gulf of Fonseca, Honduras, it was suggested that Darwin’s speci-
mens came from an area south of its present-known range.

Confirmation of the presence of Hexacreusia durhami in Central America
is based on the recent acquisition of two lots from Panama. A beachworn piece
of Porites californica collected by Dr. Leo G. Hertlein of the California
Academy of Sciences on December 22, 1931, at Bahia Honda, Veragua, Panama
(CAS-Geology locality no. 27229) contains three shells of Hexacreusia durhami
without opercular valves. The second lot, made available by Professor J. Wyatt
Durham of the University of California Museum of Paleontology (UCMP),
consists of about two dozen specimens in a large fragment of a Porites question-
ably identified as P. lobata (Dana). The coral was collected by Dr. Peter W.
Glynn of the Smithsonian Tropical Research Institute (Canal Zone, Panama)
from depths between 10 and 20 feet off the Las Secas Islands, Golfo de
Chiriqui, Panama (UCMP locality no. D-4138). These barnacles were alive
when taken and still retain opercular valves and bodies, although the latter
are unsuitable for dissection because of bleaching and drying during processing
of the coral.

Hexacreusia durhami is probably present in Porites throughout the Panamic
Province, but apparently is absent from the faunas of offshore Eastern Pacific
islands. Extensive examination of hermatypic corals from the Galapagos Archi-
pelago, Cocos Island (Costa Rica), and Clipperton Island has failed to yield
coral barnacles, although a second species, to be described at a later date, has
been discovered in ahermatypic corals from deeper water (90 meters) in the
Galapagos.

REFERENCES CITED

Barnes, H.
1953. The effect of lowered salinities on some barnacle nauplii. Journal of Animal
Ecology, vol. 22, no. 2, pp. 328-330.
Cartton, J. T., anD V. A. ZULLO
1969. Early records of the barnacle Balanus improvisus Darwin from the Pacific coast
of North America. Occasional Papers of the California Academy of Sciences,
no. 75, pp. 1-6.
Crisp, D. J., AND J. D. CostLow
1963. The tolerance of developing cirripede embryos to salinity and temperature.
Oikos, vol. 14, no. 1, pp. 22-34.
Darwin, C.
1854. A monograph on the sub-class Cirripedia, Balanidae, Verrucidae. Ray Society,
London, viii + 684 pp., 30 pls., 11 text figs.
Des Mouvutins, Cu.
1866. Liste des principaux fossiles recueilles par les membres de la Société a Cazeneuve
dans le calcaire de Bazas. Actes de la Société Linnéene de Bordeaux, tome
26, ser. 3; tome 6, pp. 293-344.
Hanna, G D.
1966. Introduced mollusks of Western North America. Occasional Papers of the
California Academy of Sciences, no. 48, pp. 1-108, figs. 1-85, pls. 1-4.

74 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Henry, Dora P.
1942. Studies on the sessile Cirripedia of the Pacific coast of North America. Univer-
sity of Washington Publications in Oceanography, vol. 4, no. 3, pp. 95-134,
pls. 14, figs. 1-5.
1958. Intertidal barnacles of Bermuda. Journal of Marine Research, vol. 17, pp.
215-234, pls. 1-9.
HERTLEIN, L. G.
1951. Invertebrate fossils and fossil localities in the San Francisco Bay Area. In
Geologic guidebook of the San Francisco Bay Counties. State of California
Division of Mines, Bulletin 154, pp. 187-192, figs. 1-2.
Martsu1, T., G. SHANE, AND W. NEWMAN
1964. On Balanus eburneus Gould (Cirripedia, Thoracica) in Hawaii. Crustaceana,
vol. 7, pt. 2, pp. 141-145, text fig. 1.
MonammMap, Murap-B. M.
1962. Larval distribution of three species of Balanomorpha in relation to some
chemico-physical factors. Proceedings of the First National Coastal and
Shallow Water Research Conference, October, 1961, pp. 360-361.
NEwMan, W. A.
1967. On physiology and behaviour of estuarine barnacles. Proceedings of Symposium
on Crustacea, Marine Biological Association of India, pt. 3, pp. 1038-1066,
figs. 1-9.
Piussry, H. A.
1907. Hawaiian Cirripedia. Bulletin of the Bureau of Fisheries, vol. 26, pp. 179-190,
pls. 4—S.
ProxopovicyH, N. P.
1968. Organic life in the Delta-Mendota Canal, Central Valley Project-California.
United States Department of the Interior, Bureau of Reclamation, Region 2,
Sacramento, California, x + 126 pp., 5 appendices.
Ross, A.
1962. Results of the Puritan-American Museum of Natural History Expedition to
Western Mexico, 15. The littoral balanomorph Cirripedia. American Museum
Novitates, no. 2084, pp. 1-44, figs. 1-24.
SHATOURY, H. H.
1958. A freshwater mutant of Balanus amphitrite. Nature, vol. 181, no. 4611, pp.
790-791, fig. 1.
UtTiInomiI, H.
1966. Recent immigration of two foreign barnacles into Japanese waters. Proceedings
of the Japanese Society of Systematic Zoology, no. 2, pp. 36-39, text figs. 1-2.
WELTNER, W.
1897. Verzeichnis der bisher beschriebenen recenten Cirripedienarten. Archiv fir
Naturgeschichte, Jahrgang 1897, Band 1, pp. 227-280.
ZWELEO IN ee Ne
1961. A new subgenus and species of coral-inhabiting barnacle from the Gulf of
California. The Veliger, vol. 4, no. 2, pp. 71-75, pl. 17, text figs. 1-2.
1967. On the identity of some specimens assigned by Darwin, 1854, to Balanus allium
Darwin (Cirripedia, Thoracica). Crustaceana, vol. 13, pt. 1, pp. 126-128.

PROCEEDINGS

OF THE

CALIFORNIA ACADEMY OF SCIENCES
FOURTH SERIES

Vol. XX XIX, No. 7, pp. 75-86, 4 figs; 3 tables. September 5, 1972

A KEY, BASED ON SCALES,
TO THE FAMILIES OF NATIVE
CALIFORNIA FRESHWATER FISHES

By

Richard W. Casteel

Department of Anthropology, University of California, Davis 95616

Asstract: Much interdisciplinary interest has been shown with regard to fish scales
in addition to their use in fisheries biology. To aid future workers, a key to the
scales of the native California freshwater fish families is presented along with photo-
micrographs of scales from each group.

INTRODUCTION

Fish scales have been used in fisheries biology and systematic ichthyology
for many years. Within fisheries studies, emphasis has been placed upon the use
of scales in age and growth studies (Cable, 1956; Cating, 1954; Chugunova,
1959; Cooper, 1951, 1952; Fry, 1943; Hile, 1936; Hogman, 1970; Jensen and
Wise, 1961; Miller, 1955; Phillips, 1948; Rush, 1952; Schuck, 1949; Taylor,
1916; Whitney and Carlander, 1956; Meehean, 1935). Various keys, based upon
the morphology of scales, have been published dealing with species identification
within families and with the identification of families comprising regional fish
faunas (Batts, 1964; Lagler, 1947; Koo, 1962).

Fish scales have been used in palaeontological work (David, 1944, 1946a,
1946b), sediment analysis (Lagler and Vallentyne, 1956; Pennington and Frost,
1961; Soutar and Isaacs, 1969), and archaeology (Follett, 1967a, 1967b; Hubbs
and Miller, 1948). Even within fisheries work, scales have been encountered
during analysis of the stomach contents of various fishes (Greenfield, Ross, and
Deckert, 1970; Kimsey, 1954). Based upon this evidently wide interdisciplinary
interest in and use of fish scales, it is felt that a scale-based key to the families
of native freshwater fishes of California would be a useful aid.

[75]
Marine Biological Laboratory
LIBRARY
SEP 15 1972
Woods Hole, Mass.

76

CALIFORNIA ACADEMY OF SCIENCES

TABLE 1. Species examined.

[Proc. 4TH SER.

Species

Source

Thaleichthys pacificus
Oncorhynchus tshawytscha
O. kisutch

Salmo gairdnerii

S. g. gairdnerii

S. g. stonei

S. g. gilberti

S. g. aquilarum

S. clarkii clarkii

S. c. henshawi

S. c. seleniris

S. aguabonita aguabonita

S. a. whitei

Salvelinus malma parkei
Prosopium williamsoni
Xyrauchen texanus
Catostomus luxatus

. platyrhynchus

. Santaanae

. rimiculus

. latipinnis

. occidentalis occidentalis

. 0. humboldtianus

. mniotiltus

. tahoensis

Rhinichthys osculus klamathensis
Mylopharodon conocephalus
Orthodon microlepidotus
Pogonichthys macrolepidotus
Lavinia exilicauda exilicauda
L. e. harengus
Ptychocheilus grandis

PD WY Py OS) B} &)

Hesperoleucas symmetricus symmetricus
H. s. subditus

H. s. venustus

H. navarroensis

— H.. parvipinnis

~ Gila bicolor bicolor
., -G~.b>, obesa

psy GeO. pectinifera

G. mohavensis
>» G. crassicauda
G. orcutti :

CAS*
VED
CAS, UCD
UCD
UCD
UCD
CAS
CAS
CAS
CAS
CAS
CAS
CAS
CAS
CAS
CAS
CAS
CAS
CAS
CAS
CAS
UCD
CAS
CAS
CAS
CAS
CAS, UCD
UCD
UCD
UCD
CAS
UCD
UCD
CAS
CAS, UCD
CAS
CAS
CAS
CAS, UCD
CAS
CAS
CAS
CAS

1 CAS = specimens from California Academy of Sciences, San Francisco.

2UCD = specimens from the author’s personal collection, presently at University of California, Davis.

VoL. XXXIX] CASTEEL: KEY TO CALIFORNIA FISHES 77

TABLE 1. (continued)

Species Source
G. elegans CAS
Richardsonius balteatus egregius CAS
Cyprinodon macularius californiensis CAS
C. nevadensis nevadensis CAS
C. n. calidae CAS
C. n. shoshone CAS
C. salinus CAS
Fundulus parvipinnis CAS
Mugil cephalis CAS
Archoplites interruptus UCD
Hysterocarpus traskii CASTUCED

For the purposes of this paper, native freshwater fishes will refer to those
fishes which occur exclusively in freshwater or spend a significant portion of
their life-cycle in freshwater and which occurred in California prior to the known
introduction of exotic species during and after the nineteenth century (Kimsey
and Fisk, 1960; Shapovalov, Dill, and Cordone, 1959; Walford, 1931). The
only exception has been the inclusion of the Mugilidae because of their impor-
tance in the Colorado River.

METHODS AND MATERIALS

The scales studied come from 54 species of native fishes (table 1) and
represent specimens collected by the author or by members of the California
Department of Fish and Game, and specimens from the California Academy of

TABLE 2. Scale sampling locations.

. Row anterior to dorsal fin.
. Right side, below dorsal fin, above lateral line.
. Left side, below dorsal fin, above lateral line.

. Right side, below dorsal fin, below lateral line.

Marine Biological Laboratory
LIBRARY
SEP 15 1972

Woods Hole, Mass.

. Left side, below dorsal fin, below lateral line.
. Right side, caudal penduncle, above lateral line.

. Left side, caudal penduncle, above lateral line.

ncluifoptel= Iles ipaleoiel i acial =

. Right side, caudal penduncle, below lateral line.
I. Left side, caudal penduncle, below lateral line.

J. Row posterior to dorsal fin.

78 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

TaBLeE 3. Scale characteristics of California
freshwater fish families.

Family Ctenoid Scales Cycloid Scales Scutes Neither

Petromyzonidae xX
Acipenseridae x

Osmeridae
Salmonidae

Catostomidae

me mM

Cyprinidae
Gasterosteidae x
Cyprinodontidae x x?

Mugilidae
Centrarchidae x

Embiotocidae xe

Cottidae x

a

3 Scales of these families, while being cycloid, should be oriented as shown for ctenoid scales in figure 1 b.

Sciences, San Francisco. Table III indicates the general scale characteristics
of the fishes in this study.

The fish were sampled for scales from ten different body locations (table
2) on the author’s specimens. For reasons of future use, the specimens from the
California Acadamy of Sciences could only be sampled from six locations on the
right-hand side of the fish. Wherever possible, samples were taken from several
individuals of different sizes within a species in order to allow for ontogenetic
variations. All scale samples were mounted in glycerine jelly on microscope
slides (Weesner, 1960) and examined under a dissecting microscope at between
ten and thirty magnifications.

DEFINITIONS

The terms used here to describe the surface features of scales are taken from
Lagler (1947, pp. 150-151) and are illustrated in figure 1.

Circuli — “Elevated markings on the outer surface; usually appearing as lines
which more or less follow the outline shape of the scale.”
Focus — “First part of scale to appear in growth; often central.”
Radi — “Grooves, usually more or less radiating from focus to one or more
margins.”
Primary Radu — “Radii that extend from focus to margin.”
Secondary Radi — “Radii that begin outward from, not at, focus.”

Ctenu — “Tooth-like structures on posterior portion of scale.”

Vor. XXXIX] CASTEEL: KEY TO CALIFORNIA FISHES 79

CIRCULI

LATERAL
FIELD

PRIMARY
RADII

SD
ae Dy iy POSTERIOR
ZB FIELD
Zz
ANTERIOR
FIELD
SECONDARY
RADII
LATERAL
FIELD
SECONDARY
RADII
Ay Bie S _/cTENII
oe Die } ve a =
ANTERIOR K nt Lae WEN 3 eS =
FIELD am MEE gow Soe “Ss FOCUS
aD ; “iy ae ae Ge
aS De Dw
p me Zee —- Soe
bs SS POSTERIOR
PRIMARY wy SIS ae
RADII i WSS. z
: ; - ma

: ——
Se
LATERAL CIRCULI

FIELD
Figure 1. Top. Cycloid Scale. Cyprinidae. Mvylopharodon conocephalus. UCD
5040 C. Bottom. Ctenoid Scale. Centrarchidae. Archoplites interruptus. UCD 5021 E.

80 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

yi
wy,

FicureE 2. a. Cyprinodontidae. Cyprinodon macularius californiensis. UCD 5063 G.
b. Centrarchidae. Archoplites interruptus. UCD 5021 E. c. Mugilidae. Mugil cephalus.
UCD 5056 I.

Fields — ‘Areas of the outer surface of the scale, either real as delimited by
angulation of the ridges (circuli) at levels of the four principal
corners or imaginary if the corners or configuration of the circuli are
wanting. Adjectives of direction applied to fields are based on their
positions when the scales are normally situated on the side of the

fish.”

Anterior Field — “Bounded by imaginary lines connecting the anterolateral
corners, or their equivalent points on scales which are
rounded (dorsal and ventral) with the focus.”

Posterior Field — “Bounded by imaginary lines connecting the posterolateral
corners (dorsal and ventral) with the focus.”

Lateral Fields — “Dorsal and ventral fields remaining after delimitation of

anterior and posterior ones.”

Vou. XXXIX] CASTEEL: KEY TO CALIFORNIA FISHES 81

Ficure 3. a. Salmonidae, Salmo gairdnerii gairdnerii. UCD 5002 D. b. Salmonidae.
Prosopium williamsoni. UCD 5016 C. c. Catostomidae. Catostomus occidentalis
occidentalis. UCD 5026 C.

SCALE-BASED KEY TO THE FAMILIES OF

CALIFORNIA FRESHWATER FISHES

The following is a key to the family level with the exception of Prosopium
williamsoni which is identified to species.

The Cyprinodontidae are characterized by both ctenoid and cycloid scales in
the same individual. Lagler (1947, pp. 156-157) classified the Cyprinodontidae
of the Great Lakes as having cycloid scales and the same appears true of the
genus Fundulus in California. However, the other members of this family may
also possess ctenoid scales (Lagler, Bardach, and Miller, 1962, p. 114). For this
reason the key identifies this family twice, once on the basis of ctenoid scales and
again on the basis of cycloid scales. This same case appears true of the Centrar-
chidae (Lagler, Bardach, and Miller, 1962, p. 114). Based upon data from my
collections, however, I find cycloid scales to occur only once in 32 samples and

82 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Ficure 4. a. Cyprinidae. Mylopharodon conocephalus. UCD 5040 C. b. Cyprinodon-
tidae. Cyprinodon macularius californiensis. UCD 5063 A. c. Embiotocidae. Hysterocarpus
traskiz. UCD 5099 C.

then only from restricted areas of the body in centrarchids (table 2, A). This is a
rather low frequency and if one considers the total number of scales over a fish’s
body, it appears that occurrence of cycloid scales in Archoplites interruptus will
be even more rare. I agree, therefore, that ‘These fishes may still be considered
as predominantly ctenoid in their squamation but the degree and extent of
development of the ctenii varies from place to place on the body” (Lagler,
Bardach, and Miller, 1962, p. 114). In the rare event that a cycloid scale from
this family should present itself in isolation it will key out as representing the
Embiotocidae instead of the Centrarchidae.

Figures 2, 3, and 4 illustrate each of the families or species separated by the
key. Each illustration is oriented with the anterior field to the observer’s left.
iva) Ctentiy present omisposterion) eile cl ee ee)

b)! (Ctenit “absent on ‘posterior field == ee eee
2. a) Ctenii numerous and evenly spaced

VoL. XXXIX] CASTEEL: KEY TO CALIFORNIA FISHES 83

b) Ctenii not numerous and irregularly spaced — eee Cyprinodontidae
(figure 2, a)

3. a) Radii converge toward focus ae Se ee RR PowresNGAaD Centrarchidae
(specifically Archoplites interruptus ; figure 2, b)

I eacimerouchiveunarallel) <2. 2 ee ee Mugilidae
(specifically Mugil cephalus; figure 2, c)

Pe mSCHeMmCVClOllmwithouts Tag) 2 a Be 5

Peal CmGy Cloldenwiliwna Cie sso. 88 ee ee ee 6
5. a) Scale with anterior and posterior fields only or without fields

coe neipene tee aE cee eee Osmeridae and Salmonidae

(figure 3, a)

b) Scale with four fields; focus centrally located Prosopium williamsoni

(figure 3, b)

6. a) Primary radii on both posterior and anterior fields = Catostomidae

(also includes the cyprinid genera Rhinichthys and Orthodon; figure 3, c)

b) Primary radii absent on either anterior or posterior field = 7

7. a) Primary radii present on anterior field, but absent on posterior field — == 8
b) Primary radii absent on anterior field, but present on posterior field = =

3 nc pec ME cade oo RS DA PE 8 Cyprinidae

(figure 4, a)

Sa ewartewes circuly im lateral than in anterior field —22-- = Cyprinodontidae

(figure 4, b)

b) Number of circuli in lateral field approximately equal to number in anterior
TT en eS a a Embiotocidae
(specifically Hysterocarpus traskiz; figure 4, c)

ACKNOWLEDGMENTS

I wish to express my thanks to Mr. W. I. Follett and Mrs. L. Dempster of the
California Academy of Sciences for their advice and cooperation and to Dr. W. G.
Kinzey, City University of New York, for his support and encouragement during
the initial phases of this work. My thanks also to Dr. R. W. Brocksen and
Messrs. L. Courtois, W. Wurtsbaugh, and H. W. Li, Department of Animal
Physiology, University of California, Davis; to Mr. Lyons, U. S. Department of
Reclamation; and to Messrs. A. Calhoun, L. Fisk, E. Armstrong, M. Coots, E. P.
Pister, R. Reavis, and J. Burns, California Department of Fish and Game, for
their aid in obtaining specimens. I also appreciate the helpful criticisms of the
manuscript given by Mr. W. I. Follett, California Academy of Sciences; Dr. D. L.
True, Department of Anthropology, University of California, Davis; and Dr.
J. D. Hopkirk, Department of Biology, Sonoma State College. Finally, my thanks
go to my wife for her help during all phases of this project.

LITERATURE CITED

Batts, Bitty S.
1964. Lepidology of the adult pleuronectiform fishes of Puget Sound, Washington.
Copeia, 1964, pp. 666-672.

84 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Caste, L. E.
1956. Validity of age determination from scales, and growth of marked Lake
Michigan lake trout. U. S. Fish and Wildlife Service, Fish Bulletin, no. 107,
pp. 1-59.
CatInG; J: P-
1954. Determining age of Atlantic shad from their scales. U. S. Fish and Wildlife
Service, Fish Bulletin, no. 85, pp. 187-199.
Cuucunova, N. I.
1959. Age and growth studies in fish: a systematic guide for ichthyologists. Academy
of Sciences of the U.S. S. R., 132 pp.
Cooper, E. L.
1951. Validation of the use of scales of brook trout, Salvelinus fontinalis, for age
determination. Copeia, 1951, pp. 141-148.
1952. Body-scale relationships of brook trout, Salvelinus fontinalis, in Michigan.
Copeia, 1952, pp. 1+.
Davin, Lore R.
1944. Use of fossil fish scales in micropalaeontology. Carnegie Institution of Washington
Publication, no. 551, pp. 25-43.
1946a. Some typical Upper Eogene fish scales from California. Carnegie Institution
of Washington Publication, no. 551, pp. 45-79.
1946b. Upper Cretaceous fish remains from the western border of the San Joaquin
valley, California. Carnegie Institution of Washington Publication, no.
551, pp. 81-112.
Fottett, W. I.
1967a. Fish remains from coprolites and midden deposits at Lovelock Cave, Churchill
County, Nevada. Annual Report of the University of California Archaeological
Survey, no. 70, pp. 94-115.
1967b. Fish remains from Salinas La Blanca, an archaeological site on the Pacific
coast of Guatemala. Smithsonian Contributions to Anthropology, vol. 3,
pp. 129-134.
Fry, F. E. J.
1943. A method for the calculation of the growth of fishes from scale measurements.
Ontario Fisheries Research Laboratory, Publication no. 61, pp. 5-18.
GREENFIELD, D. W., S. T. Ross, and G. D. DECKERT
1970. Some aspects of the life history of the Santa Ana Sucker, Catostomus (Pantosteus)
santaanae (Snyder). California Fish and Game, vol. 56, no. 3, pp. 166-179.
Hirer, R.
1936. Age and growth of the cisco, Leucichthys artedi (Le Sueur), in the lakes of the
northwestern highlands, Wisconsin. U. S. Fish and Wildlife Service, Fish
Bulletin, no. 19, pp. 211-317.
Hocman, W. J.
1970. Early scale development on the Great Lakes coregonids, Coregonus artedii
and C. kiyiz. In C. C. Lindsey and C. S. Woods (editors), Biology of Coregonid
Fishes. Winnipeg, pp. 429-435.
Hupss, C. L., and R. R. Mier
1948. Correlation between fish distribution and hydrographic history in the desert
basins of western United States. Jn The Great Basin, with emphasis on glacial
and postglacial times. Bulletin of the University of Utah, vol. 38, no. 20, pp.
17-166.

VoL. XXXIX] CASTEEL: KEY TO CALIFORNIA FISHES 85

Jensen, A. C., and J. P. WIsE
1961. Determining age of young haddock from their scales. U. S. Fish and Wildlife
Service, Fish Bulletin, vol. 61, pp. 439-450.
Kinsey, J. B.
1954. The life history of the tui chub, Siphateles bicolor (Girard), from Eagle Lake,
California. California Fish and Game, vol. 40, no. 4, pp. 395-410.
Kinsey, J. B., and L. O. Fisk
1960. Keys to the freshwater and anadromous fishes of California. California Fish
and Game, vol. 46, no. 4, pp. 453-479.
Koo, T.S. Y.
1962. Differential scale characters among species of Pacific salmon. Jn T. S. Y. Koo
(editor), Studies of Alaska Red Salmon. Seattle, pp. 123-135.
LAGLeR, K. F.
1947. Scale characters of the families of Great Lakes fishes. American Microscopical
Society Transactions, vol. 66, no. 2, pp. 149-171.
LActer, K. F., and O. R. VALLENTYNE
1956. Fish scales in a sediment core from Linsley Pond, Connecticut. Science, vol.
124, no. 3217, p. 368.
LAGLER, K. F., J. E. BArpAcH, and R. R. MILLER
1962. Ichthyology. New York, 545 pp.
MEEHEAN, O. L.
1935. The life history of the bluegill sunfish [Helioperca incisor (Cuvier and Valenci-
ennes) ] as determined from the scales. Proceedings of the Los Angeles Academy
of Science, vol. 2, pp. 139-145.
Muter, D. J.
1955. Studies relating to the validity of the scale method for age determination of the
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Game, Fish Bulletin, no. 101, pp. 6-34.
PENNINGTON, W., and W. E. Frost
1961. Fish vertebrae and scales in a sediment core from Estwaite Water (English
Lake District). Hydrobiologia, vol. 17, pp. 183-190.
Purtrps, J. B.
1948. Comparison of calculated fish lengths based on scales from different body
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1952. Observations of age and growth in the menhaden (Brevoortia tyrannus) as
determined by scale examination. Copeia, 1952, pp. 208-209.
ScHuck, H. A.
1949. Problems in calculating size of fish at various ages from proportional measurements
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1959. A revised checklist of the freshwater and anadromous fishes of California.
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1969. History of fish populations inferred from fish scales in anaerobic sediments
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1916. The structure and growth of the scales of the squeteague and the pigfish as

86 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

indicative of life history. U. S. Bureau of Fisheries, Bulletin, no. 34, pp.

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1960. General zoological microtechniques. Baltimore.
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1956. Interpretation of body-scale regression for computing body length of fish.
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PROCEEDINGS

OF THE

CALIFORNIA ACADEMY OF SCIENCES
FOURTH SERIES

Vol. XXXIX, No. 8, pp. 87-103. September 5, 1972

THE LYGAEIDAE OF THE GALAPAGOS ISLANDS
(Hemiptera:Heteroptera)

By
Peter D. Ashlock

University of Kansas, Lawrence

With the publication of this paper, reports on the Heteroptera of the Gala-
pagos International Scientific Project are nearly complete. Published papers
cover the Anthocoridae (Herring, 1966), the Tingidae (Drake and Froeschner,
1967), the Saldidae and Veliidae (Polhemus, 1968a, b), the Cydnidae ( Froeschner,
1968), the Nabidae (Kerzhner, 1968), the Miridae (Carvalho and Gagné, 1968),
and the emesine Reduviidae (Villiers, 1970). The families Pentatomidae, Corei-
dae, Rhopalidae, Stenocephalidae, Berytidae, Pyrrhocoridae, Reduviidae, Gerri-
dae, and Corixidae also are found in the Galapagos Islands, and members of the
expedition collected all but Pyrrhocoridae and Stenocephalidae. Hopefully, the
many new island records in these families will be summarized in another contri-
bution. The collections of only the Rhopalidae may still prove to contain new
species.

Before the expedition, Dr. Usinger and I had hopes of greatly increasing
the number of Lygaeidae recorded from the islands, especially in the subfamily
Orsillinae, a group well known for its ability to colonize islands. Only four species
of Lygaeidae, including two orsillines, had previously been recorded from the
Galapagos (Linsley and Usinger, 1966). Herein I am able to report only nine
species of lygaeids—five are orsillines—of which two, including one of the
orsillines, are possibly introduced by man. While more lygaeids may be found in
the archipelago, I seriously doubt that the known endemic fauna of the group
will double again. Because the lygaeid fauna of Hawaii contains forty percent
of the world orsillines, the sparse fauna of the Galapagos deserves some explana-
tion.

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88 CALIFORNIA ACADEMY OF SCIENCES [Proc. 47H Ser.

MacArthur and Wilson (1967) discuss the factors that determine the intro-
duction and extinction rates of island faunas as they develop the concept of an
island colonization rate. Further, they postulate that each island or archipelago
has a fairly fixed capacity for numbers of species. Among islands populated
from American sources, the Galapagos have a far better chance of receiving
introductions than do most other islands of the eastern Pacific, primarily because
they form an archipelago. The Hawaiian Islands, a larger archipelago, would
have received fewer introductions because of its great isolation.

Some examples will illustrate the point. The smallest orsilline faunas are
those of Guadelupe Island, 180 miles off Baja California, and San Felix, 550
miles west of Chile, each having a single undescribed endemic species (belonging
to Ortholomus and Nvysius, respectively). The Juan Fernandez Islands, two
islands about 450 miles off the coast of Chile, have two endemic orsillines, V ysius
baeckstroemi and Robinsonocoris tingitoides, the latter an endemic genus. The
Galapagos Islands have five species, resulting from four introductions, including
the endemic genus Darwinysius with two species. The Hawaiian fauna by
contrast is rich. Of the 92 species of Orsillinae, the tribe Metrargini | probably
of American origin (Ashlock, 1967) | contains 58 species (in six endemic genera)
resulting from but two introductions. The remaining Hawaiian species (in
Nysius and the endemic Nesomartis, a probable derivative of Nysius) are of
unknown origin. Clearly, the Galapagos have a much higher introduction rate
than Hawaii, but the larger, more hospitable Hawaiian Islands have a lower
extinction rate than the dryer and smaller Galapagos.

Thus, possible ecological situations on the Galapagos are nearly saturated
by naturally introduced orsillines. Hawaii’s isolation has allowed few intro-
ductions, but its low extinction rate, combined with the large number of ecological
situations, permitted explosive radiation.

Linsley and Usinger (1966) acknowledge those that made the G.I.S.P. expe-
dition possible. I must add my own thanks to the late R. L. Usinger, who made
my participation possible. Much of the work on this paper was done at the
B. P. Bishop Museum, Honolulu, with financial help from two NSF grants
(GB-3105 and GB—5860). Types of new species are deposited in the California
Academy of Sciences, San Francisco, and specimens upon which the study was
based will be found in that institution and in the Bishop Museum; the U. S.
National Museum; the California Insect Survey, Berkeley; and the collections
of J. A. Slater, G. G. E. Scudder, M. H. Sweet, and myself. All measurements are
in millimeters, and the year of collection is 1964 unless otherwise noted.

Kry To GALAPAGOS GENERA OF LYGAEIDAE

1. Suture between abdominal segments IV and V (second visible suture) curving anteriorly,
not reaching lateral margin of abdomen —____ Rhyparochrominae: Myodochini — 2
Suture between segments IV and V not curving anteriorly, clearly reaching lateral margin
Olgabdom ery es eee jee 3

VoL. XXXIX] ASHLOCK: GALAPAGOS LYGAEIDAE 89

2. Head constricted behind eye; eye removed from base of head by a distance greater than
NEI EOBITMeEO Teme UTM CV CMe eee eee ee ee Heraeus
Head not constricted behind eye; eye removed from base of head by a distance less than
[enc meO lame GE Cy Camere st Le ft. eee ee ee Pachybrachius

3. Scutellum bifid apically; clavus basally opaque, apically hyaline —— Cyminae:
Af 111 neem me pt ee Cymoninus
Scutellum not bifid apically; clavus not divided into distinct opaque and hyaline areas __
i @rsillinae yete 2 as a A

4. Costal margin of corium straight, exposing connexivum of abdomen laterally _
ai rr @rsillint 222 2 a ee ee Ori Volomus
Costal margin of corium straight at most for a distance less than length of scutellum,
then arcuately curved to apex; connexivum of abdomen completely covered — 5

5. Buccula not punctate; lateral margin of abdomen and corium without cross-striated
Giei@haliitieg, 2 ee ee INST 22-2 eee ee ee Nysius
Either buccula punctate or lateral margin of abdomen and corium with cross-striated
SUINIGIUIT ic, Se Metrarr gin .2 222 i ate eee 6

6. Buccula punctate, not tapering, ending abruptly at base of head; antenniferous tubercle
acutely produced; abdomen and corium without stridulatory structures ——
oper ee rrr rr a ee aS ee Darwinysius
Buccula impunctate, tapering to a low carina well before base of head; antenniferous
tubercle not acutely produced; abdomen and corium laterally with cross-striated stridu-
PROVES ENUGEU LC meee eeaeeee ae Se et ee eee Xyonysius

Genus Nysius Dallas
Nysius DALLAS, 1852, p. 551.

Although two species from the Galapagos were originally described in
the cosmopolitan genus \Vysius, both have since been placed in other orsilline
genera in the tribe Metrargini. Nysius (Ortholomus) naso Van Duzee is placed
in Xyonysius (Ashlock and Lattin, 1963) and Nysius (?) marginalis Dallas is
placed in Darwinysius (Ashlock, 1967). The following is the first true Nysius
to be reported from the Islands.

Nysius usitatus Ashlock, new species.
(Figures 1, 3.)

Head nearly flat between eyes, densely punctate, sparsely clothed with
appressed pale hairs, length 0.66, width 0.88, anteocular length 0.27, eye
length 0.29, eye width 0.19, interocular space 0.48; buccula widest anteriorly,
tapering posteriorly, and ending abruptly just before base of head; labium
extending to between hind coxae, first segment not reaching base of head but
just exceeding buccula, segment lengths from base 0.39, 0.39, 0.37, 0.29;
antenna with first segment exceeding clypeus by nearly half its length, segment
lengths from base 0.26, 0.60, 0.49, 0.49.

Pronotum moderately clothed with fine curved subappressed hairs, disk
densely punctate, distance between punctures from one-half to one diameter
of a puncture, sides nearly straight; length 0.59, width 1.02. Scutellum

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90 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

FicureE 1. Dorsal views of (left) Nysius usitatus and (right) Ortholomus usingeri.

with vestiture and punctation similar to those of pronotum; Y-shaped carina
appearing as a tumid pyramid; length 0.48, width 0.58.

Hemelytron exceeding abdomen, clavus and corium with surface almost
dull, moderately clothed with subappressed curved hairs, with additional
hairs projecting laterally along base of costal margin, clavus obscurely punctate
only at base of claval suture, costal margins subparallel and paralleling vein R+M
to level of apical two-thirds of scutellum, then gently becoming divergent
and arcuate to apex of corium, veins evident but not prominent; length of
claval commissure 0.34; length of corium 1.50; membrane normal with
veins evident but not distinct, basal length to level of corial apex 0.65, apical
length from corial apex 0.77.

Color. Light reddish brown, with broad stripe passing through each
ocellus, margins of clypeus, three spots at base of clypeus, ventor except
buccula, and spot below eye black. Buccula, labium, and antenna yellowish
brown, first segment of antenna a little darker. Pronotum light yellowish brown,

VoL. XXXIX | ASHLOCK: GALAPAGOS LYGAEIDAE 91

with partial indistinct median stripe, broad complete stripe behind each ocellus,
and narrow lateral stripe dark brown, callosities black. Scutellum black or
nearly so, apex white. Hemelytra light yellowish brown with indistinct brown
spots on veins and apical margin. Membrane hyaline, embrowned medially
except veins obscurely opaque white, ventor black except anterior and posterior
margins of propleura, acetabula, scent gland area, metapleural plate, and spots
laterally and apically on abdomen pale. Legs pale yellowish brown, with usual
brown spots on femur, tibia with a brown knee band.

Hototyre. 6, Santa Cruz Island, 1.5 mi. north of Academy Bay, 13 Febru-
ary, under Portulaca (P. D. Ashlock).

PaRATYPES. Santa Cruz Island, 9 ¢,10 2, same data as holotype; 1 ?, same
data but 11 February; 32 ¢,17 2, same data but 15 February; 3 ¢,5 2, same
data but 25 February; 3 2, Horneman Farm, 220 m., 2 April (D. Q. Cavagnaro).

Most specimens of this species were collected at the foot of the barranca
(cliff) on the new trail from Academy Bay, Santa Cruz Island, under Portulaca
and in company with Darwinysius marginalis. Since Nysius usitatus was
collected only near Academy Bay on Santa Cruz and not at all from the
Portulaca habitat on all other islands visited, it may be a recent introduction
to the Galapagos. Specimens fit Dallas’ (1852, p. 553) description of NV. nubilis
from Colombia fairly well, but the V. nubilis type is not extant. Dr. Usinger
searched in vain for it at the British Museum in 1964, and found (corre-
spondence) that the British Museum copy of the Dallas description has a note:

)

“type missing.” I have a short series of Nysius from Guayaquil, Ecuador,
and another from Palmira, Colombia, that also fit Dallas’ description, but the
spermathecae differ substantially from that of NM. usitatus (fig. 3). Since
there is only slight evidence that VV. usitatus is introduced, and its identity with
N. nubilis is open to question, I have described the Galapagos form as new. The
problem of the identity of VV. nubilis can best be solved by the judicious choice
of a neotype when adequate series of NV ysius are available from northwestern

South America.

Genus Ortholomus Stal

Nysius (Ortholomus) STAL, 1872, p. 43.
Ortholomus Stal, BAKER, 1906, p. 134.

Ortholomus usingeri Ashlock, new species.
(Figures 1, 3.)

Head flattened between eyes, obscurely punctate, densely covered with
flattened appressed hairs, vertex carina straight, eye prominent, slightly
raised above vertex; length 0.85, width 1.11, anteocular length 0.44, eye
length 0.22, eye width 0.29, interocular space 0.61; buccula widest anteriorly,

92 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Ficure 2. Dorsal views of (left) Darwinysius wenmanensis and (right) Xyonysius naso.

gradually narrowing posteriorly to mid eye level without abrupt change in
width; labium reaching between posterior coxae, first segment exceeding
buccula, not reaching base of head, segment lengths from base 0.46, 0.43, 0.46,
0.39; antenna with fine erect pubescence, first segment slightly exceeding
clypeus, segment lengths from base 0.29, 0.46, 0.44, 0.49.

Pronotum clothed with appressed flattened somewhat silky hairs, with
occasional short erect hairs, moderately punctate, distance between punctures
from one-half to one diameter of a puncture, sides straight, slightly swollen,
length 0.78, width 1.28. Scutellum with vestiture and punctation like those
of pronotum, Y-shaped carina swollen on upper arms, stem not prominent,
length 0.54, width 0.75.

Hemelytron slightly exceeding abdomen, clavus and corium moderately
clothed with short silky appressed hairs, with a few short erect hairs projecting

VoL. XXXIX] ASHLOCK: GALAPAGOS LYGAEIDAE 93

laterally along base of costal margin; veins evident but not prominent; length
of claval commissure 0.49; length of corium 1.80; membrane irregularly
wrinkled transversely; veins distinct, basal length to level of corial apex 0.77,
apical length from corial apex 0.78.

Abdomen with connexival spiracles raised, very prominent.

Color. Head very dark brown, clypeus, median stripe on posterior half
of vertex, buccula, labium, and antenna paler except antennal segment IV very
dark brown. Pronotum and scutellum reddish brown, callosities and punctures
a little darker. Hemelytron pale yellowish brown, darkened apically on clavus
and corium and on obscure spots on veins; vestiture nearly white, silky.
Connexival segments pale, darker around pale spiracles. Legs yellowish brown,
femur with obscure, slightly darker spots. Ventor reddish brown, acetabula and
scent gland paler.

Hototype. ¢, Santa Cruz Island, grassland, 1,800 ft., north of Academy Bay,
20 February, on Hypericum pratense (P. D. Ashlock).

PARATYPES. Santa Cruz Island, 6 ¢, 12 2,4 nymphs, same data as holotype;
feo, ) 2, same data but 2,100 it.; 1 ¢, 1 9%, Bella Vista, 26 February (R. L.
Usinger); 15 ¢, 15 2, Table Mountain, 440 m., 16 April (D. Q. Cavagnero).
Floriana Island, 1 ¢, 1 2, 15 February, Verbena (R. L. Usinger); 57 6, 48 2,
Whitmer’s Farm, 15 February, Verbena (R. L. Usinger); 1 ¢,2 2, same data but
Cordia species; 1 ¢, 1 2, moist forest 200 m. above Black Beach, 15 February,
Cordia tree with yellow flowers (R. L. Usinger); 7 ¢, 10 2, 18 February
(R. L. Usinger). San Cristobal Island, 3 ¢,4 2, Progresso, 23 February, Verbena
(R. L. Usinger).

Ortholomus usingeri is less elongate than most of the described species
in the genus, and the connexivum is more broadly exposed, being approached
only by O. gibbus (Berg) in this respect. The spiracles are more prominent
and enlarged than in any other species. All series of the species are variable
in color, ranging from the pale form illustrated (fig. 1), to specimens that are
highly maculated on the corium, clavus, and membrane. Some specimens have a
very dark, contrasting claval apex. None of these variations are correlated
with island of origin or with host plant. The new species of Ortholomus from
Guadelupe Island (see Introduction) may be very closely related to O. usingeri.

It is with devoted appreciation that I dedicate this Galapagos orsilline to
R. L. Usinger, who was the leading authority in the Orsillinae, and whose
leadership of the G. I. S. P. expedition he considered the high point of his life.

Genus Darwinysius Ashlock
Darwinysius ASHLOCK, 1967, p. 42.
Darwinysius is the only endemic genus of Lygaeidae—and one of the very
few in the Heteroptera—found in the Galapagos. It is closely related to

94 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

D. wenmanensis

N. usitatus

O. usingeri

X. naso

C. notabilis

Ficure 3. Spermathecae of the named species.

Robinsonocoris in the Juan Fernandez Islands, differing principally in features
correlated with the flightlessness of Robinsonocoris. At the time of the original
description of Darwinysius for Nysius (2?) marginalis Dallas, it was mentioned
that a second species had been found on the Galapagos.

Key TO GALAPAGOS SPECIES OF DARWINYSIUS

Larger species, male more than 3.2 long, female more than 3.8 long; clothed above with
short, appressed hairs; vertex of head raised above level of top of eye; Wolf Island —
Bren oN Ele Te eek D. wenmanensis Ashlock, new species

VoL, XXXIX] ASHLOCK: GALAPAGOS LYGAEIDAE 95

Smaller species, male less than 3.1 long, female less than 3.5 long; clothed above with mixed
erect and appressed hairs; vertex of head not raised above level of top of eye; main islands __
_____... D. marginalis (Dallas)

Darwinysius marginalis (Dallas).

Nysius (2?) marginalis DALLAS, 1852, p. 556.

Cymus galapagensis STAL, 1959, p. 252 (synonymy by Butler, 1877).
Darwinysius marginalis (Dallas), ASHLOCK, 1967, p. 42.

In the summer of 1964, Dr. R. L. Usinger checked the types of Cymus
galapagensis Stal and Nysius (?) marginalis Dallas at my request. In Stockholm
he confirmed Butler’s (1877) synonymy of Stal’s species with that of Dallas, but
in London, he found Dallas’ series of eight specimens to contain two species: four
of the eight belonged to Xyonysius naso (Van Duzee). Since Dallas did not
select a holotype, and no lectotype had since been chosen, Usinger selected the
best specimen of the remaining four as a lectotype. He wrote that the lectotype
‘Gs rather broad and well marked and has no antennae.’ The specimen is
numbered 77—2 (as were the X. aso) and is from Charles Island (Floreana).
The remaining specimens were from James Island (Santiago) and “were very
poor and not suitable to be made a lectotype.” The Charles Island specimen
(no. 77-2) is here formally selected as the lectotype of Nysius (?) marginalis
Dallas.

Published records of Darwinysius marginalis are from three islands: Floreana,
Santiago, and Daphne Major. Members of the G. I. S. P. expedition collected
well over 300 specimens and can add five islands to the list: Isabella, Fernan-
dina, Rabida, Pinzon, and Santa Cruz. Almost all of my specimens were collected
under Portulaca, where they were feeding on fallen seeds of the plant. Dr.
Usinger collected 55 specimens on Euphorbia viminea at Black Beach, Floreana.

Darwinysius marginalis is generally similar to D. wenmanensis; differen-
tiating characters can be found in the key to species. A dorsal view of D.
marginalis, a side view of the head, and drawings of the aedeagus and sperma-
theca are given in Ashlock (1967).

Darwinysius wenmanensis Ashlock, new species.
(Figures 2, 3.)

Head elevated above eye about one-third height of eye, sparsely clothed
with flattened appressed hairs, length 0.71, width 0.80, anteocular length 0.46,
eye length 0.20, eye width 0.15, interocular space 0.48, buccula widest anteriorly,
narrow, but not tapering to base of head, slightly projecting past posterior
point of attachment, not reaching base of head, densely punctate; labium just
reaching posterior coxae, first segment not reaching posterior end of buccula,
segment lengths from base 0.46, 0.43, 0.44, 0.32; antenna with flattened sub-
appressed hairs on first segment, semierect fine hairs on fourth segment, second

96 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

and third segments intermediate, first segment exceeding clypeus by nearly half
its length; segment lengths from base 0.39, 0.56, 0.48, 0.41.

Pronotum with flattened appressed hairs, deeply and densely punctate, cica-
trices depressed, area anterior to cicatrices somewhat inflated; length 0.78, width
1.19. Scutellum with vestiture and punctation similar to those of pronotum,
with a strongly elevated Y-shaped carina, length 0.51, width 0.68.

Hemelytron exceeding abdomen, clavus and corium moderately dotted with
appressed hairs, without erect hairs at base of lateral margin, surface overall
ridged, punctate on both sides of claval suture and outer edge of vein R+M for
entire length, all veins prominent and elevated, length of claval commissure 0.54,
length of corium 2.04, apex attaining abdominal segment VII; membrane with
surface irregularly ridged transversely, veins prominent, basal length to level of
corial apex 0.90, apical length from corial apex 0.54.

Color. Head, labium, antenna very dark brown, apices of first to third
antennal segments and lower edge of buccula pale. Pronotum, scutelum, and
hemelytron mottled dark and yellowish brown; pronotum with a patch of white
flattened hairs behind cicatrix, corium with lateral margin alternately light
and dark, apical margin beyond intersection of vein Cu very dark brown,
membrane including veins mottled brown and transparent. Ventor dark brown
except acetabula, posterior lobe of propleura, mesothoracic scent gland, and
lateral posterior angle of metapleuron pale, legs with femur very dark brown,
apex pale, tibia and tarsi pale with dark band dorsally and subapically on tibia.

HototyPe. 6, Wolf Island (Wenman). 31 January, under Portulaca (P. D.
Ashlock).

ParATyPeEs. Wolf Island, 15 6,15 2, same data as holotype; 1 4, 24 Septem-
ber 1906 (F. X. Williams).

Darwinysius wenmanensis is abundantly distinct from the type species of
the genus as indicated by characters in the key. Both species are variable in
color pattern, ranging from nearly immaculate on the clavus and corium to
quite dark with few pale maculations, but in general D. wenmanensis tends to
be darker. Such features of both species as the spotted costal margin, the acute
antenniferous tubercles, the general body shape, and, in D. wenmanensis, the
raised vertex of the head, make both look very much like miniature Nesoclimacias
(Hawaiian Islands). This similarity was noted earlier (Ashlock, 1967, p. 35);
however, I now think these similarities—of Robinsonocoris and Darwinysius
with the Hawaiian Metrarga, Nesoclimacias, and Nesocryptias—represent
parallelisms rather than a monophyletic origin.

Specimens of D. wenmanensis from Wenman Island were the first members
of the genus I collected, and it was on Wenman that the “under Portulaca”
habitat of Darwinysius was discovered. Unfortunately, it was then too late to

VoL. XXXIX |] ASHLOCK: GALAPAGOS LYGAEIDAE 97

search the Portulaca on Darwin Island (Culpepper), the most remote of the
archipelago.

Genus Xyonysius Ashlock and Lattin
X yonysius ASWLOcCK and LATTIN, 1963, p. 702.

Xyonysius naso (Van Duzee).

(Figures 2, 3.)

Nysius (Ortholomus) naso VAN DUZEE, 1933, p. 27.

Ortholomus naso (Van Duzee), BARBER, 1934, p. 285.

Nystus naso Van Duzee, USINGER, 1941, p. 131.

Xyonystus naso (Van Duzee), ASHLOCK and LATTIN, 1963, p. 702.

Xyonysius is a New World genus of metragine Orsillinae whose species
are found from Canada to Argentina and Chile. The endemic Galapagos species
X. naso is one of the more variable ones, ranging from forms with immaculate
hemelytra to forms with three irregular dark brown longitudinal stripes on
the corium, the middle one continuing onto the membrane, and dark brown
claval apices. Figure 1 shows a specimen with intermediate coloration. This
variation was found in all series of the species collected, and does not appear
to be correlated with island or host plant. Unlike X. californicus (Stal), a
wide-spread North American species found on many composites, X. naso
seems to be confined to species of the endemic composite genus Scalesia. The
species was described from a single specimen taken on Floreana (Charles Island).
Dr. Usinger and I collected 59 specimens on the following islands and host plants:
Santa Cruz (S. affinis, S. helleri), Fernandina (Scalesia species), Barrington
(S. helleri), Isabella (S. affinis, S. gummifera), and Floreana (S. affinis).
Nowhere was the species abundant.

Genus Cymoninus Breddin

Cymoninus BREDDIN, 1907, p. 38.

Cymoninus notabilis (Distant).

(Figures 3, 4.)

Ninus notabilis D1isTaNT, 1882, p. 191.

Cymoninus notabilis (Distant), VAN DUZEE, 1917, p. 163.

This species is here reported from the Galapagos for the first time. Wide-
spread, it ranges from the southern United States through Central America and
the Antilles south to Argentina and Brazil. I collected 55 specimens on Santa
Cruz Island on the trail north of Academy Bay and Bella Vista at about 1,300
feet on a sedge identified by Ira Wiggins as Cyperus confertus Swartz.
D. Q. Cavagnaro also collected a few specimens on Santa Cruz at the Horneman
Ranch, just north of Bella Vista. This species may be a recent introduction to
the Galapagos. I collected two specimens in the harbor at Guayaquil, Ecuador,
at light, on board ship.

98 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH Ser.

Genus Pachybrachius Hahn

Pachybrachius HAHN, 1826, p. 18.
Orthaea DALLAS, 1852, p. 580.

Key To GALAPAGOS SPECIES OF PACHYBRACHIUS

Smaller species, less than 4 long; usually brachypterous; corium with long erect hairs and an
Oval whitejspot neaainner angle 2 ee P. nesovinctus Ashlock, new species
Larger species, more than 4 long; never brachypterous, fully winged; corium with short
appressed hairs and without an oval white spot near inner angle —

Pachybrachius insularis (Barber).

(Figure 5.)

Orthaea insularis BARBER, 1925, p. 246.

Pachybrachius insularis (Barber), SLATER, 1964, p. 1127.

Barber’s Galapagos Pachybrachius has been recorded from Isabella, Santiago,
Baltra, and Santa Cruz Islands. Members of the expedition collected the species
from all of these but Isabella and Baltra, but also collected it on Fernandina
and Floreana. Most of the material was collected from lights, though occasional
specimens were collected from the forest floor. The two specimens from Floreana,
both females (Kuschel and Usinger, collectors) were unusual, since both had
the labium extending onto the abdomen. Barber (1925) described the labium
as “reaching just past intermediate coxae.’” However, labial length in a
36-specimen sample varied as follows: past front but not to mid coxae, 5 4,
O 2; between mid coxae, 14 6, 7 2; past mid but not to hind coxae, 3¢,
4 2: between hind coxae, 0 6, 1 2; behind hind coxae onto abdomen, 0 ¢,
2 2. This character is clearly variable, and females tend to have the longer
labia. Although the two females with the longest labia came from the same
island, there was no other correlation of labial length with island of origin.

Pachybrachius nesovinctus Ashlock, new species.
(Figures 4, 5.)

Head slightly elevated between eyes, densely but shallowly punctate, sparsely
dotted with inconspicuous subappressed and long erect hairs, length 0.71,
width 0.75, anteocular length 0.37, eye length 0.17, interocular space 0.44;
buccula high, short, bucculae joined as a single low carina at level of antenniferous
tubercle; labium not quite reaching mid coxae, first segment not reaching base
of head, segment lengths from base 0.44, 0.44, 0.27, 0.29; antenna with first
segment exceeding clypeus by nearly half its length, segment lengths from base
0.34, 0.63, 0.49, 0.94 (fourth segment drawn too thick in figure 3).

Pronotum with sparse short subappressed hairs and long erect fine hairs, with
a single transverse row of widely spaced shallow small punctures on collar,

VoL. XXXIX] ASHLOCK: GALAPAGOS LYGAEIDAE 99

fi

f

ye Kj
ih

Cymoninus notabilis Pachybrachius nesovinctus

Ficure 4. Dorsal views of the named species, and anterior view of fore femur of the
holotype of P. nesovinctus.

and similar punctures scattered over posterior lobe, length 0.83, collar length
0.09, width 0.56, anterior lobe length 0.29, width 0.88; divisions between collar
and two lobes very distinct, anterior lobe globular. Scutellum with vestiture
and punctation like those of hind lobe of pronotum; length 0.46, width 0.46.
Brachypterous, hemelytron reaching midway onto abdominal segment VI,
clavus and corium with surface subshining, vestiture as in pronotum and
scutellum, clavus with three linear rows of punctures and a confused fourth
row between medial and scutellar rows, corium with two linear rows of punctures

100 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

P. nesovinetus

le,

insularis

Ficure 5. Spermathecae of the named species.

VoL. XXXIX] ASHLOCK: GALAPAGOS LYGAEIDAE 101

paralleling claval suture, randomly punctate laterally, with a few punctures
posterior to level of apex of clavus; length of claval commissure 0.31, length of
corium 1.36; membrane dull, veins not visible, basal length to level of corial
apex 0.63, apical length from corial apex 0.07.

Fore femur armed beneath with spines in two ranks as figured (fig. 4); fore
tibia unarmed, slightly curved.

Color. Head black, clypeus and juga dark reddish brown, antenna, buccula,
and labium light yellowish brown. Pronotum black, collar anteriorly pale,
posterior lobe reddish brown with dark median line. Hemelytron with clavus and
corium light yellowish brown, punctures reddish brown, corium with white spot
in inner angle and the black apical margin of P. vinctus group. Venter dark with
anterior and posterior margins of propleuron, acetabula, scent gland auricle,
and metapleural plate pale. Abdomen ventrally and dorsally dark reddish
brown. Legs light yellowish brown, fore femur dark brown except apically.

Hototyree. ¢, Santa Cruz Island, grassland, 2,100 ft., north of Academy
Bay, 20 February (P. D. Ashlock).

PaRATYPES. Santa Cruz Island, 10 ¢, 12 2, same data as holotype; 1
(macropterous), Table Mountain, 440 m., 16 April (D. Q. Cavagnaro).
Fernandina Island, 1 2, southwest side, 1,000 ft., 4 February (P. D. Ashlock).

This description adds another species to the confusing Pachybrachius vinctus
complex, which badly needs revision. The new species is distinct in having
long erect hairs on the dorsum. The New World P. vinctus (Say) and the
closely related Pacific P. pacificus (Stal) both have small, inconspicuous
appressed hairs on the dorsum. Pachybrachius nesovinctus is usually found
in the brachypterous state (only one macropterous specimen has been collected),
as are many populations of P. pacificus from the western and central Pacific.
Brachypterous forms are less common in the Western Hemisphere species P. vinc-
tus. The Galapagos species show more brown on the hemelytron than do speci-
mens of P. vinctus from the North and South American mainlands.

Genus Heraeus Stal
Heraeus STAL, 1862, p. 315.

Heraeus pacificus Barber.
(Figure 5.)

Heraeus pacificus BARBER, 1925, p. 21.

Barber described this species from Santiago, and it has previously been
recorded only from this island. Members of the expedition collected it in large
numbers from Santa Cruz, and a single specimen was collected from Floreana.
Many of the specimens were collected at light between Academy Bay and the
Horneman Ranch on Santa Cruz, and I collected a large series in the so-called
grassland area of the island at 1,800 feet under Jaegeria hirta.

102 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH Serr.

LITERATURE CITED

ASHLOCK, P. D.

1967. A generic classification of the Orsillinae of the world (Hemiptera—Heteroptera:
Lygaeidae). University of California Publications in Entomology, vol. 48;
pp. 1-82.

AsHLock, P. D., and J. D. Lattin.

1963. Stridulatory mechanisms in the Lygaeidae, with a new American genus of
Orsillinae (Hemiptera: Heteroptera). Annals of the Entomological Society of
America, vol. 56, pp. 693-703.

Baker, C. F.

1906. Notes on the Nysius and Ortholomus of America. Invertebrata Pacifica,

vol. 1, pp. 133-140.
Barser, H. G.

1925. Hemiptera-Heteroptera from the Williams Galapagos expedition. Zoologica, vol.
5, pp. 241-254.

1934. The Norwegian zoological expedition to the Galapagos Islands 1925, conducted by
Alf Wollebaek. XI. Hemiptera—Heteroptera. Nyt Magazin for Naturvidens
kaberne, vol. 74, pp. 281-289.

BreEpDDIN, G.

1907. Berytiden and Myodochiden von Ceylon aus der Sammelausbeute von Dr. W.

Horn (Rhynch. het). Deutsche Entomologische Zeitschrift 1907, pp. 34-47.
Butter, A. G.

1877. Account of the zoological collection made during the visit of H.M.S. “Petrel’”
to the Galapagos Islands. X. Lepidoptera, Orthoptera, Hemiptera. Proceedings
of the Zoological Society, London 1877, pp. 86-91.

CarRVALHO, J. C. M., and W. C. Gacne.

1968. Miridae of the Galapagos Islands (Heteroptera). Proceedings of the California

Academy of Sciences, vol. 37, no. 7, pp. 147-219.
DALLAs, W. S.

1852. List of the specimens of hemipterous insects in the collection of the British

Museum, London, Pts. I and II. 592 pp.
Distant, W. L.

1880-1893. Insecta. Rhynchota. Hemiptera-Heteroptera. Vol. I. Biologia Centrali-

Americana. xx + 462 pp. 36 pls.
DRAKE, C. J., and R. C. FROESCHNER.

1967. Lace bugs of the Galapagos Archipelago (Hemiptera: Tingidae). Proceedings of

the Entomological Society of Washington, vol. 69, no. 1, pp. 82-91.
FROESCHNER, R. C.

1968. Burrower bugs from the Galapagos Islands collected by the 1964 expedition of the
Galapagos Scientific Project (Hemiptera: Cydnidae). Proceedings of the
Entomological Society of Washington, vol. 70, no. 2, p. 192.

Hann, C. W.
1826. Icones ad monographium Cimicum. Nurnberg, Lechner.
HERRING, J. L.

1966. The Anthocoridae of the Galapagos and Cocos islands (Hemiptera). Proceedings

of the Entomological Society of Washington, vol. 68, no. 2, pp. 127-130.
KERZHNER, I. M.

1968. Insects of the Galapagos Islands (Heteroptera, Nabidae). Proceedings of the

California Academy of Sciences, vol. 36, no. 4, pp. 85-91.

VoL. XXXIX] ASHLOCK: GALAPAGOS LYGAEIDAE 103

LINSLEY, E. G., and R. L. USINGER.
1966. Insects of the Galapagos Islands. Proceedings of the California Academy of
Sciences, vol. 33, no. 7, pp. 113-196.
MacArruur, R. H., and E. O. Wison.
1967. The theory of island biogeography. Princeton, New Jersey, Princeton University
Press. xi + 203 pp.
PoLHEMUS, J. T.
1968a. A report on the Saldidae collected by the Galapagos International Scientific
Project 1964 (Hemiptera). Proceedings of the Entomological Society of
Washington, vol. 70, no. 1, pp. 21-24.
1968b. A new Microvelia from the Galapagos (Hemiptera: Veliidae). Proceedings
of the Entomological Society of Washington, vol. 70, no. 2, pp. 129-132.

SLATER, J. A.
1964. A catalogue of the Lygaeidae of the world. Storrs, University of Connecticut.
1,668 pp.
Span, (C-

1859. Hemiptera species novas descriptsit. Konglika Svenska Fregattens Eugenies resa
omkring jorden. III (Zoologi, Insecter) 1859, pp. 219-298.
1862. Hemiptera Mexicana enumeravit speciesque novas_ descripsit. Stettiner
Entomologische Zeitung, vol. 23, pp. 81-118; 273-281; 289-325; 437-462.
1872. Genera Lygaeidarum Europae disposuit. Ofversigt af K. Vetenskaps-akademiens
for handlingar, Stockholm vol. 29, no. 7, pp. 37-62.
Usincer, R. L.
1941. The present status and synonymy of some orsilline species (Hemiptera, Lygaeidae).
Bulletin of the Brooklyn Entomological Society, vol. 36, pp. 129-132.
Van DuzeE, E. P.
1917. Catalogue of the Hemiptera of America north of Mexico. University of
California Publications in Entomology, vol. 2, i-xiv, 1-902.
1933. The Templeton Crocker expedition of the California Academy of Sciences,
1934. No. 4. Characters of twenty-four new species of Hemiptera from the
Galapagos Islands and the coast islands of Central America and Mexico.
Proceedings of the California Academy of Sciences, vol. 21, no. 4, pp. 25-40.
Viriters, A.
1970. Les Emesines des Iles Galapagos (Hemiptera, Reduviidae). Jn: Mission
zoologique belge aus iles Galapagos et en Ecuador (N. et J. Leleup, 1964-1965),
vol. 2, pp. 227-237.

8 TTP | ’ Leg) 7 A
i he ais

PROCEEDINGS

OF THE

CALIFORNIA ACADEMY OF SCIENCES
FOURTH SERIES

Vol. XXXIX, No. 9, pp. 105-110. September 5, 1972

A NEW LIZARD OF THE GENUS EMOIA
(SCINCIDAE) FROM THE MARIANAS ISLANDS

By
Walter C. Brown

Menlo College, Menlo Park, California, and Division of Systematic Biology,
Stanford University. Research Associate, Department of Herpetology,
California Academy of Sciences.

and

Marjorie V. C. Falanruw

University of Guam, Territory of Guam.

INTRODUCTION

Brown (1956) recognized eight species of the Papuan-Oriental genus of
lizards, Emoia, as occurring in Micronesia. Two were noted as apparently
undescribed, of which one, a moderate-sized species, was represented by only
four specimens from the Marianas Islands in the northwestern part of Micro-
nesia. One of us (the junior author) has recently obtained a large series of
the species from the same island group, and this larger sample permits a more
adequate diagnosis of the species. It apparently has its closest affinities
with Emoia boettgeri (Sternfeld) from central and eastern Micronesia and with
Emoia arnoensis Brown and Marshall from the Marshall Islands in eastern
Micronesia. Emoia atrocostata (Lesson), a widely distributed species-complex
in the western Pacific basin; Emoia flavigularis Schmidt from the Solomon
Islands to the south; and the very large Emoia nigra Jacquinot and Guichenot,
the range of which extends from the Solomon to the Tonga Islands in the southern
Island chain, are also representatives of the same evolutionary group.

Measurements were made with a caliper to the nearest 0.1 mm. Middorsal
scale rows were counted between the parietals and a point opposite the vent;

[105]

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Vor. XXXIX] BROWN & FALANRUW: A NEW SPECIES OF EMOIA 107

midbody scale rows at a point approximately equidistant between the fore and
hind limbs; fourth-toe lamellae from that proximal rounded lamella which is
at least two-thirds the breadth of the toe to the most distal ventral scale inclusive.

Emoia slevini Brown and Falanruw, new species.

Hototyre. United States National Museum no. 192781, Cocos Island (a
small island off the southern end of Guam), Marianas Islands, collected by
M. V. C. Falanruw.

PaRATYPES. United States National Museum nos. 122645-122646, Rota
Island; 122470, Ritidain Point, Guam Island, collected by L. P. McElroy;
128028, Mt. Lasso, Tinian Islands, collected by H. K. Townes, Jr.; University
of Guam nos. 547-552, 1305-1306, 1393-1395, 1448, 1450-1451; California
Academy of Sciences nos. 129138—-129143; Museum of Comparative Zoology,
Harvard University no. 128164; Field Museum of Natural History no. 171832;
and British Museum of Natural History no. 1971.1027, from the same locality
as the holotype.

Dracnosis. An Emoia species of moderately large size, 58-85 mm. in
snout-vent length for 18 mature specimens; length of hind limb less than 50
percent of the snout-vent length; simply rounded lamellae on under surface of
digits, numbering 30-37 beneath the longest toe; 34-38 midbody scale rows and
61—74 dorsal scale rows between parietals and base of tail (for a sample of 27
specimens); prefrontals separated by the frontal; frontoparietals distinct;
interparietal moderate in size; ground color (in life) of dorsum and upper
lateral surfaces light to dark brown with variable scattered dark and light
flecks.

DeEscriPpTION. An Emoia species of moderate size; snout-vent length of 6
mature females 63-75 mm., of 12 males 69-84 mm.; snout round-pointed, of
moderate length (36-44 percent of head length); supranasals much broader
anteriorly than posteriorly, in contact with the anterior loreal which is shorter
and broader than the posterior; prefrontals not in contact (separated by the
frontal which is in contact with the rostral); length of the frontal about equal
to that of the fused frontoparietals; interparietal of moderate size, length
about equal to breadth at the base to nearly one and one-half times as great;
4 or 5 supralabials anterior to the enlarged one beneath orbit; 4 supraoculars plus
a small one posteriorly; a single pair of large nuchals (fig. 1), dorsal scales
smooth; midbody scale rows 34-38 for twenty-eight specimens; transverse
rows along the middorsal line from parietals to base of tail 61-74; number of
rows across the nape from ear to ear 12; 30-37 smooth, rounded lamellae

<
Ficure 1. Emoia slevini. a. Lateral view of head of paratype; b. Dorsal view of head of
paratype, CAS.

Biological Laboratory

LIBRARY
SEP 15 1972

Marine

[Proc. 47TH Ser.

CALIFORNIA ACADEMY OF SCIENCES

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Vor. XXXIX] BROWN & FALANRUW: A NEW SPECIES OF EMOIA 109

beneath the fourth toe for 27 specimens; limbs rather well developed, but hind
limb less than 50 percent of the snout-vent length (table 1) and usually slightly
less than distance from axilla to groin.

Cotor. The ground color dorsally (in life) is irridescent medium brown
to dark brown with some darker flecks and occasionally light flecks, limbs,
especially the hind limbs, often lighter with more numerous flecks; venter
whitish to gray or cream anteriorly, more yellow posteriorly, sometimes orange
about the vent; in recently preserved specimens the ground color is brown or
somewhat reddish brown with scattered darker markings, in some specimens
forming a vague pattern of narrow, broken, transverse bands. The yellow and
orange of the venter fade rapidly in preservative.

EtymoLocy. The species is named for Mr. Joseph R. Slevin, former Curator
of Amphibians and Reptiles at the California Academy of Sciences.

ComPARISONS. Emoia slevini is probably most closely related to Emoia
boettgert which occurs in the Caroline Islands. The two species are very
similar in color pattern, but F. slevini differs from the latter primarily in
the much lower number of subdigital lamellae, 31-38 (mean = 33) for 27
specimens, instead of 45-52 (mean = 47) for 6 specimens of Emoia b. boettgeri
from the Caroline Islands, and the somewhat shorter hind limbs relative to snout-
vent length, less than 50 percent (usually greater than 50 percent for EF. boett-
geri). Emoia arnoensis and E. flavigularis are also readily distinguished from
E. slevini not only on the basis of very different color patterns, but also by the
higher subdigital lamellar counts; for E. flavigularis the number of scale rows
between the parietals and the base of the tail is usually less than 65, whereas
it is usually greater than 65 for E. slevini; and the interparietal is usually fused
with the parietals in E. flavigularis, or, if distinct, relatively small.

Of the group of related species noted in the Introduction, only £. atrocostata
is sympatric with E. slevini. Populations of both species are represented in
collections from Cocos Island, with 23 examples of E. slevini and two of E.
atrocostata. Unfortunately this widespread species, described by Lesson (1830)
on the basis of a unique specimen from Oualan (= Kusaie) Island in the
eastern Carolines, is still poorly represented in collections from most islands
of Micronesia and probably does not occur in the Marshall Islands, at least
in the eastern part. The data for E. atrocostata presented in this paper are
based upon two examples from Cocos Island and several specimens from Ulithi
Atoll and the Palau Islands. Using the limited data from these two small
samples of E. atrocostata, aside from totally unlike color patterns, E. atrocostata
would appear to be most readily distinguished from £. slevini on the basis of
the slightly greater number of midbody scale rows (table 1); the longer,
narrower interparietal (length one and one-half to two times its basal breadth) ;
and the posteriorly more broadly truncate rostral.

110 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Hasitat. Based on observations of the population on Cocos Island, E£.
slevini is a forest species. The forest on this island is predominantly tall Casuar-
ina trees with some scattered coconut and other broadleaf trees. The understory
growth is sparse and the canopy permits only diffuse sunlight to mottle the
forest floor. The forested area is bordered toward the lagoon by Scaevola shrubs.
Most of the lizards observed were active on the forest floor. Occasionally
specimens were seen in low hollows of tree trunks with only their heads protrud-
ing. Two were observed on the lower part of the trunk of Casuarina trees.
In most instances escape was attempted by hiding under material on the forest
floor. A few specimens were observed at the edges of the Scaevola shrubs.

Emoia atrocostata on Cocos Island, as elsewhere (Brown and Alcala, 1967),
occupies the various marginal beach areas or the open, often sparsely grassy
or shrub-dotted areas adjacent to the beaches.

KEY TO THE SPECIES OF EMOIA IN THE MARIANAS

1. Lamellae beneath the fourth toe greatly thinned, more than 50 E. cyanura
Lamellae) rounded; not’ thinned, less than’ 50) =e 2
. Midscale rows between the parietals and the base of tail less than 60; maturity attained
ae CH) ienbont, ie JESS tin, Selanne |esayetday Pt E. caeruleocauda
Middorsal scale rows between the parietals and ihe. base of tail greater than 60; maturity
not attained at less than 45 mm., rarely less than 50 mm., in snout-vent length 3
3. Midbody scale rows 38 to 40; rostral broadly truncate posteriorly; dorsum grayish,
greenish, or tan with black or dark brown spots; upper lateral surface marked by
iresular black vor Ganrke frown" bere ee E. atrocostata
Midbody scale rows 34 to 38; rostral rounded or narowly truncate posteriorly; dorsal
and upper lateral surfaces brownish with scattered darker markings occasionally forming

bo

a vague pattern of narrow, broken, transverse bands —_..____»_»___ E. slevini

ACKNOWLEDGMENTS

This study is part of the senior author’s program concerned with the herpeto-
fauna of the Pacific Islands with emphasis on the Philippines. The latter
program is currently supported by National Science Foundation Grant GB-
16972. Illustrations were prepared by Mr. Walter Zawojski, Stanford University.

LITERATURE CITED

Brown, WALTER C.
1956. The distribution of terrestrial reptiles in the islands of the Pacific basin.
Proceedings of the Eighth Pacific Science Congress, vol. 3, pp. 1479-1491.
Brown, WALTER C., and ANGEL C. ALCALA
1967. Population ecology of the tropical scincoid lizard, Emoza atrocostata, in the
Philippines. Copeia, 1967, pp. 596-604.
Lesson, R. P.
1830. Zoologie: In M. L. I. Duperry, Voyage autour du monde .. . sur la
corvette de sa Majeste, La Coquille, pendant les années 1822-1825. Paris,
vol. 4, 743 pp. and 157 pls.

Marine B
LIBRARY
JAN® 1973
Woods Hole, Mass,

PROCEEDINGS

‘ological Laboratory

OF THE

CALIFORNIA ACADEMY OF SCIENCES
FOURTH SERIES

Vol. XXXIX, No. 10, pp. 111-120; 5 figs. December 27, 1972

TWO NEW GENERA AND TWO NEW SPECIES
OF WESTERN PACIFIC SNAKE-EELS
(APODES: OPHICHTHIDAE)

ey)
John E. McCosker!

Scripps Institution of Oceanography
La Jolla, California 92037

ABSTRACT. Two new species representing two new genera of ophichthid eels, sub-
family Ophichthinae, are described from the western Pacific Ocean. Evips
percinctus, from Palau, and Allips concolor, from Thailand, are described and
figured.

INTRODUCTION

Examination of the extensive collections of Indo-Pacific fishes deposited
at the California Academy of Sciences has disclosed the presence of two new
species of ophichthid eels, subfamily Ophichthinae, each of which is distinctly
different from any member of a known genus. Difficulty in obtaining specimens
of sand-dwelling eels has been noted in recent works (McCosker and Rosen-
blatt, 1972), yet the two individuals upon which this report is based represent
but a small fraction of the numerous ophichthids, moringuids, and xenocongrids
collected by the George Vanderbilt and Naga expeditions to the central and
western Pacific Ocean. It is the author’s intent in this study to make these
generic names available in preparation for his more thorough osteological
comparison of genera within the family Ophichthidae.

Measurements are straight-line, made either with a 300 mm. ruler with

1Contribution from the Scripps Institution of Oceanography, University of California, San Diego.

Tilia]

112

CALIFORNIA ACADEMY OF SCIENCES

[Proc. 4TH Serr.

a aU ee iy id
JANG 1973
Wosds Hole, Mass.
Vor. XXXIX] McCOSKER: NEW SNAKE EELS a | ee
0.5 mm. gradations (for total length, trunk length, and tail length) and
recorded to the nearest 0.5 mm., or with dial calipers (all other measurements)
and recorded to the nearest 0.1 mm. Head length is measured from the snout
tip to the posterodorsal margin of the gill opening; trunk length is taken
from the end of the head to mid-anus; body depth does not include the fin.
Branchiostegal and vertebral counts (which include the hypural) were made
using radiographs. Gill arches were prepared by means of the Taylor (1967)
trypsin technique. Material used in this study is housed in the California
Academy of Sciences (CAS) or the Marine Vertebrates Collection of the Scripps
Institution of Oceanography.
I thank Richard H. Rosenblatt for reading this manuscript critically,
Carl L. Hubbs for suggestions, and William N. Eschmeyer of the California
Academy of Sciences for permission to utilize specimens in his care.

TAXONOMY
Evips McCosker, new genus

Driacnosis. Body nearly cylindrical, head and trunk longer than tail.
Dorsal and anal low, ending before pointed tail tip; dorsal origin above and
slightly behind gill openings. Pectoral rudimentary and less than eye dia-
meter, its base in upper corner of gill opening. Gill openings lateral and
shorter than isthmus breadth. Underside of snout not grooved. Anterior
nostril tubular, posterior nostrils open into mouth. Teeth pointed; inter-
maxillary teeth largest and depressible, vomerine teeth smaller and_ fixed.
Eye large. Other characters those of the single species.

Type species. Evips percinctus McCosker, new species.

EtymMotocy. From the Greek «i (eu, latinized to ev for euphony before a
vowel), good, and iy (ips, masculine), a worm, in reference to the general
appearance of this charming eel.

RELATIONSHIPS. Evips percinctus appears most closely related to the more
generalized ophichthines such as species of Ophichthus, Microdonophis, and
Pogonophis. Similarities among these genera include the retention of the
pectoral and median fins, the lateral slightly restricted gill opening, the
posterior nostril and head pore conditions, and the generally bold coloration.
Osteologically, E. percinctus also appears similar to them in its pectoral
girdle and hyoid arch conditions (viewed from radiographs and cleared and
stained specimens) and in the condition of the gill arches, primarily the
retention of the fifth ceratobranchial and the separation of UP; and UP,

<

Ficure 1. Evips percinctus McCosker, new species, CAS no. 13966, holotype, 125.5
mm. TL. Arrows indicate the origin and termination of the median fins.

114 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Ficure 2. Evips percinctus McCosker, new species, CAS no. 13966, holotype. Arrows
indicate location of median interorbital and temporal pores.

(Nelson, 1966; personal observation). The new genus is separable from
these and other ophichthines on the basis of its rudimentary pectoral fin,
robust body, and short tail.

Evips percinctus McCosker, new species.
(Figures 1-3.)

Holotype and only known specimen, CAS no. 13966, a 125.5 mm. juvenile
from Kayangel Island, Palau Islands, Southern Caroline Archipelago
(8°02’18’N., 134°43’21”E.), collected in shallow water (0-4”) by H. A.
Fehlmann and party during the George Vanderbilt Foundation 1956 Palau
Islands Expedition, on 8 October 1956.

DESCRIPTION. (Measurements are in millimeters.) Total length 125.5,
head 14.0, trunk 60.0, tail 51.5, predorsal 17.2, body depth behind gill
opening 5.2, at anus 4.5, snout length 2.8, upper jaw 4.8, eye diameter 1.4,
interorbital width 1.5, gill opening height 0.9, isthmus width 3.1. Vertebrae
132; 69 to anal fin origin.

Body stout (not exceedingly elongate as in many ophichthids) and nearly
cylindrical. Depth behind gill openings 24 times in total length, and at
anus 28; width behind gill openings 31 times in total length, and at anus
34. Snout blunt. Lower jaw included, its tip beneath a line drawn from
anterior nostril base. Eye large, about 3.4 times in upper jaw, its center
opposite midpoint of upper jaw. Anterior nostrils tubular, about 1.5 in eye
diameter. Posterior nostrils open into mouth and lie ahead of anterior margin
of eye. Tongue adnate. Branchial basket expanded and supported by numerous
branchiostegals and jugostegalia which broadly overlap along the ventral
midline. Numerous papillae on snout, beneath eye, and on anterolateral
flanges of upper lip (fig. 2). Dorsal origin above and slightly behind gill

VoL. XXXIX] McCOSKER: NEW SNAKE EELS 115

Ficure 3. Dentition of holotype of Evips percinctus McCosker, new species.

opening. Pectoral fin rudimentary, a small semicircular flap, about equal
in length to anterior nostril and attached above midpoint of gill opening.
Dorsal and anal low, disappearing in advance of tail tip. Caudal rays
lacking.

Head pores conspicuous; preoperculomandibular, temporal, suborbital,
postorbital, and supraorbital series present (fig. 2). A single median inter-
orbital and temporal pore. Two postocular pores, lying within faintly pig-

116 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

mented spots. Left lateral line pores 125; 9 before gill opening and 73
before the anus, terminating 0.3 head length before tail tip.

Teeth pointed and depressible (fig. 3). Four anterior peripheral inter-
maxillary teeth covered by skin folds, flanking a median intermaxillary pair,
followed by seven uniserial vomerine teeth. Maxillary teeth biserial, the
outer row covered by a skin fold and larger anteriorly, the inner 4 prom-
inent. Mandibular teeth uniserial, about 10 on each side and grading smaller
posteriorly.

Gill arches removed, and cleared and stained. First basibranchial os-
sified, second cartilaginous, third rudimentary, fourth absent. Hypobranch-
ials 1 and 2 ossified, 3 cartilaginous. Ceratobranchials 1—5 ossified, the
fifth a slender filament. Second and third infrapharyngobranchials ossified.
Lower pharyngeal tooth plate slender with about 25 conical biserial teeth.
Upper pharyngeal tooth plate (UP; and UP, of Nelson, 1966) separate
(unfused) and subrectangular, with about 15 conical teeth.

Color in isopropanol yellow, overlain dorsally with 16 brown saddles.
First saddle just behind occiput, does not reach sides of head; remaining
15 extend below lateral midline but do not meet ventrally. The type has
retained traces of larval pigmentation, visible as 9 minute black pigment
patches evenly spaced along the ventral midline of the trunk. Median fins
unpigmented.

EtymoLocy. From Latin, signifying banded throughout.

Allips McCosker, new genus

Dtacnosis. Body very elongate, nearly cylindrical for most of its length;
head and trunk longer than tail. Dorsal and anal low and lying within a
shallow groove, ending before bluntly pointed tail tip; dorsal origin well
behind gill openings. Pectoral rudimentary, a tiny flap in upper rear corner
of gill opening. Gill openings lateral and low on sides, separated by an
isthmus wider than their length. Underside of snout grooved. Jaw teeth
and vomerine teeth small and pointed. Eye minute. Other characters those
of the single species.

Type species. Allips concolor McCosker, new species.

Etymo.ocy. From the Greek adAos (allos), another, and iy (ips, masculine),
a worm.

RELATIONSHIPS. Allips concolor appears most similar to extant species of
Bascanichthys, Phaenomonas, and Gordiichthys. The last is a poorly known
ophichthid excluded from the recent generic treatment of Rosenblatt and
McCosker (1970). It will be redescribed on the basis of new material by
James E. Bohlke (personal communication). Allips differs from Bascanichthys
in having a more posterior dorsal fin origin, a more rounded snout, and
a more cylindrical trunk and tail. The two genera are quite similar in having

Vor. XXXIX] McCOSKER: NEW SNAKE EELS 117

Ficure 4. Head of holotype of Allips concolor McCosker, new species, CAS 13967,
375 mm. TL.

small nearly fixed teeth, comparable gill arch configurations (although the
fifth ceratobranchial, absent in Adlips, is very reduced but retained in certain
species of Bascanichthys), low crescentic gill openings, rudimentary pectoral
fins, a grooved snout, reduced eyes, highly rugose skin regions, a nearly
uniform coloration (although certain species of Bascanichthys are often darkly
pigmented dorsally), and similar head and body shapes and _ proportions.
Phaenomonas differs in its extreme dorsal fin reduction, complete loss of
the pectoral and anal fins, smaller eye, and the elongation of the trunk
region.

Allips concolor McCosker, new species.
(Figures 4-5.)

Holotype and only known specimen, CAS no. 13967, a 375 mm. specimen
from Goh Phi, Ranong Province, Thailand (10°57’42’”N., 98°35'18”E.),
north of Ban Parknam Ranong. Collected in shallow water (0-3) at the
mouth of Pakehan river by H. A. Fehlmann and party during the 1959-1961
Naga Expedition, on 1 June 1960.

DESCRIPTION. (Measurements are in millimeters.) Total length 375,
head 21.0, trunk 205, tail 149, predorsal 48, body depth behind gill openings
5.6, at anus 5.0, snout length 3.0, upper jaw 5.2, eye diameter 0.6, interorb-
ital width 2.6, gill opening height 1.7, isthmus width 2.6. Vertebrae 174;
96 to anal fin origin.

Body elongate and nearly cylindrical, becoming laterally compressed only
near tail tip. Depth behind gill openings 67 times in total length, and at
anus 75; width behind gill openings 83 times in total length, and at anus
86. Snout subconical, rounded at tip. Lower jaw included, its tip behind
anterior nostrils. Eye small, 8.5 in upper jaw, and faintly visible under
skin; its midpoint closer to corner of mouth than snout tip. Anterior

118 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

\

busy

a

Ficure 5. Dentition of holotype of Allips concolor McCosker, new species.

nostrils in a short tube; posterior nostrils open into mouth beneath eye.
Tongue adnate. Branchial basket expanded and supported by 21 pairs of
branchiostegals and jugostegalia which broadly overlap along ventral midline.
Sides of head, throat, chin, and body flanks markedly rugose. Dorsal fin
origin well behind gill openings. Pectoral fin rudimentary, a small flap
attached to upper rear margin of gill opening. Dorsal and anal fins low,
lying within a groove and disappearing 0.85 head length before finless caudal
tip. The median and posterior caudal edges are covered with a band of
numerous small papillae.

Head pores reduced, but preoperculomandibular, temporal, suborbital,

VoL. XXXIX] McCOSKER: NEW SNAKE EELS 119

postorbital, and supraorbital series present (fig. 4). A single median inter-
orbital and temporal pore. Lateral line canal and pores present, but impossible
to discern because of their reduced state and a waxy precipitate which has
formed in preservation.

Teeth small, pointed, and close set (fig. 5). Jaw teeth and posterior
vomerine teeth uniserial.

Gill arches removed, and cleared and stained. First basibranchial os-
sified, second cartilaginous, third absent, fourth cartilaginous but rudimentary.
Hypobranchials 1 and 2 ossified, 3 cartilaginous. Ceratobranchials 1—4 os-
sified, 5 absent. Second and third infrapharyngobranchials ossified. Upper
and lower pharyngeal tooth plates are subrectangular patches with conical
teeth; UPs and UP, separate.

Color in isopropanol nearly uniform brown although slightly darker on
upper half due to small dark punctations.

EtymoLtocy. From Latin, in reference to the nearly uniform coloration.

LITERATURE CITED

McCosker, JoHN E., and R1icHarp H. ROSENBLATT
1972. Eastern Pacific snake-eels of the genus Callechelys (Apodes: Ophichthidae).
Transactions of the San Diego Society of Natural History, vol. 17, no. 2,
pp. 15-24, 5 figs.
NELSON, GARETH J.
1966. Gill arches of teleostean fishes of the order Anguilliformes. Pacific Science,
vol. 20, no. 4, pp. 391-408, 58 figs.
ROSENBLATT, RicHarD H., and Joun E. McCosker
1970. A key to the genera of the ophichthid eels, with descriptions of two new
genera and three new species from the eastern Pacific. Pacific Science, vol.
24, no. 4, pp. 494-505, 8 figs.
TayiLor, WILLIAM RALPH
1967. An enzyme method of clearing and staining small vertebrates. Proceedings of
the United States National Museum, vol. 122, no. 3596, 17 pp.

PROCEEDINGS 0005 Hole, Vase.

OF THE

CALIFORNIA ACADEMY OF SCIENCES
FOURTH SERIES

Vol. XXXIX, No. 11, pp. 121-140; 6 figs.; 3 tables. December 27, 1972

FIVE NEW INDO-PACIFIC PIPEFISHES

By

Earl S. Herald
Steinhart Aquarium, California Academy of Sciences
San Francisco, California 94118

and

John E. Randall
Bernice P. Bishop Museum, P.O. Box 6037
Honolulu, Hawaii 96818

During the 1946 fish investigations at Bikini Atoll in the Marshall Islands,
V. E. Brock and Herald carried out the first moderate-depth rotenone stations
for the collection of fishes. Since this was before the development of scuba,
the diving gear consisted of Brock’s modified Monson lung rebreather with
other divers receiving air supplied through a standard dive line attached to
a small compressor. It was soon evident from these collections that an
important break occurred in the vertical distribution of the fishes. Many
were confined to the intertidal zone or a few feet below; others were common
in water of about 15 feet or greater and rarely, if ever, were seen in
shallow water. The Syngnathidae provided a good example of this faunal
break. Of the ten species of pipefishes (47 specimens from 26 stations)
which were collected in the Marshall Islands, five were found only at
depths below 15 feet, and three of those were undescribed.

With the exception of Dentirostrum janssi, the other four species of pipe-
fishes described as new herein appear to belong to the deeper water group,
being taken at depths ranging from 20 to 160 feet. One species was
collected in the Hawaiian Islands, one in the Marianas, the third in the

[121]

122 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

TABLE 1. Meristic comparisons for the Dunckerocampus complex.

Num- Trunk
ber of Rings Tail Rings Dorsal Fin Rays
eee 15) 16117 16) 7 18 19: 20) 20) 22 2320022023678 550
Dunckerocampus
multiannulatus* g* ipa i fF il 1? Seas
D. pessuliferus 1 1 1 1
D. c. caulleryi 1 1 1 1
D. c. chapmani 3 I 2 LZ: ih
D. dactyliophorus 59 1 58 10 37 10 2 21318 14 8 5
D. baldwini 20 20 Wo @ 1 ey 2 2
Dentirostrum janssi 34 34 1 426 3 1 3 16 14

‘Includes literature holotype data for D. multiannulatus and D. ben-tuviae. It is probable that trunk
ring counts for these two types should be recorded as 16 rather than 17.

Ryukyus, and the fourth at Easter and Pitcairn islands. These are island
groups in which considerable fish collecting has been carried out in shallow
water in recent years.

The following abbreviations are used for institutions in this paper: AMS
(Australian Museum-Sydney); ANSP (Academy of Natural Sciences of Phil-
adelphia); BM(NH) [British Museum (Natural History)]; BPBM (Bernice
P. Bishop Museum); CAS (California Academy of Sciences); LACM (Los
Angeles County Museum of Natural History); SMF (Senckenberg Museum) ;
and USNM (United States National Museum).

Technical assistance in making counts and measurements has been pro-
vided by Mr. D. Anderson.

THE DUNCKEROCAMPUS COMPLEX

Seven long-nosed species of very spiny belly-pouch Indo-Pacific pipefishes
are members of the Dunckerocampus complex. All of these including the
related Doryrhamphus and Oostethus pipefishes have a first trunk ring,
i.e., the one bearing the pectoral fins, that is twice as long as any of the
remaining trunk rings. Although it is actually double, it is counted as a
single ring. In meristic characters these species are very similar (table
1). Based on pigment patterns or the absence thereof, members of the
Dunckerocampus complex fall into four categories: the first is the narrow-
banded group whose two members usually have 4 or 5 very narrow pigment
bands on the opercle, two or more on each trunk ring and sometimes on
each tail ring. Dunckerocampus multiannulatus (Regan) 1903, was described
from Mauritius and was characterized by a lack of banding on the snout.

VoL. XXXIX] HERALD & RANDALL: PIPEFISHES 123

Dunckerocampus ben-tuviae Fowler and Steinitz 1956, described from the
Red Sea had the same meristic characters but did have banding on the
snout. Dr. Eugenie Clark advises us that Red Sea pipefishes of the D.
multiannulatus/D. ben-tuviae group have a wide range of snout banding
variability so that D. ben-tuviae will need to be considered a synonym of
D. multiannulatus.

The remaining species in the narrow-banded group, Dunckerocampus
pessuliferous, is distinct from other Dunckerocampus in having a high dorsal
fin count of 30 as compared with a normal range of 20-25. It is known
from the holotype taken by dredge at a depth of 144 feet at Sulade Island
in the southern Philippines.

The two species in the wide-band group usually have a single pigment
ring for each trunk and tail ring. The most common species is the central
and western Pacific D. dactyliophorus; it has a single ring around the opercle
which differentiates it from the twin opercular rings of D. caulleryi Chabanaud
known only from Amboina and New Caledonia. This latter species has two
subspecies: D. c. caulleryi with 19 tail rings, and D. c. chapmani Herald
with 16 or 17 tail rings. More material is needed to determine the status
of these two.

The third category is represented by the striped Hawaiian endemic described
as new herein: Dunckerocampus baldwini. This species extends the generic
range from the Austral Islands northward to the Hawaiian region, a distance
of about 2800 miles. And, finally, the fourth category is that group of pipe-
fishes that, unlike Dunckerocampus, developed the brood pouch folds; the
new genus and species for this is Dentirostrum janssi described as follows:

Dentirostrum Herald and Randall, new genus

Diacnosis. Spiny, long-snouted Doryrhamphine pipefish with abdominal
brood area protected by lateral membranous folds but without protecting
plates. Lateral trunk ridge continuous with inferior tail ridge at anal ring;
superior trunk and tail ridges discontinuous at end of dorsal fin; lateral
tail ridge ending free at anal ring. Tail non-prehensile. Dorsal fin rays
21-24, anal 4, pectoral 19-21, caudal 10; trunk rings 16; tail rings 20-
23. Named Dentirostrum in reference to the magnificent spines present on
the median snout ridge of males and to a lesser extent on females.

Discussion. Dentirostrum is most closely related to Dunckerocampus
from which it differs, (1) in having brood pouch flaps rather than having
the eggs nakedly attached to the abdomen, (2) in its extremely spinose
nose ridge, and (3) in the presence of a secondary spine behind the primary
spine at the ring juncture of the superior and inferior trunk ridges. Den-
tirostrum and Doryrhamphus are similar in that both have brood-pouch
folds, but the latter differs in its small size and its short snout.

124 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH Serr.

The type species of Dentirostrum, D. janssi, is new and is the only
known member of the genus. The description of D. janssi based on 34
specimens (14 males 61-126 mm. SL and 20 females 52-110 mm. SL)
from 19 localities in the Indo-Malayan region is as follows:

Dentirostrum janssi Herald and Randall, new species.
(Figure 1.)

From the Patau IsLaAnps: 13 types, 6 localities.

Hototype. CAS 14139, male 125 mm. SL, with 126 brood patch sockets—
a few with eggs; Arappu Point of Koror to Ho Island, south side of reef;
0-50 feet; H. DeWitt; November 19, 1957 (GVF 1442).

ParATYPES. CAS 14140, male 123 mm. SL; same data as holotype. CAS
14141, male 105 mm. SL; south entrance to small bay on west side of
Ngalab Point, Koror Island; 0-50 feet; H. DeWitt; November 21, 1957
(GVF 1445). CAS 14142, female 89 mm. SL; Iwayama Bay, off Kaibakku
Island; 0-45 feet; H. DeWitt; October 3, 1957 (GVF 1408). AMS 16144-001,
male 103 mm. SL; female 105 mm. SL; LACM 32123-1, male 105 mm.
SL; SMF 11425, male 111 mm. SL; channel between Sanryo Island and
Ngatkumer Island, Iwayama Bay; 0-50 feet; H. DeWitt; October 31, 1957
(GVF 1433). CAS 14143, 3 females, 53 mm., 97 mm., 103 mm. SL; channel
between Sanryo Island and Kamori Island, Iwayama Bay; 0-42 feet; H.
DeWitt; November 18, 1957 (GVF 1439). BPBM 11937, male 102 mm.
SL; female 83 mm. SL; western tip of Ngargol Island; 20-40 feet, coral;
J. E. Randall, E. 5. Heliman, O. Custer; June 8, 1968.

From THAILAND: 16 paratypes, 9 localities.

BPBM 11938, female 91 mm. SL; NW. side of Koh Kroi Island about
5.5 miles ESE. of Ban Pae Fisheries Training Center, Rayong Province;
0-4 meters; H. A. Fehlmann; April 30, 1960 (GVF 2183). CAS 14146, male
77 mm. SL; NW. side of Goh Samet Island, Rayong Province; 0-2 meters;
B. Bronson; April 29, 1960 (GVF 2180). CAS 14147, 3 females, 62 mm.,
64 mm., 83 mm. SL; NW. side of Goh Raed about 1.5 miles ESE. of
Prachuap Khiri Khan town, Prachuap Khiri Khan Province; 0-15 feet
(photo 3-79 a-c); H. A. Fehlmann; June 19, 1961 (GVF 2651). USNM
206654, male 126 mm. SL; female 78.5 mm. SL; CAS 14148, 2 males 80
mm. and 61 mm. SL; female 59 mm. SL; W. side of Goh Luem ca. 3
miles SE. of Prachuap Khiri Khan Town, Prachuap Khiri Khan Province;
0-15 feet; H. A. Fehlmann; June 18, 1961 (GVF 2648). CAS 14149, 2
females 95 mm. and 102 mm. SL; W. side of Goh Luem, Prachuap Khiri
Khan Province; 0-15 feet; H. A. Fehlmann; June 17, 1961 (GVF 2646).
CAS 14150, female 111 mm. SL; on fringing reef at head of Gulf of Siam,
Goh Sak Island, about 35 miles NE. of Prachuap Khiri Khan town, Prachuap

VoL. XXXIX] HERALD & RANDALL: PIPEFISHES 125

PAPA saay.
aoaaeaaaners

a a a ae

mm

Ficure 1. Dentirostrum janssi Herald and Randall, Holotype; male 125 mm. SL
(CAS 14139); (a) lateral view; (b) expanded ventral view of brood patch area; (c)
head detail male holotype compared with (d) head detail female 111 mm. SL paratype
(CAS 14150). Drawing by H. Hamman.

Khiri Khan Province; 0-20 feet; H. A. Fehlmann and R. Rofen; October
24, 1957 (GVF 1460). CAS 14151, female 58 mm. SL; SW. point on Goh
Chorakhay, Chumphon Province; 0-16 feet; B. Bronson; May 25, 1960
(GVF 2199). CAS 14152, male 91 mm. SL; fringing reef on NW. side
of Goh Maprao, Chumphon Province; 0-12 feet; H. A. Fehlmann; May
18, 1960 (GVF 2186). CAS 14153, male 105 mm. SL; Prond Bay at SW.

126 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

corner of Goh Samed, ca. 100 meters from shore, Chumphon Province; 0-
10 feet; H. A. Fehlmann; May 17, 1960 (GVF 2185).

From OTHER LOCALITIES”: 5 paratypes.

CAS 14145, 2 females 96 mm. and 107 mm. SL; Bay of Nhatrang
SW. of Hon Long, Vietnam, South China Sea; 1-15 meters; R. Bolin;
March 14, 1960 (GVF 2116). BPBM 11939, female 93 mm. SL; Bay of
Nhatrang, Hon Long, Ving Damlon, Vietnam, South China Sea; 1-6 meters;
R. Bolin; February 23, 1960 (GVF 2072). CAS 14144, male 93 mm.
SL; Surabaja, Java, Indonesia; Hilmi Oesman; February 1961. ANSP 119933,
female 99 mm. SL; Little Hope Island, NE. end, Queensland, Australia;
5-8 meters; J. Tyler, C. L. Smith, and G. Bettle; January 3, 1969.

It will be noted that the distribution of Dentirostrum janssi does not
include the Philippines although it is known from west, east, and south
of that area. It is probable that it will be found there when rotenone
collecting is conducted by diving methods using scuba.

Diacnosis. Dorsal fin rays 21-24, usually 23-24; dorsal covering a total
of 5—6% rings of which 1-2 are trunk rings and 4-5 are tail rings; usual
count: 1 + 5. Trunk rings 16, tail rings 20-23, usually 22; pectoral 19-
21, usually 20; anal 4, caudal 10. Head-in-standard length 4.2—5.0; snout-
in-head 1.5—1.9; dorsal-base-in-head 1.9-2.8; brood pouch folds extending
from second trunk ring to anal ring, and occasionally as far as 4th tail
ring. Eggs large, about .75 mm. diameter; largest number in belly brood
patch: 195 (105 mm. male). Ridge system and brood patch folds typical
of Dentirostrum.

DescripTION. Nearly all ridges of head, trunk, and tail are spinose. The
median snout ridge of the males has 15-21 large spines, and on each side
there is a lateral ridge with smaller spines. The spines on females are less
accentuated. The supranasal ridge is present. The anterior ocular ridge
is lacking, and the posterior ridge is smooth. The orbit is ringed with small
spines. The opercle has a smooth median ridge over its entire length, and
beneath it may have 5-7 additional radiating ridges (compare figs. le and
1d). The pectoral cover plate has 1 or 2 spines on its upper anterior
edge, and a single spine in the center. Superior and inferior trunk ridges have
double spines at each ring juncture, but on the tail only a single spine
at each ring. The lateral trunk ridge has double spines for the first 5-7
rings, and then a single spine for the next 9-11 rings.

Between 62 and 80 mm. the future brood patch folds show their presence
by a pair of darkly pigmented lines on the abdominal surface of the male.
Among the 14 males only two had eggs attached to the brood patch: the

2A new male Dentirostrum janssi (106 mm. SL) has just arrived; it is a depth record (100 feet)
captured at Kranket I., Madang, New Guinea; G. R. Allen, May 15, 1972.

VoL. XXXIX] HERALD & RANDALL: PIPEFISHES 127

125 mm. male holotype (126 brood patch sockets, a few with eggs) and a
103 mm. male (21 eggs and 135 empty sockets = 156). These were collected
in October and November in the Palaus. Another male of 111 mm. collected
at the same time and place showed 156 empty egg sockets. Two males,
91 and 105 mm. from Chumphon Province, Gulf of Thailand (May collection),
showed 106 and 195 empty sockets. All 33 specimens had 16 trunk rings,
and the male brood patch folds usually cover 15 of the 16. However, five
of the 14 males had the folds extending upon the tail, the furthest distance
being 3% tail rings. Although the range of brood pouch coverage was 15
to 19% rings there was no indication of vascularity on the tail portion of the
pouches. It is probable that this small tail section is not used as a brood
area.

The overall color of Dentirostrum janssi is light brown with two pairs
of dark brown lines extending from the head to the dorsal fin with one
on each side of each superior trunk ridge. The first pair extends from above
the eye dorsally on the inside edge of the superior trunk ridge, whereas the
second pair extends from the center part of the eye over the opercle and
along the upper lateral side of the superior trunk ridge. The tail fin has
a clear center section surrounded by a dark reddish black area. The junior
author’s field notes on two males collected in the Palaus at Ngargol Island
indicate that the body is blackish anteriorly and posteriorly with orange in
the middle. Also, the pigment around the tail fin shows a small amount
of white on the outer edge.

Discussion. From the upper west side of the Gulf of Siam (GVF 64:
BanAangtong Bay; CAS 14155), we have three small specimens (33, 33.5,
and 42 mm. SL) that have all of the characteristics of D. janssi except that
they have a single spine on each ring of trunk and tail and are lacking the
double spines of the trunk of D. janssi. Our smallest specimen of the latter
species is 58 mm. SL (CAS 14151), and it has the physical characteristics
of the adults. Among the syngnathids, spines are usually lost with growth,
not added. Because of the lack of spines on these three specimens, we are
faced with the possibility of another species of Dentirostrum existing in this
area. However, we will not describe these as new at this time, but will
await the collection of adult material from the same area.

At this same GVF station 64, juvenile representatives of two additional
species of syngnathids were collected: 8 specimens (average length 44 mm.)
of Corythoichthys species (CAS 14318) and a 61 mm. specimen tentatively
identified as Syngnathus maxweberi (CAS 14317).

Named ‘janssi? in honor of Mr. Edwin Janss, Jr., whose keen interest
in tropical marine biology has resulted in important field investigations in
many regions.

128 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

FicurRE 2. Dunckerocampus baldwini Herald and Randall, Holotype; male 125 mm.
SL (CAS 24734); cave SE. of Pokai Bay, Waianae coast, Oahu; 70 feet. Photograph
by J. E. Randall.

Dunckerocampus baldwini Herald and Randall, new species.
(Figure 2.)

Twenty type specimens from the Hawaiian Islands: Oahu (17), Hawaii
(2), and Maui (1). Capture methods: by hand, quinaldine, and chemfish
ichthyocide. Depth range 20-160 feet; size range 63-131 mm. SL.

HototypPe. CAS 24734, male 120.5 mm. SL (125.5 mm. total length);
cave SE. of Pokai Bay, Waianae coast, Oahu; 70 feet; J. E. Randall and
S. Swerdloff; July 20, 1969.

PARATYPES. CAS 24735, female 125 mm. SL; same data as holotype.
LACM 32126-1, female 126 mm. SL; CAS 14138, 2 males 106.5 and 129
mm. SL; 3 females 129, 131, and 131 mm. SL; Waianae coast off Makaha
Shores Condominium, Oahu; 45 feet; J. E. Randall and A. R. Emery; April
26, 1970. BPBM 11941, 2 females 63 and 128 mm. SL: Waianae coast
off Lahilahi Point, Oahu; 40 feet; J. E. Randall and P. M. Allen; July
11, 1970. SMF 11426, female 128 mm. SL; Kawaihoa Point, Maunalua

VoL. XXXIX] HERALD & RANDALL: PIPEFISHES 129

Bay, Oahu; 35 feet, lava caves; Jane Culp; November 15, 1971. AMS
15607-001, female 123.5 mm. SL; CAS 14137, female 134 mm. SL; Kawaihoa
Point, Maunalua Bay, Oahu; 35 feet, lava caves; Jane Culp and James
Moore; December 18, 1971. USNM 204683, female 92 mm. SL; off north
side at base of dropoff, Moku Manu, Oahu; 160 feet; J. E. Randall, T.
Stark, W. Baldwin; October 9, 1969. BM(NH) 970.1.26:1, female (?)
72 mm. SL, Moku Manu, Oahu; 85 feet; J. E. Randall and W. Baldwin;
October 6, 1969. BPBM 7783, male 124 mm. SL; female 125 mm. SL;
Kanoehe Bay at channel entrance, Oahu; 95 feet; J. E. Randall, D. Chave,
W. Hashimoto; October 10, 1969. BPBM 11940, male 102 mm. SL (tail
broken at 11th ring; caudal regenerated); north end of Honaunau Bay, Kona
coast, Hawaii; 150 feet; J. E. Randall, E. S. Hobson, J. R. Chess; August
16, 1969. BPBM 11942, male 130 mm. SL; first point north of Hanakahau
Boat Harbor, Kona Coast, Hawaii; 100 feet; J. E. Randall; August 18,
1970. CAS 14979, female 117 mm. SL; 1% miles north of Lahina, Maui,
Hawaii; 20 feet; Steinhart Divers; June 29, 1972.

Diacnosis. Holotype belly brood area covering % 12 % trunk rings with
143 egg sockets—73 on right and 70 on left. Dorsal fin rays 21-23; pectoral
19-21; anal 4; caudal 10; dorsal fin covering %—1 trunk ring and 3% tail
rings; usually “4 + 3; total rings covered by dorsal 3%—4%; trunk rings
16; tail rings 20-21; head-in-standard length 4.5—5.2; snout-in-head 1.5—
1.7; dorsal fin base-in-head 2.9-3.5. Ridge system typical of Dunckerocampus,
i.e., lateral trunk ridge continuous with inferior tail ridge; median tail ridge
extending forward to anal ring. Tail fin long, 5-7 mm. in adults of 100
mm. or larger. General appearance except color similar to other members
of genus Dunckerocampus. Color in life reddish, fading to dark with preserva-
tion. Red stripe along upper part of body from snout to tail. Edges of
caudal fin white.

DeEscriPTION. Median snout ridge with many serrations; orbital, opercular,
nuchal, and pectoral cover plate crests faintly visible. Eye very large, the
orbit diameter contained almost two times in minimum snout depth or %
length of opercle. Intermedial scutella between rings small and equal in
width to one-half distance between scutella. Body ridges well developed with
sharp spine projecting at posterior edge of juncture of each ring.

The following color description was made from 102 mm. Honaunau Bay
male shortly after capture:

“light pinkish gray with broad dorso-lateral bright orange-red stripe becoming
blackish red and narrower on snout; also a broad orange-red stripe midventrally
extending forward on snout as red and blackish red. Snout yellowish, white
dorsally and yellow between lateral and ventral blackish red stripes; tail nearly
entirely red (red stripes close together); caudal fin red with a narrow white
edge at sides, and a black area distally on 2 upper central interradial membranes;
dorsal colorless.”’

130 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

In preservative the males apparently retain the color stripe along the
head and trunk much better than do the females. The 125 mm. female
from Kaneohe Bay (BPBM 7783) has a conspicuous blackish tip on the
outer margin of the last five rays of the dorsal fin. Other specimens did
not show this character.

Discussion. Capture data and dive observation records indicate that
specimens of Dunckerocampus baldwini are often found in pairs. In captivity
longevity has been short, usually no more than 2 months. This compares
with about three months longevity for the banded pipefish, D. dactyliophorus.
In recent years this latter species has often been imported into the United
States from the Philippines.

Cleaning activity by Dunckerocampus baldwini has been observed by
Dr. Edith Chave. While diving at Milolii, Hawaii (August 4, 1970), in a
cave at a depth of 75 feet, she watched an adult redstripe pipefish clean
a cardinal fish, Apogon evermanni, then a moray eel, Gymnothorax species,
and finally the pipefish attempted to clean Dr. Chave’s wrist.

Only two other species of pipefishes are known to act as cleaners. Randall
(1962) reported D. P. Wilson’s observations at the Plymouth Aquarium of
a John Dory being cleaned by the snake pipefish, Entelurus aequoreus. The
junior author has also observed the flagtail pipefish, Doryrhamphus mela-
nopleura, cleaning reef fishes, specifically a moray and a cardinal (Randall
and Helfman, 1972).

In recognition of his study of Pacific fishes, the redstripe pipefish,
Dunckerocampus baldwini, is named in honor of Wayne J. Baldwin who
with the junior author collected the first specimens of this new species.

Dunckerocampus dactyliophorus (Bleeker) 1853.
Banded Pipefish.

Syngnathus dactyliophorus Bleeker, Nat. Tijdschr. Nederl. Indiee, vol. 4, p. 506, 1853
(type locality, Onrust Island, Djakarta [Batavia], Java).

Herald (1953, p. 252) studied geographic variability of meristic characters
in a small series of Dunckerocampus dactyliophorus. He postulated sexual
dimorphism in the trunk color ring pattern with females having a greater
number than males. Tables 2 and 3 present data on the 62 specimens from
22 localities now available. Unfortunately, there does not appear to be any
recognizable pattern that can be correlated with sex or area. It will be
noted that the trunk ring count is remarkably constant at 16 with only one
of 62 having a different count, i.e., 15. Size range for the 62 specimens
was 41 to 159 mm. SL. The smallest mature male showing egg sockets on
the brood patch area was 90 mm.

The banded pipefish has some interesting color ring variations. The typical

VoL. XXXIX] HERALD & RANDALL: PIPEFISHES 131

TABLE 2. Geographic variation of Dunckerocampus dactyliophorus.

Trunk
Rings Tail Rings Dorsal Fin Rays
ye) ly iy IG Koy a aa 2s AON Bil BRP Bs ab DS

Australs 2 2 2
Marshalls 6 5 gS
Kapingamarangi 1 1 1
Guam 1 1 1
Ulithi 2 2 2
Palaus 1 29 9 21 o) 12 7
Philippines 6 tee v4 1 IT the Sh as
Solomons 2 il 1 1
Celebes 6 to § 3 2 1
Java 1 1
New Guinea 5 5 Se,
Totals (62) i “Oil 10 40 10 2} 2 SS 2a Oe

pattern shows about 5 rings on the snout (range 4-7), 1 over, under, and
rarely through the eye, another around the opercle, 1 at the pectoral fins,
then (table 3) 6-13 on the trunk, and 7-14 on the tail. The pattern for
any given specimen is recorded as 5-1-1-1-7-9 = 24. The width of the
individual color band is usually 42 to “% of an individual ring. However,
the two Ulithi specimens (BPBM 8746) have remarkably narrow color
bands; in width they are only equal to about % to ™% of a trunk or tail
ring. Another extreme is shown by a 91 mm. specimen from Urukthapel
in the Palaus (BPBM 7352); the color bands were very wide, 1 to 1%
rings in width.

Dunckerocampus dactyliophorus ranges through the western and central
Pacific. The type locality, Djakarta, is the westernmost point in the distribu-
tional pattern, and until recently the easternmost was 4300 miles away at
Rongelap Atoll in the Marshall Islands. However, in 1971 the junior author
diving at a depth of 185 feet collected two specimens at Rurutu (lat. 24°
S.) in the Austral Islands. This extended the range westward for another
2700 miles and southward for 900 miles from the latitude of the Solomons
(lat. 9° S.). The northernmost locality is Guam (lat. 12° N.).

Recently Dr. Eugenie Clark advised us that she has examined Dunckero-
campus dactyliophorus from the Red Sea (Eilat, Gulf of Aqaba) which is
some 8800 miles (by water) from Djakarta. This break in distribution is
similar to that which occurs in Corythoichthys flavofasciatus with the sub-
species of C. f. flavofasciatus in the Red Sea and C. f. conspicillatus many
miles away in the Central Pacific (Herald, 1953, p. 275).

132 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

TaBLe 3. Geographic variation of Dunckerocampus dactyliophorus.

Trunk Color Bands Tail Color Bands

7 Be —@) lO. abl 22 ie 7 8 9) DOI Stas
Australs i ail 1 1
Marshalls Se nlee2 1 292 1
Kapingamarangi 1 1
Guam 1 1
Ulithi i! 1 1 1
Palaus A S1S s6r 4: 5. 4)» 59 2
Philippines Wee 2e> 2k et See
Solomons 2 2
Celebes a ae 1 31 ee 1
Java 1 1
New Guinea 1 4 2M aes
Totals (62) aks ale alk alloy ao) 1 6 16° 18> PIS:

The specific collection localities for Dunckerocampus dactyliophorus listed
for the general areas of tables 2 and 3 are as follows: Marshall Islands:
Rongelap, Bikini, and Eniwetok atolls; Palaus: Koror, Auluptagel, Babelthaup,
Urukthapel, and Arakabesan islands; Solomons: New Georgia and Florida
islands; Philippines: Cebu, Jolo, Pandanan Island, and Sibutu. Although
the Philippine localities are all from the southern section, we have been advised
by Mr. Earl Kennedy, who is the major Manila live fish jobber, that the
species is fairly well distributed throughout the islands; most of his specimens
come from Luzon (Batangas), Mindoro, and Palawan. Amboina is the only
recorded locality from which we have not examined specimens.

THE PIPEFISH OF EASTER AND PITCAIRN ISLANDS

On the map of the world, Easter Island is a small dot in the eastern
south Pacific some 2000 miles west of Caldera, Chile, and about 1100 miles
east of Pitcairn, the nearest inhabited island. This isolated volcanic outcrop
has a total area of 46 square miles, being about 14 miles long and 7 miles
wide (27° 05’ S. Lat. and 109° 20’ W. Long.). Although the island is inter-
nationally famed because of the giant stone statues, the published record of
its ichthyological fauna and relationships is less well known (Randall, 1970).
In recent years three field parties of biologists (1958, 1965, and 1969) have
made collections at various sites around the island. All have used ichthyocides,
and among them they have taken ten specimens from three localities of the
first syngnathid to be collected in the area. In December 1971 the junior

VoL. XXXIX] HERALD & RANDALL: PIPEFISHES 133

author visited Pitcairn Island and was able to collect three additional specimens.
The type series consists of 13 specimens: 5 males (69-85.5 mm. SL) and
8 females (67-95 mm. SL).

Several years ago Dr. David K. Caldwell studied the Ramsey Parks
1958 series of 6 specimens at the Los Angeles County Museum of Natural
History. He tentatively determined that they represented an undescribed
species. This we are now able to verify, and we name this species in his
honor.

Syngnathus caldwelli Herald and Randall, new species.
(Figure 3.)

Hototyer. LACM 6560-3, 76 mm. standard length male with eggs in
pouch. Anakena Cove, Easter Island; Ramsey Parks, Yacht Chiriqui; boulders
and brown algae, rotenone to 4.6 m.; October 1, 1958.

PARATYPES. (5 from same collection as holotype): LACM 6560-41, 69
mm. male and 82 mm. female; CAS 24202: 71 mm. male and 65.5 mm.
female; and USNM 203409, 79 mm. female. Another collection made at
type locality 6% years later—BC 65440, 84 mm. male (pouch empty);
closed tidepools, depth % to 8 m., rotenone; Ian E. Efford and Jack A.
Mathias, Jan. 15, 1964. Ten days later at nearby Vinapu on southwest
side—BC 65449, 2 females 93 and 93.5 mm.; large boulders and rocks; depth
2-3 m., rotenone, also Efford and Mathias, Jan. 25, 1965. Finally, 10%
years after first colection—BPBM 6596, 95 mm. female (fig. 2); southwest
coast between Hanga-Roa and Hanga-piko, inshore; depth 61 cm.; boulder
bottom with brown algae; chemfish; J. E. Randall and G. R. Allen; January
26, 1969. BPBM 10856, 82 mm. male and 80 mm. female. Pitcairn Island,
off “the Rope” 20 ft., large boulders with brown algae, sand and small rocks
in low places; J. E. Randall, Dean B. Cannoy, Steve Christian, and Noggie
Young; December 23, 1970. CAS 13922, 75 mm. female, same data as BPBM
10856.

Diacnosis. Dorsal fin: 28-31; 7 specimens with 28 rays; 3 with 29;
2, 30; and 1, 31; pectoral 14-16; usually 14; anal 3; caudal 10; dorsal
fin covering 6%4—7% rings, i.e., 72-1 trunk ring and 6-6% tail rings usually
1 + 6; trunk rings 16-17; tail rings 34-36; head-in-standard length 8.5—
9.2; snout-in-head 2.2—2.6; head-in-dorsal fin base 1.06—-1.15; dorsal fin
base-in-head .87—.95. Lateral body ridges typical for genus Syngnathus: i.e.,
lateral trunk ridge interrupted at anal ring, then subcontinuous with lateral
tail ridge (65.5 mm. female has the two ridges continuous on one side
only). Brood pouch covering first 13-15 tail rings; eggs large, 1 mm. diameter,
arranged in 3—4 single rows across pouch (holotype) or 2 rows wide and 2
rows deep (69 mm. male). Holotype egg count 88; for 69 mm. male: 19

134 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Ficure 3. Syngnathus caldwelli Herald and Randall, Paratype; female 93 mm. SL
(BPBM 6596); Easter Island between Hanga-Roa and MHanga-piko. Photograph by
J. E. Randall.

and 21 eggs for dorsal rows and 14-14 for ventral rows or about 68 for
total pouch count. Brood pouch folds almost in contact but without overlap
or indentation, merely slightly thickened at free end of individual flap (modified
Open Brood Pouch Closure: O-BPC). Color in preservative, brownish some-
times with many narrow dark lines running length of body. Eye may have
dark band extending obliquely postero-ventrally over opercle. Dark spots
sometimes present at juncture point between rings on inferior trunk ridge,
and to lesser extent on lateral trunk ridge. Spots on Pitcairn pipefishes,
larger and very pronounced; also with wide whitish areas on upper surface:
about 4 on trunk and 7 on tail.

DescrRIPTION. Median snout ridge smooth, extending forward from inter-
orbital area for about half snout length. Superior ocular ridge extends

VoL. XXXIX] HERALD & RANDALL: PIPEFISHES 135

posteriorly for distance equal to eye width but is absent anteriorly. Eye
very large, in diameter about equal to 2% of snout length. Nuchal plates
bilobed, indistinct. Upper and lower pectoral ridge plates evident but not
pronounced; opercular ridge extends over % to *% length of opercle. Body
ridges evident, smooth, and not strongly pronounced. Lateral flanges of brood
pouch slightly developed.

Fretp Notes. The junior author and G. R. Allen collected the most
recent Easter Island paratype and also made color and black and white
photographs at the time of capture. From field notes the near-living color
of Syngnathus caldwelli from Easter Island is:

“light brown with row of red dots along anterior lateral ridge and full length
of ventral flange; a small red spot at front and at rear of dorsal fin; ventral
part of body light yellowish with a midventral row of red dots along trunk;
large irregular light gray blotches on back and upper side; a dark brown band
running from snout through lower half of eye where it broadens on lower opercle
and continues as a broad zone on chest; caudal fin yellowish with brown rays;
dorsal fin clear.”

Any distinctive marks on the 1958 specimens have been largely lost
following preservation. The two 1965 specimens from Vinapu are the only
ones to show clearly the narrow lines on the body, about 12 on the top
of the trunk and approximately 10 on the trunk sides. The 1965 Anakena
specimen is very dark and its markings are suggestive of the 1969 specimen
whose near-life colors are described above.

Comparisons. In the Indo-Pacific from Africa to the Americas there
are about 34 members of the genus Syngnathus, excluding Corythoichthys and
Bombonia. None of these approach Syngnathus caldwelli in their numerical
or other characteristics. The nearest relative is probably Syngnathus balli
of Hawaii. However, the latter is a smaller species (58 mm.) with fewer
tail rings (32 vs. 35-36) and fewer dorsal fin rays (21-23 vs. 28-31).

Syngnathus banneri Herald and Randall, new species.
(Figure 4.)

HototyrEs. BPBM 8695, 39 mm. SL (40 mm. TL) undet. sex; Ryukyu
Islands, Ishigaki; reef about ’2 mile off harbor of Ishigaki City; depth 20-
35 feet; chemfish ichthyocide; J. E. Randall and A. H. Banner, May 22,
1968.

PaRATYPE. CAS 14375, 26.5 mm. SL (27.3 mm. TL) immature; Marshall
Islands, Eniwetok Atoll, lagoon off Eniwetok Island; 25 feet, coral and rubble
patch; quinaldine; J. E. Randall, March 31, 1972.

Diacnosts. Dorsal fin rays 16-18 located on a total of 3'2—-4 rings (%4-—
1 trunk and 3-3% tail rings); pectoral 11; anal 2; caudal 10. Trunk rings

136 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

5 mm

Ficure 4. Syngnathus banneri Herald and Randall, Holotype; imm. 39 mm. SL
(BPBM 8695); reef near Ishigaki City, Ryukyus. Drawing by L. Sabre.

15; tail rings 27. Head-in-standard length 7.3—7.8; snout-in-head 2.5-3.2;
dorsal fin base-in-head 1.7-1.8. Body ridge pattern typical of Syngnathus,
i.e., lateral trunk ridge discontinuous with lateral tail ridge at end of anal
ring; inferior trunk and tail ridges continuous and superior trunk and tail
ridges discontinuous at end of dorsal fin. Brood pouch unknown.

DEscrIPTION. Holotype median snout ridge with two triangular flanges
just anterior to nostrils; orbital ridge markedly extended dorsally; prenuchal-
nuchal ridges present but not accentuated. Opercular ridge extending over
one-half (paratype three-quarters) of opercle. Superior and inferior pectoral
cover plate ridges sharply pronounced. Trunk and tail ridges slightly raised
and indented between individual rings so that ridgés have moderately scalloped
profile. Paratype ridges for the most part less accentuated than holotype.
Holotype color light tan with exception of dark brown spot pattern on
anterior trunk; beginning behind pectoral, 6 spots on lower trunk ridge and
5 on lateral trunk ridge on left side; spots on right side less distinct; on
dorsal surface 9 dark brown bars, one-half ring in width: 3 extending
between superior trunk ridges and 6 between superior tail ridges. Paratype
color whitish with dark area on lower sides of gill cover extending underneath
pectoral fins.

Discussion. Syngnathus banneri is a remarkable species for it has the
lowest dorsal fin count of any member of the genus (16-18 as compared
with a normal range of about 19-45), and with this is combined a very
low tail ring count of 27. When egg-bearing males are available for examination,
this species will probably prove to be a member of the subgenus Microsyngnathus
Herald 1953. These are small Svngnathine pipefishes usually less than 100
mm. in length that have overlapping brood pouch closures (O-BPC).

Named ‘banneri’? in honor of Dr. A. H. Banner whose welcome field
efforts resulted in the capture of the holotype of this species.

VoLt. XXXIX] HERALD & RANDALL: PIPEFISHES 137

Minyichthys Herald and Randall, new subgenus
of Micrognathus Duncker 1915

Type Species. Micrognathus brachyrhinus Herald 1953.

Diacnosis. Differentiated from other members of Micrognathus by in-
creased number of trunk rings (19-21 rather than 13-17), very short snout, and
small size (mature at less than 50 mm.). Named Minyichthys, small fish,
from the Greek ‘“miny” meaning small.

Discussion. Pipefishes of the genus Micrognathus have the lateral trunk
ridge continuous with the inferior tail ridge and the lateral tail ridge is
present. The tail brood pouch has everted type closure in which the outer
lip of one flap is turned back upon itself, and the other flap overlaps it
(Herald, 1959). There are three subgenera: (1) Anarchopterus Hubbs 1935,
characterized by smooth body ridges and absence of the anal fin, has two
species limited to the Gulf of Mexico and Caribbean; (2) Minyichthys, defined
above, has two Pacific species; and (3) the type subgenus Micrognathus with
its typical sharp body ridges, has two Atlantic American species and nine
Pacific species.

Micrognathus (Minyichthys) myersi Herald and Randall, new species.
(Figures 5 and 6.)

HorotyPe. CAS 13918, 42.5 mm. SL mature male (43.5 mm. TL);
Guam, NW. Cocos Island, outside of reef, 70-100 feet depth; rotenone;
June 30, 1969; J. E. Randall, e¢ al.

PARATYPE. BPBM 8759, 41 mm. SL female (43 mm. TL); Guam; south
of Uruno Point, about 10 mi. NE. of Agana; depth 60-90 feet; reef edge
adjacent to sand; June 27, 1968; J. E. Randall and H. Kami.

Diacnosis. Dorsal fin rays 29-31 covering 9-10% rings, i.e., 2%-3 trunk
rings and 64-77% tail rings; trunk rings 19, tail rings 40-41; pectoral 11-
12; anal 2, caudal 8; head-in-standard length 7.2—8.92; snout-in-head 2.36-
2.7; dorsal fin base-in-head .75—.93; pectoral base-in-pectoral length 1.6. Lat-
eral ridge pattern typical of Micrognathus, i.e., median trunk ridge continuous
with inferior tail ridge; lateral tail ridge extended forward onto 2 trunk
rings (holotype) or 2% (paratype); superior trunk and superior tail ridges
interrupted at posterior edge of dorsal fin. Distinctive spike on dorso-
median snout ridge just ahead of nostrils. Brood pouch covering first 13
tail rings with embryos to 10th ring; 7 large embryo sockets; pouch
protecting plates slightly developed with pouch closure of everted type (E-
BPC). Mouth extremely vertical.

DEScRIPTION. With exception of internasal spike, all head and body
ridges faint. Orbital ridge mildly accentuated with pronounced spine on
anterior border (holotype) or moderate (paratype). Opercular ridge extending

138 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

5 mm

Ficure 5. Micrognathus myersi Herald and Randall, Holotype; male 42.5 mm. SL
(CAS 13918); NW. Cocos Island, Guam. Drawing by L. Sabre.

over half of opercle. Pectoral cover plate with faint superior and inferior
ridges. Trunk and tail ridges with indentation between rings, and ridges
with finely toothed edge. Base of dorsal fin resting in slight ‘V.’ Color
light tan: holotype with indication of 4 pigment streaks extending downward
from opercle and dark area on side of snout just anterior to eye. Paratype
with indication of color bands formed by occasional dark spots: 7 bands
or bars on trunk and 17 on tail.

Discussion. Although the two type specimens are almost the same size,
the holotype snout length is much greater than that of the paratype, with
snout-in-head values being 2.36 and 2.7. If the other characters were not
so similar, one would be tempted to consider them as closely related but
separate species.

Short snouts are the mark of the subgenus Minyichthys as is shown even
better by the two known specimens of the genotype M. (Minyichthys)
brachyrhinus (snout-in-head 3.17). A comparison between the two species
of Minyichthys follows:

M. (M.) brachyrhinus M. (M.) myersi
Dorsal fin rays 23-25 29-31
Rings covered 1%-1%+5%- 2%-3+6%4-
by dorsal 6%4=74-8%4 74%2=9-10%
Trunk rings 19-21 19
Tail rings 37-39 40-41
Snout-in-head 3.17-3.18 2.3-2.7
Max. known size 31 mm. 42.5 mm.
Distribution Oahu; Ticao I. Guam

Philippines

This new species is named in honor of Stanford faculty member Dr.
George Myers (retired) who has been the major professor for many of
today’s ichthyologists.

VoL. XXXIX] HERALD & RANDALL: PIPEFISHES 139

5 mm

Ficure 6. Micrognathus myersi Herald and Randall, Paratype; female 41 mm. SL
(BPBM 7579); south of Uruno Point, Guam. Drawing by L. Sabre.

COMMENTS ON OTHER MICROGNATHUS

Since the senior author’s Micrognathus study of 1953, there have been
two additional new species, one in the Atlantic and one in the Pacific. In
1964 Gilbert Whitley described Micrognathus boothae from Australia’s Lord
Howe Island (off New South Wales). This species is closely related to
M. brocki discovered at Bikini Atoll in the Marshall Islands, about 2570
miles north of Lord Howe. The two species differ mainly in the shorter
snout of M. boothae along with its 42 rather than 37 tail rings. Three
specimens are known for the two species: the female holotypes of each
plus a second specimen of M. brocki captured by the junior author in 1968
at Ishigaki in the Ryukyu Islands. This 82 mm. female (BPBM 8755;
20-70 feet) represents a westward range extension of about 2800 miles.
When more material is available of both species, it is possible that the two
may prove identical.

In April and May 1971 the junior author collected four heavily-banded
specimens of Micrognathus edmonsoni in the Marquesas. Three were taken
at Nuka Hiva at a depth of 70 feet (BPBM 10857, 71 and 70 mm. SL,
and CAS 13977, 62 mm.) and one was captured at 115 feet in Vaitahu Bay
at Tahuata (BPBM 11936, 72 mm. SL). This is startling because previously
this species had been considered a rare Hawaiian endemic. Although it was
described more than 42 years ago (1930) only 7 other specimens are known
from 3 localities on Oahu and Maui. This new locality extends the range
about 2500 miles southward from the Hawaiian archipelago. It is entirely
possible that this species is one of the deeper water syngnathids, and thus
has escaped collections made by non-diving ichthyologists.

Another problem deals with a pipefish originally described as [chthyocampus
annulatus Macleay 1878. The two type specimens are actually Micrognathus
brevirostris as shown by reexamination in 1971 by Dr. John R. Paxton

140 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

at the Australian Museum. Unfortunately the artist used a specimen of
Yozia species to make the illustration for the original description, and in
some unknown way this fact escaped the notice of the author.

LITERATURE CITED

HERALD, EArt S.

1953. Family Syngnathidae: Pipefishes. In: Leonard Schultz et al. Fishes of the
Marshall and Marianas Islands. United States National Museum, Bulletin
no. 202, pp. 231-278, 9 figs.

1959. From pipefish to seahorse—a study of phylogenetic relationships. Proceedings
of the California Academy of Sciences, ser. 4, vol. 29, no. 13, pp. 465-473,
3 figs.

1969. A new pipefish from the Virgin Islands, Micrognathus dawsoni. Occasional
Papers of the California Academy of Sciences, no. 73, 4 pp., 1 fig.

RANDALL, JOHN E.

1962. Fish service stations. Sea Frontiers, vol. 8, no. 1, pp. 47.

1970. Easter Island, an ichthyological expedition. Oceans, vol. 3, no. 3, pp. 48-59.
RANDALL, JOHN E., AND E. HELFMAN

In Press. Dipractacanthus xanthurus, a cleaner wrasse from the Palau Islands with
notes on other cleaning fishes. Tropical Fish Hobbyist.

SmMitH, J. L. B.

1963. Fishes of the family Syngnathidae from the Red Sea and western Indian

Ocean. Rhodes University Ichthyological Bulletin no. 27, pp. 515-543.

PROCEEDINGS

OF THE

CALIFORNIA ACADEMY OF SCIENCES

FOURTH SERIES

Vol. XXXIX, No. 12, pp. 141-184; 11 figs. July 9, 1973

THE GENUS MECAS LECONTE
(COLEOPTERA: CERAMBYCIDAE)

By
John A. Chemsak and E. G. Linsley
University of California, Berkeley

The genus Mecas is a group of lamiines generally distributed from Guatemala
to southcentral Canada. A single species occurs in North America west of the
Rocky Mountains, but none are known from northeastern North America. Based
upon present knowledge, most of the species are associated with weedy com-
posites. A few of them are also lampyrid and/or cantharid and lycid mimics.
Although the genus has been recently revised (Breuning, 1955), available in-
formation and material have necessitated a further reclassification of the group.

CLASSIFICATION AND NOMENCLATURE

The first named species of Mecas as currently recognized (M. pergrata, M.
cana, M. cinerea) were assigned by their authors to the genus Saperda Fabricius
(Say, 1824; Newman, 1840). LeConte (1852, 1859a) transferred S. pergrata
to Stenostola Mulsant, a generic name he also used for his ‘saturnina’ (LeConte,
1859a), but left ‘cana’, which he did not know, in Saperda (he had no reason
to refer to ‘cinerea’ which was from Mexico). In 1847, Haldemen described
‘femoralis’ in the genus Phytoecia Mulsant, which LeConte (1852) made the
monobasic type of his genus Mecas. Twenty-one years later, LeConte (1873a)
added a second species, M. marginella, and defined the genus (1873b) in such
a manner that all of the above species could be included, although he did not
mention them by name. Unfortunately, Lacordaire (1872) overlooked Mecas
which would have fallen in his Tribe IV (Phytoecides), although he included the
related Dylobolus Thomson (which Bates later regarded as a synonym of Mecas)
in his Groupe IV (Arenicides) and Pannychis Thomson (which we regard as a

[141]

142 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

subgenus of Mecas) in his Groupe III (Amphionychides). LeConte’s expanded
concept of the genus was first applied to the North American species by Horn
(1878) and to the Mexican species by Bates (1881). Casey (1913) provided a
key to the species in his collection and Breuning (1955) published a revision of
the genus. However, this revision is incomplete, since three of the species, M.
inornata of authors, M. cineracea Casey, and M. bicallosa Martin were omitted.
These were incorrectly transferred by Breuning to the genus Saperda, although
their published characters (including the cleft or toothed claws in both sexes)
and recorded host plants (weedy composites) are typical of the species currently
and historically referred to Mecas (the species of Saperda are wood borers).

Although the evolution of the generic concept of Mecas until the revision by
Breuning was fairly straightforward, there has been considerable uncertainty
regarding the identity of some of the species. The most serious confusion in-
volves the name ‘Saperda inornata’ described from Missouri Territory by Say
(1824), who used the broad concept of Saperda current in his time. LeConte
(1852), not knowing what species Say had before him, included it in his treat-
ment of Saperda, quoting from Say’s description as he did for other species
which he had not identified. Obviously uncertain, he speculated that it might
be the male of his S. concolor from Sante Fe, New Mexico, which he described
in the same paper (LeConte, 1852). However, he apparently decided later
that ‘imornata Say’ belonged to the genus Mecas, and the specimens standing
under that name in his collection belong to this genus. Further, the name ‘in-
ornata Say’ does not appear in his key to the species of Saperda (LeConte,
1873a). Horn (1878), in his key to the species of Mecas, included the name
‘inornata, applying it to the species previously described by LeConte as ‘satur-
nina, which Horn regarded as a synonym of ‘inornata.’ However, Blanchard
(1887) found two species differing in claw structure standing under the name
‘saturnina’ in his collection. These had not been differentiated in Horn’s Key.
This discovery prompted Horn (1888) to apply the name ‘saturnina’ LeConte
to the species with the claws moderately deeply cleft with the inner division
lobe-like. On this basis of distinction Gahan (1888) placed Saperda cinerea
Newman and Mecas senescens Bates as synonyms of Mecas inornata (Say) and
Mecas saturnina (LeConte) as a synonym of Mecas cana (Newman). We agree
that M. saturnina and M. cana are conspecific, although in the material before
us the two appear to be allopatric. Mecas cinerea (Newman) and M. senescens
Bates also appear to be conspecific, but are not closely related to any species
from the United States and thus could not under any circumstances be called
‘inornata Say.’ Although various species of Mecas have been treated taxonomi-
cally in the intervening years as ‘M. inornata Say’ (see synonymical bibiog-
raphies), the species which most closely fits Say’s very brief description is M.
confusa (described below) or, possibly, M. cineracea Casey.

VoL. XXXIX] CHEMSAK AND LINSLEY: MECAS 143

In 1924, Martin named a species MW. bicallosa on the assumption that ‘inornata
Say’ was Saperda in the modern sense and a senior synonym of S. concolor Le-
Conte (see above), although the arguments advanced for this decision were not
conclusive (Martin also assumed that he was formally naming the species that
previously had been called M. inornata; however, the species before him was from
the Great Basin and the ranges accorded to M. inornata by Horn and Blanchard
are to the east of this area). Martin’s placement of S. concolor LeConte as a
synonym of S. inornata Say was accepted by Breuning (1952) and Nord and
Knight (1970). These latter appealed to the International Commission on
Zoological Nomenclature to use its plenary powers to designate as neotype of
Saperda inornata Say the type of Saperda concolor unicolor Felt and Joutel,
and place ‘inornata’ on the Official List of Specific Names in Zoology. While
such an action would eliminate some nomenclatural confusion in the genus Mecas,
its impact on the literature of the North American Saperda will be unfortunate.
Proposing to relegate the name ‘inornata Say’ to the list of nomina dubia might
have been a preferable solution to the problem.

DISTRIBUTION

The known species of Mecas occur in the area from western Canada to south-
eastern United States, Mexico, and Guatemala. Of the 15 species recognized by
us, five are restricted to northern and central Mexico and another ranges into
Guatemala; six occur in both the United States and Mexico and three are thus
far known only from the United States.

A single species, M. bicallosa, is found west of the Rocky Mountains, occupy-
ing the Great Basin and extending into British Columbia and northern Baja
California. Most of the species occurring in the United States are found in
either the southeastern and/or southwestern portions of the country. Only three
species, M. femoralis, M. marginella, and M. confusa, are not yet known from
Mexico but probably only M. femoralis does not occur there. The most widely
distributed species, M. rotundicollis, ranges from Costa Rica to Kansas ana
Arizona. Of the remaining four species common to both countries, M. cana
saturnina, M. pergrata, and M. cineracea apparently occur only in northeastern
Mexico but are more widely distributed in the United States. The other species,
M. menthae, ranges from Distrito Federal north along the western slope of
the Sierra Madre to Arizona and New Mexico. Five of the six Mexican species,
M. sericea, M. humeralis, M. cinerea, M. cirrosa, and M. ambigena are apparently
largely restricted to central Mexico from Chihuahua and San Luis Potosi to
Puebla. The sixth, WM. obereoides, extends into Guatemala.

In many respects the distribution of Mecas is comparable to that of Elytro-
leptus (Linsley, 1962; Chemsak and Linsley, 1965). Of the 17 species of Elytro-
leptus, 9 are known only from Mexico, 7 from the United States and Mexico,

144 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

and one from the United States only. The majority of the species, as in Mecas,
are found in central and northern Mexico and southwestern United States. Since
nothing is known of the biology of the species of Elytroleptus, the significance
of the distributional similarity, if any, is not evident.

FORM AND COLORATION

Most of the species of Mecas are concolorous with the integument black and
densely clothed with pale recumbent pubescence. A few of the species, M. fem-
oralis, M. pergrata, and M. cinerea frequently or always possess reddish femora
and the latter two often have the elytra and/or pronotum partly reddish. Some
of the members of the genus are distinctive by the narrow, densely pubescent
white bands on the suture and epipleurae of the elytra.

The most striking divergence from the typical appearance is to be found in
M. sericea, M. rotundicollis, and M. obereoides. The first of these, M. sericea,
is definitely lycid-like in aspect usually possessing two lateral dark bands at
the base and apex of the elytra. The basal band may be reduced or absent
and occasionally both bands are lacking. Since this species is mimetic, the
variation in color is probably an expression of resemblance to different lycid
models within the range of the species. Lycus sallei Gorham is a possible model
with two dark elytral bands and we have another similarly marked Lycus from
Sinaloa and Colima.

Mecas rotundicollis is unquestionably a lampyrid mimic throughout most
of its range, although in some areas a cantharid may be the model. As is the case
in M. sericea, M. rotundicollis apparently mimics different species of models
in different parts of its range. Variation is expressed primarily by the presence
or absence of yellow pubescence on the apical abdominal sternites and yellow
pubescent bands on the suture and epipleurae of the elytra. We have been unable
to find a geographical trend in these characteristics but most of the available
specimens from Arizona lack the yellow pubescence.

The third species, M. obereoides, may be involved in a mimetic ring with a
cantharid model. Very little variation in color is expressed in this species but
it is one of the most distinctive in the genus. Field studies will be necessary to
confirm involvement with mimicry.

The remaining species of Mecas are all quite similar in form and coloration.
Differences between species involve such characteristics as tarsal claw structure,
number of glabrous calluses on the pronotum, and relative lengths of antennal
segments.

BIOLOGY

Little has been recorded on the biology of species of Mecas. The most complete
accounts known to us are those of Baerg (1921) and Stride and Warwick (1962).
Baerg’s report describes injury to Jerusalem artichokes in Arkansas and probably

VoL. XXXIX] CHEMSAK AND LINSLEY: MECAS 145

refers to M. cana saturnina (LeConte) or the species named below as Mecas
confusa, new species. Baerg’s account, under the name “Mecas inornata Say”
is as follows:

This beetle, half an inch long, of a light gray color, is a girdler that attacks arti-
choke (Helianthus tuberosus). The beetles begin ovipositing early in July. The females
when laying eggs girdle the main stem about six inches from the top. Two girdles are
made, about 1-144 inches apart. Immediately above the lower girdle is the egg punc-
ture. This is exactly similar to the method followed by the Raspberry cane girdler
(Oberea bimaculata). The girdles are not clean cuts such as we find in woody plants,
but rather a series of holes encircling the stem. Apparently one female will deposit
in a large number of plants. In spite of the fact that only a few beetles could be
located, practically all the plants in the field were attacked in the course of a few
days.

As a result of the injury, the leader in the plant dies and the plant develops a
bushy type of branching.

The young larvae upon hatching begin to feed between the girdles and later
proceed towards the base of the plant. They confine their injury largely to the pith.
Apparently under certain weather conditions the artichoke is not well fitted as a
host plant. In only one out of four or five plants showing egg punctures was there
a full grown larva. In most of the other plants the larva had begun to feed and
some time later died, presumably it had been injured by the growing stalk.

The larvae attain full growth, that is about seven-eights of an inch in length,
some time in November. At this time the larvae are found at the very base of the
stalk, about two inches below the surface of the ground, in an enlargement of the
tunnel which has been padded with fine bits of pulp.

The pupal stage has not been observed but since the adults appear early in July,
the larvae will presumably pupate some time in May or early in June.

It seems reasonable to assume that this species will attack most of the species in
the genus Helianthus. None of these were near the artichokes, and no data have been
secured. The only host plant other than artichoke that could be located is the
common ragweed (Ambrosia artemisiifolia) .

The observations of Stride and Warwick were made on M. saturnina in Aus-
tralia where that species had been introduced as a biological control agent for
Xanthium (Wilson, 1960). The habits as they reported are almost identical with
Baerg’s account and an additional observation was the plugging of the oviposition
hole with a gummy substance after the single egg had been laid. Stride and
Warwick advance a hypothesis whereby the double-girdling behavior of M.
saturnina may be regarded as a device originally evolved to permit the use of
succulent green shoots of otherwise woody plants as food for cerambycid larvae.
Presumably, it has been retained in M. saturnina because it promotes advan-
tageous changes in the herbaceous host plant attacked, possibly increased
pithiness.

Earlier reports also refer to stem- and root-boring habits of Mecas (Riley,
1880; Beutenmuller, 1896; Leng and Hamilton, 1896). Most aspects of adult be-
havior are lacking from the older literature but Townsend (1884) reported that

146 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

‘Mecas inornata’ takes wing and flies away when it observes someone approach-
ing, but drops to the ground and feigns death when unexpectedly disturbed.

Adults of M. menthae, new species were found in numbers by their collectors
on the upper leaves of the mint, Agastache, during the day.

Host RELATIONSHIPS

Precise host data for most species of Mecas are lacking. In part this results
from uncertainty regarding the identity of the species of Mecas associated with
published host records and in part from the fact that most records, both pub-
lished and unpublished, are based upon collections of adults from plants and
not upon reared material. Nevertheless, there is an interesting consistency among
the records that are available. With very few exceptions the plants involved
are weedy, herbaceous composites which have special chemical characteristics
expressed in terms of aromatic, medicinal, irritant, or toxic properties. The
list of known or suspected hosts based upon field collections, with notations of
some of their properties as reported by Blake (1951) and Kingsbury (1967) is
as follows:

COMPOSITAE

Ambrosia (Ragweed).
A. artemisiifolia Linnaeus (Common Ragweed). One of the most widespread causes
of hayfever.
Artemisia (Sagebrush).
A. tridentata Nuttall (Big Sage). Used medicinally by Indians and early white
settlers in the West; a hay fever plant; toxic to livestock if eaten in excess.
Aster (Aster).
A. tanacetifolius Humboldt, Bonpland, & Kunth (Tansyleaf Aster). Used medic-
inally by the Indians; some species of aster absorb Selenium and become toxic.
Baileya (Baileya).
B. multiradiata Harvey & Gray (Desert Marigold). Causes mortality in sheep, par-
ticularly, but not exclusively, on over-grazed land.
Gaillardia (Blanket-flower) .
G. pulchella Fougeroux de Bondaroy. A related species, G. pinnatifida Torrey.
was used by the Hopi Indians as a diuretic.
Guardiola (Guardiola).
G. tulocarpus Gray.
Helenium (Sneezeweed).
H. hoopesti Gray (Orange Sneezeweed). Contains a toxic glucoside (dugaldin)
which causes spewing sickness in sheep.
H. microcephalum WDeCandolle (Sneezeweed). Toxic to livestock.
H. tenuifolium Nuttall (Bitter Sneezeweed). Toxic to livestock and suspected of
poisoning humans.
Helianthus (Suntlower).
H. annuus Linnaeus (Common Sunflower). Toxic to cattle in large amounts; the
seeds are edible.
H. tuberosus Linnaeus (Jerusalem Artichoke). Roots edible.

VoL. XXXIX] CHEMSAK AND LINSLEY: MECAS 147

Verbesina (Crown-beard).
V. encelioides (Cavanilles) Bentham & Hooker (Golden Crownbeard). Used by
Indians and White pioneers in the West for treatment of boils and skin disease;
Hopis reported to bathe in water in which plant has been soaked to relieve pain
of spider bites.
Xanthium (Cocklebur).

X. spinosum Linnaeus (Spiny Cocklebur). Seeds and seedlings contain a glucoside
(Xanthostrumarin) poisonous to swine and poultry.

LABIATAE

Agastache (Giant-hyssop)
A. species (Horsemint).

Genus Mecas LeConte

Mecas LEContTE, 1852, Jour. Acad. Sci. Philadelphia, vol. 2, no. 2, p. 155; 1873, Smith-
sonian Misc. Coll., no. 265, p. 347; Horn, 1878, Trans. American Ent. Soc., vol. 7, p. 44;
Bates. 1881, Biologia Centrali-Americana, Coleoptera, vol. 5, p. 203; LeContTEe and
Horn, 1883, Smithsonian Misc. Coll., vol. 507, p. 332; Lrenc and Hamiurton, 1896,
Trans. American Ent. Soc., vol. 23, pp. 151, 152; Casry, 1913, Memoirs on the
Coleoptera, vol. 4, p. 360; BREUNING, 1955, Mem. Soc. Roy. Ent. Belgique, vol. 27, p. 138.

Form elongate, usually parallel. Head with front convex, interantennal area
usually concave; mandibles rather short, apices curved, acute; palpi slender,
maxillary pair longer than labial; eyes rather small, finely faceted, deeply emar-
ginate, upper lobe small; antennae usually slender, sparsely or densely fringed
with long hairs beneath, particularly on basal segments; third segment usually
longer than first, fourth subequal to or shorter than third, outer segments short or
long. Pronotum wider than long, sides usually rounded; disk variably pubescent,
often with glabrous calluses; prosternum short, intercoxal process narrow,
expanded at apex, coxal cavities closed behind; mesosternum with coxal cavities
open; metasternum with episternum broad in front, narrowing behind. Legs
short; intermediate tibiae with a dorsal sinus; tarsal claws bifid. Abdomen
normally segmented; last sternite deeply impressed in the male, linearly im-
pressed in the female.

Type spectEs. Phytoecia femoralis Haldeman (monobasic).

This genus is distinctive from others in its tribe by the proportions of the
antennal segments, the frequent presence of dorsal calluses on the pronotum, the
shape of the metepisternum, and by the nature of the impressions of the last
abdominal sternite, which are concave in the male and linear in the female. Many
of the species resemble Saperda but the bifid claws will readily separate them.

There are at present 15 known species of Mecas assignable to three subgenera,
all occurring in the New World.

148

2(1).

4(3).

5(4).

6(5).

7(6).

8(4).

CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

KEY TO THE KNOWN SPECIES OF THE GENUS Mecas

Pronotum with sides rounded or subparallel; elytra not expanded apically

behind! middle; appearance’ not ly ciionn) 22 Ea 2
Pronotum with sides obtusely produced at middle; elytra expanding slightly

toward apices; integument yellow and black; appearance lycid-like. Mecas

(Pannychis). 14-20 mm. Chihuahua to Veracruz _..__ SS M. sericea

Pronotum not densely fringed with short, erect, golden pubescence, erect hairs
moderately long; elytral apices broadly rounded or rotundate-truncate;
abdomen with pubescence of sternites unicolorous. Mecas, sensu stricto. 3

Pronotum densely fringed with short, erect, golden pubescence with scattered
long setae intermixed, dorsal surface with a pair of longitudinal vittae on
each side of middle composed of short, appressed, golden pubescence;
integument usually concolorous golden yellow, less commonly mottled
or vittate with black, rarely wholly black; elytral apices angulate, obliquely
truncate or emarginate; abdomen often with last three sternites margined
laterally with longitudinal bands of yellowish-white pubescence suggesting
luminescent organs of a lampyrid. Mecas (Dylobolus). 8-14 mm. South-
western WUmited states! to) Costar Rica ses ee eee M. rotundicollis

Pronotal pubescence dense, appressed, obscuring surface, intact or broken by
well defined black polished callosites and usually also a median impunctate
area. On ISK... i225 22 oe eee 4

Pronotal pubescence very sparse, erect, not obscuring surface which has five
tubercles, one median, two antemedian, and two lateral, the surface red to
yellow, concolorous or margined laterally with black and/or with black
spots on the median or median and lateral tubercles. 8-10 mm. Sinaloa and

San) JouismPotosiatoy Guatemalae = ee ee M. obereoides
Pronotum with pubescence intact, not interrupted by polished black callosities. 5
Pronotum with at least two polished black callosities and usually a median

elongate impunctate “area on) isk) eee 8

Pronotum and elytra with concolorous pubescence which obscures the surface;
Stemnum) juniformly, densely, pubescent) = ———————E—EEs 6

Pronotum and elytra with longitudinal bands of dense, appressed, often yel-
lowish pubescence at middle and sides, remaining pubescence not completely
obscuring surface; sternum margined with a row of dense, appressed,
yellowish pubescence. Length, 6.5-8 mm. Southeastern United States to

IN GW, VICxXI COM == 28 ee ee eee M. marginella
Tarsal claws with inner tooth much smaller than outer one. Smaller species,
6=0iimm-ain length: (fig. Ub), 2-2 Ee ee if
Tarsal claws with inner tooth almost as long as outer one. Length, 10-14 mm.
INansasMto.wMexas) (lig. 1a)).p «tet Eee M. confusa
Femora always reddish; pubescence finer, not completely obscuring surface.
Length, 6-8 mm. Southeastern United States. SS M. femoralis

Femora always black; pubescence thick, obscuring surface. Length, 6-11 mm.
Southeastern United States to Arizona, Colorado, and northeastern Mexico,
etait AS ata cote he le ee M. cineracea

ANCA: <2 82... 5 eee 9
Pronotum with two rounded glabrous calluses in addition to elongate median
imnipunctate area. ee EEE EE = UG)

VoL. XXXIX] CHEMSAK AND LINSLEY: MECAS 149

a b Cc

FicurE 1. Some variations in the form of the tarsal claws in the genus Mecas.

9(8). Form elongate, elytra about 3 times as long as broad; antennae at least as long
as body; elytra uniformly pubescent, margins and suture not distinctly pubes-

cent. Length, 8-13 mm. San Luis Potosi to Morelos and Nayarit. __. M. cinerea
Form rather stout, short, elytra about 244 times as long as broad; antennae
shorter than body; elytra with margins and suture densely pale pubescent.
Length, 6-12 mm. Great Plains to southeastern United States, New

Mexico; and northeastern: Wiexico. 2 == ene M. pergrata
10(8). Tarsal claws with inner tooth short, obtuse, lobe-like (fig. Ic), 0. 11
Marcoleclaws with inner tooth acute: spine=likes eee ee 12

11(10). Antennae shorter than body, densely clothed beneath with long curved hairs,
segments robust, flattened; elytra uniformly gray pubescent. Length, 10.5-

AS emiinen Gea Wat oven! @Uuereter@: se eee eee M. cirrosa
Antennae longer than body, sparsely clothed beneath with long erect hairs,
segments slender; elytra broadly clothed with brownish pubescence along

epipleurae; humeri glabrous. Length, 10.5-13 mm. Jalisco. —___ M. humeralis
12(10). Antennae much shorter than body, outer segments short. — 13
Antennae at least as long as body, outer segments elongate. = ti«i2“SH

13(12). Appressed pubescence gray, uniform on elytra; pronotum ieee Sains
punctate; elytra lacking long erect dark hairs over apical one half. Length,
10-15 mm. Washington to northern Baja California and Colorado. __ M. bicallosa
Appressed pubescence brownish, denser on margins and suture of elytra;
pronotum coarsely, confluently punctate; elytra densely clothed with dark
erect hairs over apical one half. Length, 9-10 mm. Durango to Mexico.
tet ee, | a a ee ee eee M. ambigena
14(12). Elytra sparsely, separately punctate, lacking long erect hairs; pronotal calluses
Siiallbemediansampunctates areas Via Sle. ee 15
Elytra coarsely, contiguously punctate, densely clothed with long erect hairs;
pronotal calluses large, median impunctate area distinct. Length, 8-13 mm.

AmizonaeiomNayanitrandmDistrito) Hecdetals seems sess ss een M. menthae
15(14). Elytra with distinct longitudinal pubescent bands along margins and suture.
encth-— 10-16) mma shloriday = ee M. cana cana

Elytra uniformly grayish or yellowish pubescent, suture and margins without
pubescent bands. Length, 10-16 mm. Eastern United States to South
Dakotasangenortheastenmel lexi Cp eee M. cana saturnina

Subgenus Pannychis Thomson

Pannychis THomson, 1864, Systema cerambycidarum, p. 127;1868, Physis, vol. 2, p. 197;
LAcorDaIRE, 1872, Genera des coléoptéres, vol. 9, pp. 882, 889; BaTEs, 1881, Biologia
Centrali-Americana, Coleoptera, vol. 5, p. 205; GimMmour, 1962, Rev. Biol. Trop.,
vol. 10, pp. 125, 137.

150 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

XG
: Ss

Ficure 2, Mecas (Pannychis) sericea (Thomson), ¢.

VoL. XXXIX] CHEMSAK AND LINSLEY: MECAS 151

Form moderate to large, lycid-like. Antennae short, not extending beyond
third abdominal segment, segments not gray pubescent. Pronotum broader than
long, sides obtusely produced at middle; disk convex, sparsely punctate, pubes-
cence not obscuring surface. Elytra slightly expanded behind middle, disk
costate, pubescence bicolored. Legs with tarsal claws with long inner tooth.
Abdomen with last sternite impressed at apex in females.

Type SPECIES. Pannychis sericeus Thomson (monobasic).

The lycid-like form and coloration and the obtusely produced pronotal sides
make this subgenus very distinctive. A single species is known from Mexico.

Mecas (Pannychis) sericea (Thomson).

(Figures 2, 3.)

Pannychis sericeus THOMSON, 1864, Systema cerambycidarum, p. 127; 1868, Physis, vol. 2,
p. 197; Bates, 1881, Biologia Centrali-Americana, Coleoptera, vol. 5, p. 205; Gimrmour,
1962, Rev. Biol. Trop., vol. 10, p. 137.

Pannychis ducalis BATEs, 1881, Biologia Centrali-Americana, Coleoptera, vol. 5, p. 205;
Gitmour, 1962, Rev. Biol. Trop., vol. 10, p. 137. New synonymy.

Mate. Form moderate sized to rather large, elytra slightly expanding toward
apices; integument yellowish, antennae black, legs variable, tibiae usually black,
femora often partially black, head usually with 3 black bands over vertex and
behind eyes, pronotum usually with 3 longitudinal bands joining at base and
broad black bands at sides beneath, elytra yellow or with basal and broad apical
black bands, basal bands often reduced or lacking, beneath variably colored.
Head with front convex, coarsely, shallowly punctate; interantennal area con-
cave, median line deep; vertex shallowly punctate; pubescence dense, appressed
on cheeks, sparse, short, and suberect on front; antennae extending to about
third abdominal segment, segments all clothed with very short, dark, depressed
pubescence, basal segments with a few long erect setae beneath, third segment
subequal in length to first, fourth shorter than third, remaining segments grad-
ually decreasing in length, eleventh appendiculate, segments from fifth with
vague longitudinal poriferous areas. Pronotum broader than long, sides obtusely
produced at middle; disk convex, almost impunctate; pubescence dense, golden,
appressed at sides, a little sparser on disk with a few long, erect hairs inter-
spersed; prosternum sparsely pubescent; meso- and metasternum moderately
densely pubescent, distinctly punctate at sides. Elytra about 3 times as long
as broad, slightly expanding behind middle; each elytron lightly bicostate,
epipleurae vertical; punctures at base coarse, dense, becoming fine and sparser
toward apex; pubescence short, dense, subdepressed, colored as integument,
longer suberect hairs present basally; apices rounded. Legs finely pubescent;
tarsal claws with inner tooth slightly shorter than outer. Abdomen finely
moderately pubescent; last sternite deeply impressed for most of its length.
Length, 14-19 mm.

FEMALE. Form more robust, elytra slightly more expanded behind middle.

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152

VoL. XXXIX] CHEMSAK AND LINSLEY: MECAS 153

Antennae extending to about first abdominal segment. Abdomen with last
sternite linearly impressed, concave at apex, apex broadly V-shaped. Length,
14-20 mm.

TYPE LocALITy. Of ‘sericeus,’ Mexico; ‘ducalis,’ Orizaba, Mexico.

RANGE. Chihuahua, Mexico to Veracruz (fig. 3).

FLIGHT PERIOD. July to October.

Remarks. This striking species undoubtedly is one of the lycid mimics.
It is variable in coloration and, as is found among members of Elytroleptus
(Chemsak and Linsley, 1965), all yellow individuals occur as well as ones with
only the apices of the elytra black or with both the apices and base black. It
would not be surprising to encounter individuals with all black elytra.

MATERIAL EXAMINED. MEXICO. Chihuahua: 1 2, 3 miles SE. of Temoris,
VII-25-69 (R. C. Gardner, C. S. Glaser, T. A. Sears); Jalisco: 1 ¢, 10 miles
SW. of Tecalitlan, X-10-64 (A. E. Michelbacher); 1 ¢, Cuidad Guzman, Jalisco,
IX-15-63 (M. C. Colorado); Morelos: 1 4, Tequesquitengo, VII-15-61 (R.
& K. Dreisbach); Michoacan: 1 2, near Morelia, IX-5-52 (G. H. Dieke);
Guerrero: 1 2, Iguala, VII-21-62 (H. E. Milliron); 1 2, 10 miles N. of Mercula,
VIII-26-58 (E. L. Mockford); 1 6, Thaxmalac, IX-22-42 (W. F. Fosberg) ;
Puebla: 1 6, 19 miles NW. of Cacaloapan, VII-30-65 (W. A. Foster).

Subgenus Dylobolus Thomson

Dylobolus THomson, 1868, Physis, vol. 2, p. 195; LAcorpatrE, 1872, Genera des coléopteéres,
vol. 9, pp. 897, 900.

Form slender, elongate. Antennae slender, third segment slightly curved.
Pronotum with sides rounded, disk densely fringed with short, erect, golden
pubescence. Elytra with apices angulate, usually obliquely emarginate. Legs
with tarsal claws with inner tooth almost as long as outer one. Abdomen
frequently with yellowish appressed pubescence at sides of apical sternites.

Type species. Dylobolus rotundicollis Thomson (monobasic).

This subgenus differs from the others by the pubescence of the pronotum
and emarginate or truncate elytral apices. The single known species is a
lampyrid mimic.

Mecas (Dylobolus) rotundicollis (Thomson).
(Figures 4, 5, 6.)

Dylobolus rotundicollis THomson, 1868, Physis, vol. 2, p. 196.

Mecas rotundicollis, Bates, 1881, Biologia Centrali-Americana, Coleoptera, vol. 5, p. 205;
BREUNING, 1955, Mem. Soc. Roy. Ent. Belgique, vol. 27, p. 148.

Mecas ruficollis Horn, 1878, Trans. American Ent. Soc., vol. 7, p. 44; Bates, 1881, Biologia
Centrali-Americana, Coleoptera, vol. 5, p. 205; Lenc and Hamirton 1896, Trans.
American Ent. Soc., vol. 23, pp. 152; CAsEy, 1913, Memoirs on the Coleoptera, vol. 4,
Das02:

Mecas ruficollis morpha mediomaculata BREUNING, 1955, Mem. Soc. Roy. Ent. Belgique,
vol. 27, p. 149. New synonymy.

154 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Ficure 4. Mecas (Dylobolus) rotundicollis (Thomson), ¢.

VoL. XXXIX] CHEMSAK AND LINSLEY: MECAS 155

Mecas rotundicollis morpha ruficollis, BREUNING, 1955, Mem. Soc. Roy. Belgique, vol. 27,
p. 149.

Mecas laticeps Bates, 1881, Biologia Centrali-Americana, Coleoptera, vol. 5, p. 204;
BrREUNING, 1955; Mem. Soc. Roy. Ent. Belgique, vol. 27, p. 151. New synonymy.

Mecas laticeps morpha sutureflava BREUNING, 1955, Mem. Soc. Roy. Ent. Belgique, vol. 27,
p. 151. New synonymy.

Mecas laticeps morpha mediopunctata BREUNING, 1955, Mem. Soc. Roy. Ent. Belgique, vol. 27,
p. 151. New synonymy.

Mecas mexicana BATES, 1881, Biologia Centrali-Americana, Coleoptera, vol. 5, p. 204.

Mecas rotundicollis morpha mexicana, BREUNING, 1955, Mem. Soc. Roy. Ent. Belgique,
vol. 27, p. 149.

Mecas vitticollis CAsEY, 1913, Memoirs on the Coleoptera, vol. 4, p. 362.

Mecas laticeps morpha vitticollis, BREUNING, 1955, Mem. Soc. Roy. Ent. Belgique, vol. 27,
Deez.

Mate. Form moderate sized, elongate, sides parallel; color black, head
and pronotum orange, usually with dark spots or bands, legs often orange,
thoracic sterna often orange, abdomen usually with broad bands of yellow
appressed pubescence at sides of last three sternites, elytra frequently with
narrow bands of appressed yellowish pubescence down suture and epipleurae.
Head rather small; front convex, median line extending from clypeus to neck;
interantennal area concave; vertex moderately coarsely, densely punctate;
pubescence dense, yellowish, appressed with short, dark, erect hairs numerously
interspersed; antennae shorter than body, scape finely, very densely punctate,
third segment longer than first, fourth shorter than third, fifth subequal to first,
segments from sixth gradually decreasing in length, scape rather densely clothed
with short subdepressed hairs, underside of segments densely clothed with short,
pale, appressed pubescence, basal segments with a few long erect hairs beneath.
Pronotum usually broader than long, sides rounded; disk convex, sparsely to
rather densely punctate; pubescence usually dense, consisting of short, dense,
subappressed, longitudinal bands, one on each side of middle and at lateral
margins, longer erect hairs numerously interspersed; prosternum densely pubes-
cent; meso- and metasternum finely, densely punctate at middle, coarsely at
sides, pubescence dense. Elytra over three times longer than broad; suture
and epipleurae usually with narrow bands of appressed yellow pubescence;
punctures rather coarse, dense, becoming finer and sparser toward apex; surface
clothed with fine appressed pubescence, short, recurved hairs numerously inter-
spersed, apices obliquely truncate. Legs finely, densely pubescent; tarsal claws
with inner tooth almost as long as outer. Abdomen elongate, densely pubes-
cent; last three sternites usually clothed with broad yellow bands at sides; last
sternite deeply impressed for most of its length. Length, 9-16 mm.

FEMALE. Form similar, more robust. Antennae slightly shorter than in male.
Abdomen with last sternite linearly impressed for its entire length; last tergite
strongely, obtusely conical at apex. Length, 10-19 mm.

156

CALIFORNIA

ACADEMY OF SCIENCES

[Proc. 4TH SER.

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Type LocaLity. Of ‘rotundicollis, Mexico; ‘ruficollis, Texas; ‘laticeps,’
Guanajuato, Mexico; ‘mexicana,’ Izucar, Mexico; ‘vitticollis, Durango City;
‘mediomaculata, Guerrerro; ‘sutureflava, Temax, Yucatan; ‘mediopunctata,’
Mexico.

RANGE. Oklahoma to Arizona, Texas, and south to Costa Rica (fig. 6).

FLIGHT PERIOD. May to December.

Host pLrants. Adults have been collected on flowers of Guardiola tulocarpa
(Compositae) and on Eysenhardtia polystachya (Leguminosae).

Remarks. This species is a lampyrid mimic, resembling different models in
different parts of its range, as is evident by the polychromatism expressed in the
specimens at hand. These color differences are expressed especially in the presence
or absence of yellowish longitudinal sutural and epipleural bands on the elytra and
in the yellowish apical sternites of the abdomen. There are varying combinations
of these characters but we have been unable to correlate them geographically,
although this might be possible with larger series of specimens and model-mimic
associations over the entire range.

In addition to the above differences, individuals also vary considerably in
size, color of head, pronotum, sternum, and legs.

MATERIAL EXAMINED. UNITED STATES. Oklahoma: 1 ¢, 1 2, Edmond,
VII-9-57 (D. Alexander); Arizona: 2 ¢4, Dry Canyon, Sands Ranch, SE.
end Whetstone Mts., Cochise Co., VII-10-52 (H. B. Leech, J. W. Green) ;
1 2, Tombstone, VIII-13-40 (E. S. Ross); 1 2, Madera Canyon, Pima Co.,
IX-1-54 (Menke & Stange); 1 6, 5 mi. W. Portal, Chiricahua Mts., VIII-
12-58 (P. Opler). Texas: 1 6, New Braunfels; 1 ¢, 1 2, San Antonio, May,
VI-11-36 (C. D. Orchard); 1 4, 13 mi. W. of San Marcos, Comal Co., VI-24-
25-61 (R. L. Westcott); 1 ¢, Brownsville; 1 ¢, Van Horn, V-23-32 (E. G.
Linsley). MEXICO. Tamaulipas: 1 ¢, Ciudad Victoria, VI-8-51 (P. D.
Hurd); 1 ¢, Rio Corona, 21.3 mi. N. of Cuidad Victoria, X-25-65 (G. E. Ball,
D. R. Whitehead). Nuevo Leon: 1 6,2 22, Monterrey, XI-30-65, X-12-52.
San Luis Potosi: 1 2°, El Huizache, VIII-22-61 (R. & K. Dreisbach). Veracruz:
3 66,1 2, Veracruz, XI-1-57, IX-24-61 (R. & K. Dreisbach). Chihuahua: 1 ¢,
3 mi. E. of Temoris, VIII-26-69 (Sears, Gardner, Glaser). Durango: 1 ?, Encino,
VII-27-47, 6200 ft. (Schramel); 14, 11 mi. W. of Durango, VIII-2-64, 7000 it.
(L. Kelton); 1 6, 7 mi. W. of Durango, VII-23-64 (W. R. M. Mason); 1 4, 25
mi. S. of Durango, VII-24-64 (L. Kelton); 1 6,2 22,8 mi.S. of Canutillo, VII-
9-51, on flowers of Guardiola tulocarpa (P.D. Hurd). Hidalgo: 1 4, Ajacuba, VI-
21-37 (M. A. Embury); 2 66,2 22, Pachuca, VI-15-35, VII-10-37 (Embury),
VII-31-35 (Embury); 1°, Zimapan, VII-14-68 (M. Wasbauer, J. Slansky). Dis-
trito Federal: 2 22, Temascaltepec, 1931 (G. B. Hinton). Nayarit: 6 6¢, 5
2 2, Tepic, IX-13-57 (R. & K. Dreisbach), IX-15-17-53 (B. Malkin), IX-24-47
(B. Malkin); 1 2, Campostella, IX-16-57 (R. & K. Dreisbach). Jalisco: 1%, 13

158 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH Serr.

SCALE
200 600 800 Jo0o MILES
= T om oS
400 600 800 1000 1200 1400 KILOMETERS

LAMBERT AZIMUTHAL EQUAL-AREA PROJECTION

Base map copyright 1961 by the University of Chicago. Reproduced by permission.

Ficure 6. Known occurrence of Mecas (Dylobolus) rotundicollis (Thomson).

VoL. XXXIX] CHEMSAK AND LINSLEY: MECAS 159

mi. N. of Chapala, VIII-1-63 (P. J. Spangler); 1¢, San Juan Lagos, VII-27-51,
on Eysenhardtia polystachya (P. D. Hurd); 1 2, Tlaquepaque, VI-28-45 (N.C.
H. Krauss); 1 2, Ajijic, VIT-28-54 (Cazier, Gertsch, Bradts); 1 6, 17 mi. SSW.
of Guadalajara, VIII-22-70 (M. & J. Wasbauer). Colima: 1 ¢, Pine zone, SE.
Slope Mt. Colima, XII-48 (E. S. Ross). Michoacan: 1 ¢, 5 km. W. of Zacapu,
VII-13-51 (H. E. Evans); 1 ¢, Morelia, [X-30-45; 1 ¢, 6 mi. NW. of Quiroga,
VII-11-63 (Parker, Stange); 1 4, 12, Quiroga, VII-15-56 (R. & K. Dreis-
bach); 2 6¢, 1 @%, Tuxpan, ITX-6-03 (McClendon), IX-18-57 (Scullen).
Morelos: 2 ¢¢,1 2, Hujintlan, VIII-22-56 (R. & K. Dreisbach); 1 2, Lake
Tequesquitengo, IX-13-57 (Schullen); 2 ¢4, Cuernavaca, X-02, X-29-57
(Dreisbach); 1 2, Morelos, Oct. Guerrero: 1 2, Rio Balsas (Wickham).
Puebla: 1 6, 3 mi. N. of Petalcingo, VIII-21-63 (Parker, Stange); 1 6, 55 mi.
S. of Acatlan, VII-30-63 (J. Doyen); 1 ¢, Tehuacan, VI-23-51 (Evans); 1 4,
19 mi. NW. of Cacaloapan, VII-30-63 (Foster). Oaxaca: 8 44,1 2, Oaxaca,
VII-20-37 (Embury); 2 64,1 2, Oaxaca, IX-13-20-47 (Malkin); 1 4,1 2,
Monte Alban, VIII-3-54 (P. & C. Vaurie) , X-12-63 (A. E. Michelbacher) ; 2 2 2,
18 mi. NW. of Totolapan, VII-28-63 (Doyen, Foster); 1 2, Temescal, VII-5
-65 (G. H. Nelson). Chiapas: 1 ?, Jct. Hwys. 190-195, VI-6-69 (H. F. How-
den); 3 64,2 22, San Jeronimo, Volcan Tacana, X-1-10-70, XI-7-70 (E. C.
Welling). GUATEMALA. 1 2, El Salto, Escuintla, 1934 (F. A. Bianchi).
COSTA RICA. 1 ¢, La Pacifica, 4 km. NW. of Canas, Guanacaste, XI-17-71
Gera. Opler).

Subgenus Mecas sensu stricto

Mecas LEConreE, 1852, Jour. Acad. Nat. Sci. Philadelphia, (2) vol. 2, p. 155; 1873, Smith-
sonian Misc. Coll., vol. 11, no. 265, p. 347; BratcHLEy, 1910, Coleoptera—in Indiana,
p. 1090; Knutr, 1946, Ohio Biol. Survey, Bull. vol. 39, p. 274.

Form moderate sized, parallel; body usually densely clothed with appressed
pubescence. Pronotum with or without dorsal calluses, sides broadly to narrowly
rounded. Elytra parallel, apices rounded, disk not costate. Legs with tarsal
claws variable, inner tooth long or short.

TypeE species. Phytoecia femoralis Haldemen (monobasic).

The members of this subgenus are easily recognizable by the densely pubes-
cent body, subcylindrical and usually densely pubescent pronotum which fre-
quently has glabrous dorsal calluses.

Thirteen species are presently known.

Mecas obereoides Bates.
(Figure 7.)

Mecas obereoides BATES, 1881, Biologia Centrali-Americana, Coleoptera, vol. 5, p. 204, pl. 15,
fig. 16; BREUNING, 1955, Mem. Soc. Roy. Ent. Belgique, vol. 27, p. 150.

Mecas laminata Bates, 1881, Biologia Centrali-Americana, Coleoptera, vol. 5, p. 204;
BREUNING, 1955, Mem. Soc. Roy. Ent. Belgique, vol. 27, p. 150.

[Proc. 4TH SER.

CALIFORNIA ACADEMY OF SCIENCES

160

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VoL. XXXIX] CHEMSAK AND LINSLEY: MECAS 161

Mecas laminata morpha rufobasalis BREUNING, 1955, Mem. Soc. Roy. Ent. Belgique, vol. 27,

p. 150. New synonymy.

Mecas laminata morpha discopunctata BREUNING, 1955, Mem. Soc. Roy. Ent. Belgique,
vol. 27, p. 150. New synonymy.

Mecas laminata morpha discoimpunctata BREUNING, 1955, Mem. Soc. Roy. Ent. Belgique,
vol. 27, p. 150. New synonymy.

Mate. Form moderate sized, elongate, sides parallel; color black, vertex
of head and pronotum orange, pronotum usually with three black spots, one
median and two lateral. Head with front convex, densely clothed with gray
appressed pubescence which covers basal half of mandibles, erect dark hairs
numerous on front; interantennal area convex; vertex moderately coarsely
punctate; antennae slightly longer than body, densely gray pubescent beneath
with a few long erect hairs present on basal segments, scape slightly shorter
than third segment, fourth equal to third, fifth shorter than first, segments
six to nine subequal, remaining two segments shorter. Pronotum usually a little
broader than long, sides narrowly rounded; disk convex, middle with a usually
black glabrous callus, frequently a vague callus present on each side of middle
anterior to median one, two black glabrous spots present laterally; punctation
irregular, moderately coarse; pubescence sparse, fine with a few long erect hairs
present at sides; prosternum densely clothed with gray recumbent pubescence;
meso- and metasternum densely clothed with gray recumbent pubescence, sides
moderately coarsely punctate. Elytra about three times as long as broad;
each elytron costate down middle, area between costae and suture impressed;
punctures coarse, dense, becoming finer toward apex; pubescence short, dense,
appressed, with longer suberect hairs numerously interspersed; apices rounded.
Legs densely pubescent; tarsal claws with inner tooth slightly smaller than
outer. Abdomen densely clothed with gray appressed pubescence; last sternite
shallowly impressed for most of its length. Length, 10-14 mm.

FEMALE. Form similar, slightly more robust. Antennae about as long as
body. Abdomen with last sternite linearly impressed for its entire length,
shallowly concave at middle at apex. Length, 10-15 mm.

Type LocaLity. Of ‘obereoides, Cuernavaca, Mexico; ‘laminata, not
restricted; ‘rufobasalis, Mexico; ‘discopunctata, Mexico; ‘discoimpunctata,’
Mexico.

RANGE. San Luis Potosi and Sinaloa, Mexico to Guatemala (fig. 7).

FLIGHT PERIOD. June to December.

Host pLants. Unknown.

Remarks. The sparsely pubescent orange pronotum and the dark pronotal
spots will readily separate this species from other known species of Mecas. The
coloration is almost constant in the series at hand. However, specimens from
the eastern parts of Mexico tend to have dark appressed elytral pubescence while
in those from the western portions to Guatemala this tends to be grayish.

MATERIAL EXAMINED. MEXICO. San Luis Potosi: 1 2, 30 mi. N. of

162 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Tamazunchale, X-6-57 (H. A. Scullen). Veracruz: 2 ¢6¢, 3 2%, Veracruz,
VIII-1-6-61 (R. & K. Dreisbach); 1 2, Jalapa, IX-28 to X-3-61 (R. & K.
Dreisbach); 1 ¢, 1 2, Puente Nacional, VIII-3-56 (R. & K. Dreisbach), VII-
23-24-65 (Flint & Ortiz); 2 22, San Andreas Tuxtla, X-25-57 (R. & K.
Dreisbach); 1 ¢, 12, Hueyapan, X-30-57 (R. & K. Dreisbach); 1 4, Cerro
Venado, Los Tuxtlas Range, XII-29-62 (Edmonds, Robinson). Sinaloa: 1 ¢,
Venedillo, VII-31-18; 2 6 3, 20 mi. E. of Villa Union, VIII-19-64 (M. E. Irwin,
E. I. Schlinger). Nayarit: 2 64,1 2,15 mi. N. of Tepic, VII-25-54 (Cazier,
Gertsch, Bradts); 1 ¢, 1 2, 18 mi. N. of Tepic, VIII-16-60 (D. C. Rentz);
3 66, Tepic, [X-21-24-53 (B. Malkin), IX-13-57 (R. & K. Dreisbach); 1 ¢,
14 mi. E. of San Blas, VII-21-54 (Schlinger). Colima: 1 ¢, Colima (Conradt).
Jalisco: 1 2, Puerto Los Mazos, 9 mi. NW. of Autlan, VIII-28-70 (M. & J.
Wasbauer). Guerrero: 2 ¢¢4,1 2, Teloloapan, VIII-15-21-57 (D. Douglas) ;
1 6,1 2,3 mi. S. of Acahuizlotla, XI-17-46 (E. C. Van Dyke); 1 ¢, Taxco,
VITI-16-18-56 (A. E. Lewis); 1 ¢, Highway 95, 23 mi. N. of Acapulco. VII-
30-65 (Cornell U. Mex. field party); 1 ¢, 5 mi. W. of Mex. 92, Cacahuamilpa
Caves, VITI-16-18-56 ( A. E. Lewis). Morelos: 6 6¢,2 22%, Hujintlan, VII-
22-56 (R. & K. Dreisbach); 1 ?, Lake Tequesquitengo, IX-13-57 (H. A.
Scullen); 1 ¢, Tequesquitengo, VII-15-61 (Dreisbach); 1 ¢, Xochicalco,
VII-13-61 (Dreisbach); 5 64, 1 2, Cuernavaca, VII-15-52 (G. M. Boush),
VITI-9-13-38 (L. Lipovsky), VIII-1-6-21 Sept. (Barrett), IX-8-90 (D. De-
long); 1 6,1 2, 22 mi. S. of Cuernavaca, [X-10-57 (Scullen); 1 ¢, 45 mi. S.
of Cuernavaca, IX-12-57 (Scullen). Puebla: 1 ¢, 5 mi. S. of Izucar de Mata-
moros, VIII-1-63 (Parker, Stange). Oaxaca: 1 ¢, 10 mi. NE. of Huajuapan
de Leon, VI-27-65 (Doyen). Chiapas: 1 ?, Santo Domingo, 15 mi. SE. of
Simojovel, VII-8-15-58 (J. A. Chemsak); 6 64,9 2, San Jeronimo, Volcan
Tacana, VIII-10 to X-12-70 (E. C. Welling). GUATEMALA. 1 2, El Salto,
Escuintla, 1934 (F. Bianchi).

Mecas marginella LeConte.

Mecas marginella LECoNTE, 1873, Smithsonian Misc. Coll., vol. 11, no. 264, p. 239; Horn,
1878, Trans. American Ent. Soc., vol. 23, p. 152; Bratcutry, 1910, Coleoptera—in
Indiana, p. 1090; CAsry, 1913, Mem. Coleoptera, vol. 4, p. 361; BreuNntNG, 1955, Mem.
Soc. Roy. Belgique, vol. 27, p. 147.

Mate. Form small, subparallel; color black, pronotum with three longi-
tudinal bands of yellowish to whitish appressed pubescence, elytra with narrow
bands of pale pubescence down suture and lateral margins. Head with front
convex, deeply punctate, densely clothed with appressed pale pubescence, long
erect hairs numerous; interantennal area broadly concave; vertex coarsely,
densely punctate; antennae a little longer than body, very sparsely gray pubes-
cent beneath, long erect hairs numerous on basal segments, third segment longer
than scape, fourth subequal to third, fifth shorter than fourth. Pronotum

Vot. XXXIX] CHEMSAK AND LINSLEY: MECAS 163

broader than long, sides subparallel; punctures moderately coarse, dense, calluses
absent; pubescence dense, appressed, lateral bands broad, yellowish, median
band narrower, usually whitish, remainder of surface finely pubescent, long,
erect hairs numerous; prosternum densely pubescent; meso- and metasternum
densely pubescent, rather coarsely punctate at sides, metepisternum yellow pubes-
cent over posterior half. Elytra over twice as long as broad; punctures coarse,
close, becoming finer toward apex; pubescence between longitudinal bands fine,
appressed, with longer erect hairs numerously interspersed; apices rounded.
Legs finely, densely pubescent; tarsal claws with teeth subequal in length.
Abdomen densely pubescent, narrowly yellow at sides of apical sternites; last
sternite deeply impressed for its entire length. Length, 6.5—-8 mm.

FEMALE. Form and size similar. Antennae about as long as body. Abdomen
with last sternite shallowly impressed near apex. Length, 6.5-8 mm.

TYPE LOCALITY. Western States and Texas.

RANGE. Southeastern United States to New Mexico.

FLIGHT PERIOD. March to July.

Host PLANts. Unknown. One specimen was collected on Colubrina texensis
(Rhamnaceae) in Texas, but it is very unlikely that this shrub is a host.

REMARKS. The absence of pronotal calluses and the distinctive pubescent
bands make this species easily recognizable. Breuning (1955) states that the
head, pronotum, and elytra are densely and very finely punctate but this is
an illusion produced by the pubescence. Actually, they are coarsely punctate,
the elytra less so apically.

MATERIAL EXAMINED. Twenty-one specimens from South Carolina, Alabama,
Texas, and New Mexico.

Mecas confusa Chemsak and Linsley, new species.
(Figure 8.)

Mecas inornata (not Say), BLANCHARD, 1887, Ent. Amer., vol. 3, p. 86; Horn, 1888, Trans.
Amer. Ent. Soc., vol. 15, p. 301; Lenc and HAmirton, 1896, Trans. Amer. Ent. Soc.,
vol. 23, p. 152; BratcHiEy, 1910, Coleoptera—in Indiana, p. 1090.

Mate. Form moderate sized, subparallel; color black, body densely clothed
with thick, grayish, recumbent pubescence which obscures the surface. Head
with front convex, finely densely punctate, darker suberect hairs short, about
half as long as second antennal segment; interantennal area very shallowly con-
cave; vertex sparsely punctate, large punctures well separated; antennae about
as long as body, scape finely gray pubescent, remaining segments to ninth
gray pubescent beneath, third segment longer than scape, fourth shorter than
third, fifth shorter than first, remaining segments gradually decreasing in
length. Pronotum broader than long, sides rounded, base impressed; disk
convex, calluses absent; large deep punctures irregular, well separated, each

[Proc. 4TH SER.

CALIFORNIA ACADEMY OF SCIENCES

164

Se
Sa PS Se

== = SS
a ee a ee

é.

sley,

Mecas confusa Chemsak and Lin

FIGURE 8.

VoL. XXXIX] CHEMSAK AND LINSLEY: MECAS 165

puncture bearing a long erect hair; pro-, meso- and metasterna densely clothed
with recumbent pubescence which obscures the surface. Elytra less than 2%
times as long as broad; punctures coarse, close, linearly arranged, becoming
obsolete at apex; recumbent pubescence completely obscuring surface, base with
numerous rather short suberect hairs, these becoming shorter and recurved
toward apex; apices obliquely subtruncate. Legs very densely pubescent; tarsal
claws with inner tooth almost as long as outer. Abdomen very densely pubes-
cent; last sternite impressed for its entire length. Length, 10-13 mm.

FEMALE. Similar in form and size. Abdomen with last sternite linearly
impressed, apex broadly V-shaped. Length, 10-14 mm.

MATERIAL EXAMINED. Holotype male (California Academy of Sciences)
from Luling, Gonzales Co., Texas, V-3-53 (B. J. Adelson). Allotype from
Gonzales, Gonzales Co., Texas, V-2-53 (M. Wasbauer). Paratypes as follows:
2 64, same data as holotype; 1 2,1 2, same data as allotype; 1 2, Ft. Sam
Houston, Texas (C. Grant); 1 ?, Corpus Christi, Texas, VI-28-42 (E. S. Ross) ;
2 64,1 2, Palmetto State Park, Gonzales Co., Texas, IV-11-53 (M. Wasbauer),
V-10-53 (B. J. Adelson); 1 6, Lee Co., Texas, May (R. Oertel); 1 2, Hidalgo
Co., Texas, VI-22-33 (S. Bromley). Other material, not paratypical includes:
1 2, Texas (F. Blanchard collection); 1 2°, Lee Co., Texas, June; 1 2, 49 mi.
N. of Raymondville, Kenedy Co., Texas, VI-30-61 (R. L. Westcott); 1 4,5 22,
Lake Texoma, 2 mi. E. of Willis, Oklahoma, June, July, 1965 (R. M. Bohart) ;
Que elark Co,, Kansas, June (¥. H. Snow).

This species closely resembles M. cineracea but may be separated by its
larger size, denser overall pubescence, shorter erect hairs on the front of the head,
and by the structure of the tarsal claws. In M. confusa the inner tooth of the
claws is elongate and almost as long as the outer one; in M. cineracea the tooth
is very small. The two species are sympatric, at least in parts of Texas, but it
is not now known whether they infest the same or different host plants.

Mecas femoralis (Haldeman).

Phytoecia femoralis HALDEMAN, 1847, Trans. American Philos. Soc., vol. (2)10, p. 59.

Mecas femoralis, LeContE, 1852, Jour. Acad. Philadelphia vol. (2)2, p. 155; Horn, 1878,
Trans. American Ent. Soc., vol. 7, p. 44; Lenc and Hamirton, 1896, Trans. American
Ent. Soc., vol. 23, pp. 152, 153; Casey, 1913, Memoirs on the Coleoptera, vol. 4, p. 360
(in.) ; BreuNtING, 1955, Mem. Soc. Roy. Ent. Belgique, vol. 27, p. 143.

Mate. Size small, subparallel; color black, femora reddish; pubescence
grayish. Head with front convex, densely punctate, densely clothed with ap-
pressed pubescence and numerous suberect, dark hairs; vertex rather coarsely,
closely punctate, densely pubescent; antennae about as long as body, basal seg-
ments sparsely gray pubescent beneath, long, erect hairs sparse, third segment
longer than first, fourth shorter than third, fifth shorter than fourth. Pronotum
about as long as broad; sides almost subparallel; disk convex, rather coarsely,

166 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

closely punctate; pubescence dense, short, appressed, long, erect hairs numerous;
prosternum densely pubescent; meso- and metasternum densely clothed with re-
cumbent pubescence, sides more coarsely punctate. Elytra about 2’ times as long
as broad; punctures coarse, dense, becoming finer toward apex; pubescence
dense, appressed, with longer erect hairs numerously interspersed; apices rounded.
Legs moderately densely pubescent; tarsal claws with inner tooth very short.
Abdomen densely pubescent; last sternite shallowly impressed over most of
its length. Length, 6-8 mm.

FEMALE. Very similar in size and shape. Abdomen with last sternite im-
pressed over apical one-half. Length, 6-8 mm.

TYPE LOCALITY. Not given.

RANGE. Southeastern United States.

FLIGHT PERIOD. May to July.

Host PLANTS. Unknown.

Remarks. Mecas femoralis can be recognized by its small size, rather
uniform pubescence, lack of pronotal calluses, and reddish femora. This species
appears to be rare in collections. The eleven specimens we have seen vary
very little.

MATERIAL EXAMINED. Eleven individuals from North Carolina to Florida
have been seen.

Mecas cineracea Casey.

Mecas cineracea CAsry, 1913, Memoirs on the Coleoptera, vol. 4, p. 360; Vocr, 1949, Pan-

Pacific Ent., vol. 25, p. 184. (record)

Saperda cineracea, BREUNING, 1955, Mem. Soc. Roy. Ent. Belgique, vol. 27, p. 139.
Saperda bicallosa BREUNING (not Martin), 1955, Mem. Soc. Roy. Ent. Belgique, vol. 27,

p. 140.

Mate. Form rather small, parallel; color black, body densely clothed with
gray recumbent pubescence. Head with front convex, appressed pubescence
obscuring punctures, longer erect hairs numerous; interantennal area almost
plane; vertex moderately coarsely, separately punctate; antennae about as
long as body, basal segments finely gray pubescent beneath, long erect hairs
decreasing in number toward apex, third segment longer than first, fourth shorter
than third, fifth a little shorter than first, remaining segments gradually de-
creasing in length. Pronotum broader than long, sides almost parallel, shal-
lowly impressed at base; disk convex, without calluses, punctures coarse,
separated; appressed pubescence obscuring surface, long erect hairs numerous;
prosternum densely pubescent; meso- and metasternum densely pubescent,
coarsely punctured at sides. Elytra about 2% times as long as broad; punctures
coarse, contiguous, becoming finer toward apex; pubescence obscuring surface,
long suberect hairs numerous near base; apices obliquely truncate. Legs very
densely pubescent; tarsal claws with inner tooth small. Abdomen densely

VoL. XXXIX] CHEMSAK AND LINSLEY: MECAS 167

pubescent; last sternite shallowly impressed for its entire length. Length,
6-10 mm.

FEMALE. Similar in form and size. Antennae shorter than body. Abdomen
with last sternite linearly impressed. Length, 7-11 mm.

TYPE LOCALITY. Harris Co., Texas.

RANGE. Southeastern United States to Arizona and Colorado and _ north-
eastern Mexico.

FLIGHT PERIOD. April to August.

Host pLants. Helenium microcephalum, Baileya multiradiata.

Remarks. This species may be recognized by its small size, lack of pronotal
calluses, uniform coloration and pubescence, and by the small inner tooth of
the tarsal claws. It was incorrectly transferred to Saperda by Breuning (1955).

A series of specimens from western New Mexico and Arizona have thicker
pubescence than the Texas examples. However, there appears to be a gradient
in this character from east to west as is apparent in the material at hand.

MATERIAL EXAMINED. More than 200 specimens from South Carolina
to Florida, to Arizona and Colorado.

Mecas cinerea (Newman).
(Figure 9.)

Saperda cinerea NEWMAN, 1840, Entomologist vol. 1, p. 13.

Mecas cinerea, GAHAN, 1888, Trans. American Ent. Soc., vol. 15, p. 300.

Mecas senescens BATES, 1881, Biologia Centrali-Americana, Coleoptera, vol. 5, p. 203; CAsEy,
1913, Memoirs on the Coleoptera, vol. 4, p. 360.

Mecas rubripes BATES, 1881, Biologia Centrali-Americana, Coleoptera, vol. 5, p. 203;
BREUNING, 1955, Mem. Soc. Roy. Ent. Belgique, vol. 27, pp. 140, 143. New synonymy.

Mecas rubripes morpha callosoreducta BREUNING, 1955, Mem. Soc. Roy. Ent. Belgique,
vol. 27, pp. 140, 143. New synonymy.

Mecas inornata GAHAN, 1888 (not Say), Trans. Amer. Ent. Soc. vol. 15, p. 300.

Mate. Form moderate sized, elongate, slender, parallel; color black, femora
and/or elytra often reddish, pronotum occasionally with a median longitudinal
reddish band; pubescence gray to yellowish, dense, appressed, long erect hairs
fairly numerous. Head with front convex, finely separately punctate; inter-
antennal area slightly concave; vertex deeply, separately punctate; pubes-
cence dense, obscuring surface, long erect hairs moderately numerous; an-
tennae slightly longer than body, segments gray pubescent beneath, basal
segments with a few long erect hairs beneath, third segment longer than first,
fourth subequal to third, fifth equal to first, remaining segments gradually
decreasing in length, eleventh subacute at apex. Pronotum slightly broader
than long, cylindrical; disk with two small glabrous calluses on each side of
basal median elongate callus; punctures moderately coarse, separated at
middle and subconfluent at sides; appressed pubescence obscuring surface,

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CALIFORNIA ACADEMY OF SCIENCES

168

VoL. XXXIX] CHEMSAK AND LINSLEY: MECAS 169

denser at middle and forming a vague longitudinal band, long erect hairs fairly
sparse, shorter erect hairs more numerous; prosternum densely pubescent;
meso- and metasternum densely pubescent, rather densely punctate at sides.
Elytra usually about 3 times as long as broad; punctures coarse and contiguous
basally, becoming finer and sparser toward apex; pubescence short, dense,
appressed, long erect hairs numerous basally, becoming shorter and suberect
toward apex; apices rounded. Legs moderately densely pubescent; tarsal claws
with inner tooth slightly shorter than outer. Abdomen densely pubescent; last
sternite rather shallowly impressed for its entire length. Length, 8-11 mm.

FEMALE. Form similar. Antennae about as long as elytra. Pronotum more
transverse, distinctly broader than long. Abdomen with last sternite linearly
impressed. Length, 8-13 mm.

TYPE LOCALITY. Of ‘cinerea,’ Mexico; ‘senescens,’ Puebla; ‘rubripes,’ Mex-
ico; ‘callosoreducta,’ Mexico.

RANGE. San Luis Potosi to Morelos and Nayarit (fig. 9).

FLIGHT PERIOD. June to September.

REMARKS. The number of calluses on the pronotum, elongate body form,
and the length of the antennae distinguish this species from the other Mexican
species of Mecas. The body and antennal length, and different tarsal claws
separate it from M. pergrata. It is difficult to detect a definite geographical
variational trend from the material available for study. The more northern
individuals are all black while most of the southern specimens possess reddish
femora and often, reddish elytra. Two individuals from Nayarit have a reddish
longitudinal band on the pronotum and yellowish epipleurae.

MATERIAL EXAMINED. MEXICO. San Luis Potosi: 2 646, 17 miles W. of
Xilitla, 4700 ft., VII-22-54 (Univ. Kansas Mex. Exped.); Hidalgo: 1 4,2 92,
7 miles NE. of Zimapan, VIII-15-58 (H. F. Howden); Distrito Federal: 5 64,
7 22, Temascaltepec, 1931 (G. B. Hinton); 1 ¢, 2 22, Real de Arriba,
Temascaltepec, VII-32 (H. E. Hinton), VII-33 (Hinton and Usinger); 2 4,
5 22, Tejupilco, Temascaltepec, VI-VII-33 (Hinton and Usinger); Puebla:
1 2,15 miles S. of Puebla, 6200 ft., IX-6-57 (H. A. Scullen); 1 4, Atlixco,
VII-23-56 (R. & K. Dreisbach); Mexico: 14,12, 3 miles N. of Valle de Bravo,
VI-28-29-65 (G. H. Nelson); Morelos: 7 é 6, Cuernavaca VII-6-38, VII-29-61,
7000 ft. (R. & K. Dreisbach); 1 ¢, 12, Cuernavaca-Acapulco Road, VIII-22-36
(Ball & Stone); 1 ¢, 4 miles E. of Cuernavaca, 6000 ft., VI-25-59 (H. E.
Evans); 1 6, 7 km. E. of Cuernavaca, 5700 ft., VIII-11-62 (Evans); Nayarit:
1 6,1 2, La Mesa de Nayarit, VII-21-55 (B. Malkin).

Mecas pergrata (Say).

Saperda pergrata SAy, 1824, Jour. Acad. Philadelphia, vol. 3, p. 407; HaLpEMAN, 1847,
Trans. American Philos. Soc., vol. (2)10, p. 55; LEConTE, 1859, Complete Writings of
Thomas Say, vol. 2, p. 190.

170 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Stenostola pergrata, HALDEMAN, 1847, Proc. American Philos. Soc., vol. 4, p. 373; LEConreE,
1852, Jour. Acad. Nat. Sci. Philadelphia, vol. (2)2, p. 154.

Mecas pergrata, Horn, 1878, Trans. American Ent. Soc., vol. 7, p. 44; Lenc and Hamiton,
1896, Trans. American Ent. Soc., vol. 23, pp. 152, 153; BLarcHLry, 1910, Coleoptera—in
Indiana, pp. 1090, 1091; CAsry, 1913, Memoirs on the Coleoptera, vol. 4, p. 361;
CRAIGHEAD, 1923, Dom. Canada Agr. Bull., vol. 27, p. 138; Knut, 1946, Ohio Bio.
Survey, Bull. vol. 39, pp. 274, 275, pl. 22, fig. 86; BrEUNING, 1955, Mem. Soc. Roy.
Ent. Belgique, vol. 27, pp. 140, 144, fig. 1.

Mecas pergrata morpha semiruficollis BREUNING, 1955, Mem. Soc. Roy. Ent. Belgique, vol. 27,
pp. 140, 145. New synonymy.

Stenostola gentilis LECONTE, 1852, Jour. Acad. Nat. Sci. Philadelphia, vol. (2)2, p. 154.

Mecas discovittata BREUNING, 1955, Mem. Soc. Roy. Ent. Belgique, vol. 27, pp. 140, 143.
New synonymy.

MALE. Form moderate sized, parallel; color black, femora pale reddish,
elytra occasionally partly reddish; pubescence dense, short, recumbent, grayish.
Head with front convex, punctures rather fine, well separated; pubescence dense,
appressed, long dark erect hairs numerously interspersed; appressed pubescence
thicker around eyes; vertex rather densely punctate; antennae shorter than body,
segments gray pubescent beneath, outer segments annulate, third segment longer
than first, fourth subequal to first, remaining segments gradually decreasing
in length. Pronotum broader than long, sides slightly rounded; disk convex,
four glabrous calluses present in addition to median callus; punctures rather
sparse, scattered; apex and base usually with a narrow band of dense yellowish
pubescence, remaining surface partially obscured, long, erect hairs numerously
interspersed; prosternum densely pubescent; meso- and metasternum densely
pubescent, densely punctate at sides. Scutellum densely clothed with yellowish
recumbent pubescence. Elytra about 2’ times as long as broad; punctures rather
coarse, contiguous at base, becoming finer toward apex; pubescence short, re-
cumbent, partially obscuring surface, longer suberect hairs numerous, suture and
lateral margins narrowly clothed with dense, yellowish, appressed pubescence;
apices rounded. Legs finely, densely pubescent; femora reddish; tarsal claws
with inner tooth small. Abdomen densely pubescent; last sternite shallowly im-
pressed over most of its length. Length, 6-11 mm.

FEMALE. Similar in form and size. Antennae extending to about second
abdominal segment. Abdomen with last sternite linearly impressed. Length,
6-12 mm.

TypeE LocaLity. Of ‘pergrata,’ Platte River, Nebraska; ‘gentilis,’ Missouri
Territory; ‘semiruficollis, Texas; ‘discovittata,’ Colorado.

RANGE. Great Plains to southeastern United States, New Mexico, and north-
eastern Mexico.

FLiGHT PERIOD. April to July.

Host pLAnts. Aster (roots), Helianthus.

Remarks. The five glabrous spots of the pronotum, reddish femora, and the

Vor. XXXIX] CHEMSAK AND LINSLEY: MECAS 171

densely pubescent lines on the suture and lateral margins of the elytra will
readily distinguish this species. In certain parts of the range, the elytra tend
to be reddish down the disk and frequently the pronotum is also partially
reddish.

Hasits. According to Craighead (1923) larvae feed in the stems of Aster
and down into the roots, completely hollowing the latter. Subsequently that
portion of the stem of the plant breaks off at the surface of the ground. Small
heaps of frass are exuded about the base of the plant. Only one larva is found
in each stem.

Mecas cirrosa Chemsak and Linsley, new species.

Mate. Form moderate sized, robust; color black; pubescence dense, gray,
recumbent. Head with front convex, rather finely, irregularly punctate; inter-
antennal area rather deeply impressed; vertex coarsely, separately punctate;
pubescence dense, appressed, long erect hairs numerous, pale and dark colored;
antennae slightly shorter than body, segments robust, somewhat flattened, all
segments gray pubescent beneath, dark brown above, segments from second
densely fringed with long curved hairs beneath, third segment slightly longer
than first, fourth a little shorter than third, fifth shorter than first, remaining
segments gradually decreasing in length. Pronotum broader than long, sides
broadly rounded; disk convex, each side with a small glabrous, premedian callus,
narrow median callus extending almost length of disk; punctures moderately
coarse, irregular, subconfluent; pubescence fine, appressed, long, pale, erect hairs
numerous; prosternum densely pubescent; meso- and metasternum densely
pubescent, minutely punctate. Elytra more than 2% times as long as broad;
basal punctures moderately coarse, well separated, punctures becoming fine
and quite sparse toward apex; pubescence fine, dense, appressed, basal margin
with a few long, erect, pale hairs, remainder of surface with very short, suberect,
dark hairs; apices rounded. Legs densely pubescent; tarsal claws with inner
tooth short, obtuse, lobe-like. Abdomen densely pubescent; last sternite deeply
impressed. Length, 13 mm.

FEMALE. Form similar. Antennae shorter than body, curved hairs less
numerous beneath. Abdomen with last sternite linearly impressed. Length,
10.5 mm. .

TYPE MATERIAL. Holotype male (California Academy of Sciences) from
5 miles N. of Guanajuato, Guanajuato, Mexico, VII-25-54 (E. I. Schlinger). The
specimen that we consider as the female of this species differs in several respects.
The pubescence appears thicker and the antennae have a much sparser fringe.
This individual is from Km. 320, near Hacienda Balvanera, Queretero, Mexico,
VIT-13-55.

Remarks. Although the tarsal claws are uniquely different from other

172 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

species of Mecas, we consider them to be bifid and the other morphological char-
acteristics are similar enough to place ‘cirrosa’ in the genus.

Mecas humeralis Chemsak and Linsley, new species.

Mate. Form moderate sized, parallel; color black; pubescence dense,
grayish and brownish. Head with front convex, rather finely, separately
punctate; interantennal area impressed; vertex rather finely, separately punc-
tate; pubescence dense, appressed, long, dark, erect hairs numerous; antennae
about as long as body, outer segments slightly flattened, all segments gray
pubescent beneath, dark brown above, long erect hairs sparse on basal segments,
third segment much longer than first, fourth a little shorter than third, fifth
equal to first, remaining segments gradually decreasing in length. Pronotum
slightly broader than long, sides broadly rounded; disk convex, each side with
a large glabrous callus before middle, median callus long, rather broad; punctures
moderately coarse, subconfluent; pubescence dense, appressed, long erect hairs
numerous; prosternum densely pubescent; meso- and metasternum densely
pubescent, finely densely punctate at sides. Elytra about 3 times as long as
broad; basal punctures coarse, contiguous, becoming finer and sparser toward
apex; pubescence dense, recumbent, gray on disk, humeri glabrous, epipleurae
and sides with a broad band of brown pubescence extending from under humeri
almost to apex, long erect hairs abundant at base, becoming shorter and recurved
toward apex; apices rounded. Legs densely pubescent; tarsal claws with inner
tooth short, blunt, lobe-like. Abdomen densely pubescent, first three sternites
with a narrow glabrous band at base; last sternite deeply impressed. Length,
13 mm.

FEMALE. Form similar. Antennae about as long as body. Legs with tarsal
claws having the short inner tooth slightly acute. Abdomen lacking glabrous
lines on sternites, last sternite linearly impressed. Length, 10.5 mm.

TYPE MATERIAL. Holotype male (California Academy of Sciences) from
El Molino, Jalisco, Mexico, VII-10-56 (R. & K. Dreisbach); allotype from
Guadalajara, Jalisco, VII-24-51 (P. D. Hurd).

REMARKS. This species has tarsal claws similar to those of M. cirrosa. The
different antennae and antennal pubescence will readily separate the two species.
The glabrous humeri also make M. humeralis distinctive.

Mecas bicallosa Martin.

Mecas bicallosa Martin, 1924, Ent. News, vol. 35, p. 244; Hatcu, 1971, Univ. Washington
Publs. Biol., vol. 16, p. 155.
Saperda bicallosa, BREUNING, 1955, Mem. Soc. Roy. Ent. Belgique, vol. 27, pp. 139, 140.

Mate. Form moderate sized, parallel, rather robust; color black, body
densely clothed with short, appressed, grayish pubescence. Head with front

VoL. XXXIX] CHEMSAK AND LINSLEY: MECAS 173

convex, pubescence obscuring surface, long erect hairs very numerous; inter-
antennal area plane, vertex deeply punctate; antennae extending to about third
abdominal segment, segments through fourth gray pubescent, dark at apices,
basal segments with numerous long, suberect hairs, segments from third with
long hairs beneath, these decreasing in number toward apex, third segment
longer than first, fourth subequal to first, remaining segments short, subequal
in length. Pronotum broader than long, sides rounded; disk convex, with two
glabrous calluses at middle and a smaller median one behind middle; punctures
rather fine, deep, separated; pubescence obscuring surface, very long erect hairs
numerous; prosternum densely pubescent, front coxal cavities narrowly open
behind; meso- and metasternum densely clothed with recumbent and_sub-
depressed pubescence. Elytra more than twice as long as broad; punctures at
base coarse, dense, becoming finer toward apex; pubescence obscuring surface,
long, suberect hairs abundant over basal half; apices rounded, often vaguely,
obtusely toothed. Legs very densely pubescent; tarsal claws with inner tooth
very small, short. Abdomen densely pubescent; last sternite shallowly, rather
broadly impressed. Length, 10-13 mm.

Femate. Form similar. Antennae slightly shorter. Abdomen with last
sternite narrowly linearly impressed, apex shallowly concave. Length, 10-
>) ma.

Type LocaLity. Martins Springs, Lassen Co., California.

RANGE. Washington to northern Baja California, to Colorado.

FLIGHT PERIOD. April to August.

Host PLaAnts. Artemisia tridentata.

Remarks. The bicallused pronotum and abbreviated distal antennal seg-
ments characterize this species. Breuning (1955) incorrectly synonymized
M. bicallosa with M. cineracea Casey and transferred both to Saperda.

MATERIAL EXAMINED. A total of 127 specimens were examined from
Washington, Oregon, California, Nevada, Idaho, Utah, Colorado and Baja
California.

Mecas ambigena Bates.

Mecas ambigenus BATES, 1881-1885, Biologia Centrali-Americana, Coleoptera, vol. 5, pp. 203,

426; BREUNING, 1955, Mem. Soc. Roy. Ent. Belgique, vol. 27, pp. 142, 147.

Mecas pseudambigena BREUNING, 1955, Mem. Soc. Roy. Ent. Belgique, vol. 27, pp. 142,

147. New synonymy.

Mate. Form rather small, short, parallel; color black; pubescence dense,
rather coarse, brownish, appressed, long dark erect hairs numerous. Head with
front convex, finely densely punctate; interantennal area concave; vertex
distinctly, separately punctate; pubescence short, appressed, front with numerous
very long erect black hairs; antennae extending to about apical of elytra,

174 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

segments to tenth gray pubescent beneath, segments from fourth narrowly
annulate at base, basal segments with a moderate number of suberect hairs
beneath, scape robust, a little shorter than third segment, fourth subequal to
first, remaining segments decreasing in length. Pronotum broader than long,
sides broadly rounded; disk convex with a glabrous callus on each side before
middle, elongate median basal callus often vague or absent; punctures coarse,
confluent; pubescence short, appressed with numerous long erect hairs inter-
spersed; prosternum densely pubescent; meso- and metasternum densely pubes-
cent, finely densely punctate. Elytra less than 2% times as long as broad;
punctures coarse, contiguous, becoming finer toward apex; pubescence mod-
erately densely appressed, suture narrowly lined with dense yellow-brown
pubescence, long erect hairs numerous over basal ’2, shorter and suberect toward
apex; apices rounded. Legs densely pubescent; tarsal claws with inner tooth
smaller than outer. Abdomen densely pubescent, finely punctate; last sternite
deeply impressed for about ™ its length. Length, 9 mm.

FEMALE. Similar in form and size. Antennae extending to about apical %
of elytra. Abdomen with last sternite lightly linearly impressed. Length, 10 mm.

Type LocaLity. Of ‘ambigena,’ Mexico; ‘pseudambigena,’ Mexico.

RANGE. Durango to Mexico.

FLIGHT PERIOD. June and July.

Remarks. The shorter form and short antennae will distinguish M. ambigena
from other species of Mecas.

MATERIAL EXAMINED. MEXICO. Durango: 1 ¢, 25 miles W. of Durango,
7,500 ft., VI-23-64 (J. E. Martin); Zacatecas: 2 64, 4 miles W. of Monte
Escobido, 7,800 ft., VII-19-20-54 (R. H. Brewer); Mexico: 1 ¢&, Toluca
(Wickham).

Mecas menthae Chemsak and Linsley, new species.

(Figures 10, 11.)

Mecas marginella, LINSLEY, KNULL, and STATHAM (not LeConte), 1961, Amer. Mus. Nov.,
nNOwZ20505 pas2-

Mate. Form moderate sized, subparallel; color black; pubescence dense,
short, appressed, grayish to gray-brown, long erect dark hairs numerous. Head
with front convex, rather finely, separately punctate; vertex deeply, separately
punctate; pubescence dense, short, appressed, antennal tubercles dark pubescent
above, long erect hairs numerous on front and vertex; antennae slightly longer
than elytra, segments to tenth gray pubescent beneath, segments from fifth
narrowly pale annulate at base, long erect hairs fairly numerous beneath on
basal segments, third segment longer than first, fourth shorter than third but
longer than first, fifth equal to first, remaining segments gradually decreasing
in length. Pronotum broader than long, sides broadly rounded; disk convex,
each side of middle with a rather large, glabrous callus, middle with an elongate

VoL. XXXIX] CHEMSAK AND LINSLEY: MECAS 175

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CALIFORNIA ACADEMY OF SCIENCES

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VoL. XXXIX] CHEMSAK AND LINSLEY: MECAS 177

callus toward base; punctures rather fine, separated; pubescence short, appressed,
obscuring surface, long erect hairs numerous; prosternum densely pubescent;
meso- and metasternum densely pubescent, finely densely punctate at sides.
Scutellum densely clothed with appressed pubescence. Elytra more than
2% times as long as broad; punctures coarse, contiguous to about apical %,
very fine at apex; pubescence dense, short appressed, lateral margins and
suture with a narrow band of appressed pubescence, long erect hairs numerous
basally, shorter toward apex; apices rounded. Legs densely pubescent; tarsal
claws with inner tooth slightly shorter than outer. Abdomen densely pubescent;
last sternite deeply impressed for about * of its length. Length, 8-12 mm.

FEMALE. Similar in form, slightly more robust. Antennae about as long as
elytra. Abdomen with last sternite linearly impressed. Length, 9-13 mm.

MATERIAL EXAMINED. Holotype male, allotype (California Academy of
Sciences) and 99 paratypes (60 males, 39 females) from 8 miles W. of El
Palmito, Sinaloa, Mexico, VII-19, 24, 29-64, VIII-5-64 on Agastache (J. A.
Chemsak, J. A. Powell, H. F. Howden). Additional material not paratypical
assignable to this species includes: 9 6 6,5 2 2, 6.5 miles E. of Potrerillos, Hwy.
30, Sinaloa, VIII-20-21-64 (E. 1. Schlinger, P. Rauch); 2 ¢ 4, 20 miles E. of Villa
Union, Sinaloa, VIII-20-64 (P. Rauch); 1 2, 63 miles E. of Jct. Hwy. 15 &
40 on Hwy. 40, Mexico, VIII-28-64 (D. C. & K. A. Rentz, J. A. Grant); 1 ¢,
El Pichon, Nayarit, Mexico, VI-25-63 (J. Doyen); 1 4, Southwestern Research
Station, Chiricahua Mts., Arizona, VIII-24-58 (P. D. Hurd); 1 6,1 2°, Yank’s
Spring, Sycamore Canyon, Tumacacori Mts., Santa Cruz Co., Arizona, VII-28-
65 (H. B. Leech); 1 ¢, Oak Creek Canyon, 12 miles S. of Sedona, Arizona,
VII-18-57 (C. W. O’Brien); 1 2, Swift Trail, between Ladybug Saddle and
Shannon Park, Pinaleno Mountains, Graham County, Arizona, VI-27-58 (J. M.
& S. N. Burns). Other specimens from Mexico tentatively assigned to M.
menthae; 1 2, 2 miles S. of Tlaquepaque, Jalisco, VII-11-53 (C. & P. Vaurie) ;
1 2,5 km. W. of Zacapu, Michoacan, VII-13-51 (H. E. Evans); 1 2, Real de
Arriba, Distrito Federal, VII-32 (H. E. Hinton); 1 6, 2 2%, Temescaltepec,
Distrito Federal, 1931 (G. B. Hinton); 1 2, Tejupilco, Temescaltepec, VI-33
(H. E. Hinton, R. L. Usinger).

This species differs from M. ambigena by the much longer antennae, finer
pubescence, and much less coarsely punctate pronotum.

The type series was collected during the day from the apical leaves of the
mint Agastache. The adults were found resting in the curve of the smaller top
leaves of the plant. Numerous individuals were mating and this plant is prob-
ably the larval host.

Mecas cana (Newman).

Saperda cana NewMan, 1840, Entomologist, vol. 1, p. 12; LeConrer, 1852, Jour. Acad.
Nat. Sci. Philadelphia, vol. (2)2, p. 164.

178 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Mecas cana, GAHAN, 1888, Trans. American Ent. Soc., vol. 15, p. 300; Lenc and Hamirton,
1896, Trans. American Ent. Soc., vol. 23, p. 152; Casey, 1913, Memoirs on the Coleoptera,
vol. 4, p. 360; BREUNING, 1955, Mem. Soc. Roy. Ent. Belgique, vol. 27, p. 148.

MALE. Form moderate sized, subparallel; color black, body densely clothed
with gray recumbent pubescence. Head with front convex, pubescence obscuring
punctures, longer, dark, suberect hairs numerous; interantennal area almost
plane; vertex finely, separately punctate; antennae slightly shorter than body,
segments to sixth rather sparsely grey pubescent beneath, long erect hairs de-
creasing in number toward apex; third segment longer than scape, fourth
shorter than third but longer than first, remaining segments gradually decreasing
in length. Pronotum broader than long, sides rounded to subparallel; basal
and apical margins narrowly margined; disk convex, each side with a flat
glabrous callus before middle, middle usually with a vague linear callus near
base; punctures moderately coarse, rather sparse, partially obscured by pubes-
cence; longer erect hairs pale with dark setae interspersed mostly at sides;
prosternum densely pubescent; meso- and metasternum densely pubescent;
metasternum deeply rather densely punctate at sides. Elytra over 2% times
longer than broad; punctures moderately coarse, well separated, becoming
finer and sparser toward apex; pubescence obscuring surface, longer suberect
hairs numerous; apices obliquely subtruncate. Legs densely pubescent; tarsal
claws with inner tooth small. Abdomen occasionally reddish, densely pubescent;
last sternite moderately impressed for its entire length. Length, 9-15 mm.

FEMALE. Form similar. Antennae shorter than body. Abdomen with last
sternite linearly impressed. Length, 10-16 mm.

TYPE Locatity. St. John’s Bluff, Florida.

RANGE. Southeastern United States to Texas, northeastern Mexico, and
South Dakota.

REMARKS. This species may be separated from M. confusa by the short
inner tooth of the tarsal claws. The pronotal calluses readily distinguish it from
M. cineracea and the elongate antennal segments from M. bicallosa.

Two subspecies can be recognized.

Mecas cana cana (Newman).

Saperda cana NEWMAN, 1840, Entomologist, vol 1, p. 12; LeConre, 1852, Jour. Acad. Nat.
Sci. Philadelphia, vol. (2)2, p. 164.

Mecas cana, GAHAN, 1888, Trans. American Ent. Soc., vol. 15, p. 300; Casry, 1913, Memoirs
on the Coleoptera, vol. 4, p. 360; BREUNING, 1955, Mem. Soc. Roy. Nat. Belgique, vol. 27,
p. 148.

Body densely grayish pubescent, elytra narrowly clothed at suture and
lateral margins with bands of appressed pubescence. Length, 10—-12.5 mm.

TypE LocALity. St. John’s Bluff, Florida.

RANGE. Florida.

VoL. XXXIX] CHEMSAK AND LINSLEY: MECAS 179

FLIGHT PERIOD. April to October.

Host pLrants. Ambrosia, Flaveria linearis.

REMARKS. This subspecies appears to be restricted to Florida, primarily
the southern portion. Although Breuning (1955) states that the type of M. c.
cana appears to be lost, it is in the collection of the British Museum (Natural
History).

Mecas cana saturnina (LeConte).

Stenostola saturnina LECONTE, 1859, Smithsonian Contr. Knowledge, vol. 11, p. 21.

Mecas saturnina, GAHAN, 1888, Trans. American Ent. Soc. vol. 15, p. 300; Horn, 1888,
Trans. American Ent. Soc., vol. 15, p. 301; Brreuntnc, 1955, Mem. Soc. Roy. Ent.
Belgique, vol. 27, p. 146; Wirson, 1960, Comm. Inst. Biol. Control Tech. Comm. vol. 1,
p. 62; StrmpE and WARWICK, 1962, Anim. Behaviour, vol. 10, p. 112 (habits).

Mecas inornata (not Say), Horn, 1878, Trans. American Ent. Soc., vol. 7, p. 44; KNULL,
1946, Ohio Biol. Surv. Bull. vol. 39, p. 274; Dmton and Ditton, 1961, Man. Common
Beetles of Eastern North America, p. 652, pl. 65, no. 17.

Mecas brevicollis CAsEY, 1913, Memoirs on the Coleoptera, vol. 4, p. 362.

Similar in form and size to typical subspecies. Pubescence uniformly gray
or yellowish, elytra without pubescent bands on margins and suture. Length,
9-16 mm.

TYPE LocaALity. Of ‘saturnina, Kansas; ‘brevicollis, WKansas.

RANGE. Alabama to northeastern Mexico to South Dakota.

FLIGHT PERIOD. May to August.

Host pLants. Ambrosia, Xanthium, Helianthus, Gaillarda. Adults have
also been taken on Prosopis and Salvia in Texas.

INCERTAE SEDIS

Mecas albovitticollis Breuning.

Mecas albovitticollis BrEuNtNG, 1955, Mem. Soc. Roy. Ent. Belgique, vol. 27, p. 146.
Mecas (?)———————————(?) Bates, 1885, Biologia Centrali-Americana, Coleoptera,
vol. 5, p. 427.

“Antennae a little longer than the body. Lower lobes of the eyes obviously longer
than the cheeks. Head and pronotum densely and very finely punctate. Pronotum
transverse. Elytra slightly truncate at the apices, densely and finely punctate.

“Black covered with light gray pubescence. Pronotum ornated with three straight,
longitudinal, discal, whitish bands. Scutellum having whitish pubescence. Antennae
faced with deep brown pubescence.

“Length: 10 mm; width: 2% mm.

“Type: a male from Mexico in the British Museum.”

We have been unable to place this species on the basis of the above original
description and were unable to locate the type at the British Museum. However,
we believe that this species was based on the single specimen cited by Bates in
the Biologia Centrali-Americana. Since this reference stated: “One example

180 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

of a distinct species, in bad condition, doubtfully belonging to this genus.’’, we
will follow Bates and exclude it from the genus Mecas.

Mecas marmorata Gahan.

Mecas marmorata GAHAN, 1892, Trans. Ent. Soc. London. 1892, p. 268, pl. 12, fig. 7;
BREUNING, 1955, Mem. Soc. Roy. Belgique, vol. 27, pp. 140, 152.

Mate. Form elongate, slender; color black, pronotum and elytra suffused
with pale reddish brown, antennal segments 2 to 10 reddish; pubescence dense,
yellowish, appressed, arranged in patches on elytra, forming a longitudinal
band down middle of pronotum. Pronotum cylindrical, as long as broad;
mesosternal process slightly broadened. Elytra more than 3 times longer than
broad; punctures coarse, irregular; apices produced, dentate. Legs short;
mesotibiae with sinus; tarsal claws with inner tooth smaller than outer. Abdomen
with last sternite deeply impressed.

TYPE LOCALITY. Guerrero, Mexico (not restricted further).

The elongate body, apically produced elytra, and cylindrical pronotum
appear sufficient to exclude this species from Mecas. Since we have examined
only the type in the British Museum (Natural History) and one other male
from 17 miles south of Puebla, Puebla, IX-6-57 (H. A. Scullen) we have not
attempted to clarify the generic status of ‘marmorata’ at this time.

ACKNOWLEDGMENTS

This study was made as part of a National Science Foundation sponsored
monograph of North American Cerambycidae (Grant GB-31120-X). Although
a large portion of the material we examined came from the California Academy
of Sciences and the California Insect Survey, Berkeley, we wish to gratefully
acknowledge the following institutions for the loan of specimens: American
Museum of Natural History; Canadian National Collection, Ottawa; Cornell
University; Field Museum of Natural History, Chicago; Los Angeles County
Museum; Museum of Comparative Zoology, Harvard University; Instituto
Tecnologico y de Estudios Superiores de Monterrey, Monterrey, Mexico;
Ohio State University; Oregon State University; United States National
Museum; University of California, Davis and Riverside, University of Kansas;
collections of G. H. Nelson and R. L. Westcott.

We are deeply appreciative to Celeste Green for preparing the illustrations
and to the authorities of the British Museum (Natural History) for permitting us
to examine type specimens in their collection.

LITERATURE CITED

BaArerc, W. J.
1921. A girdler on artichokes and other little-known insect pests. Journal of Economic
Entomology, vol. 14, pp. 99-100.

VoL. XX XIX] CHEMSAK AND LINSLEY: MECAS 181

Bates, H. W.
1881-1885. Biologia Centrali-Americana, Insecta, Coleoptera, Lamiidae, vol. 5, pp. 153-
224, pls. xii-xv, pp. 334-436, pls. xxii-xxv. [Mecas, pp. 203-205.]
BEUTENMULLER, W.
1896. Food-habits of North American Cerambycidae. Journal of the New York Ento-
mological Society, vol. 4, pp. 73-81.
Brake, S. F.
1951. Compositae. Jn: Kearney, T. H. and R. H. Peebles, Arizona Flora, pp. 829-971.
University of California Press.
BLANCHARD, F.
1887. Notes on Coleoptera. Entomologia Americana, vol. 3, pp. 85-88.
BLATCHLEY, W. W.
1910. On the Coleoptera known to occur in Indiana. Indiana Department of Geology
and Natural Resources, Bulletin 1, 1386 pp.
BREUNING, S.
1952. Revision einiger Gattungen aus der Gruppe der Saperdini Muls. Entomologische
Arbeiten aus der Museum G. Frey, vol. 3, pp. 107-213.
1955. Revision du genre Mecas LeConte. Memoires de la Société Royale d’Entomologie
de Belgique, vol. 27, pp. 138-152, 1 fig.
1967. Mecas. Catalogue des Lamiaires du Monde, vol. 10, pp. 830-832.
CasEy, T. L.
1913. Mecas LeC. Memoirs on the Coleoptera, vol. 4, pp. 360-363.
CuHEmMsAK, J. A., AND E. G. LINSLEY
1965. A revised key to the species of Elytroleptus with notes on variation and distri-
bution. The Pan-Pacific Entomologist, vol. 41, pp. 193-199.
CRAIGHEAD, F. C.
1923. North American cerambycid larvae. Dominion of Canada, Department of
Agriculture Bulletin No. 27, new series, 238 pp., 44 pls.
Diton, E. S. anp L. S. Ditton
1961. A manual of the common beetles of Eastern North America. Row, Peterson
and Company. 884 pp.
GaHaNn, C. J.
1888. Notes on some types of North American Cerambycidae in the British Museum.
Transactions of the American Entomological Society, vol. 15, pp. 299-300.
1892. Additions to the Longicornia of Mexico and Central America, with notes on
some previously-recorded species. Transactions of the Entomological Society
of London. 1892, pp. 255-274.
Gimour, E. F.
1962. Synopsis of the tribe Aerenicini. Revista de Biologia Tropical, vol. 10, pp. 123-147.
HALDEMAN, S. S.
1847. Material toward a history of the Coleoptera Longicornia of the United States.
Transactions of the American Philosophical Society, vol. (2)10, pp. 27-66.
1847. Corrections and additions to his paper on the Longicornia of the United States.
Proceedings of the American Philosophical Society, vol. 4, pp. 371-376.
Hatcu, M. H.
1971. The beetles of the Pacific Northwest. Part V. University of Washington Publi-
cations in Biology, vol. 16, 662 pp.

182 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Horn, G. H.
1878. Notes on some genera of Cerambycidae of the United States. Transactions of
the American Entomological Society, vol. 7, pp. 41—50.
1888. Additional notes. Transactions of the American Entomological Society, vol. 14,
pp. 300-301.
Kincssury, J. M.
1967. Poisonous plants of the United States and Canada. Prentice-Hall, New Jersey.
xiii + 626 pp., 130 figs.

Knut, J. N.
1946. The long-horned beetles of Ohio. Ohio Biological Survey Bulletin 39, pp. 133-
354, 29 pls.
LAcorDAIRrE, J. T.
1872. Genera des coléoptéres. . ., vol. 9, no. 2, pp. 411-930.
LEConrmE, J. L.

1852. An attempt to classify the longicorn Coleoptera of the part of America north
of Mexico. Journal of the Academy of Natural Sciences of Philadelphia, vol.
(2)2, pp. 139-178.
1859a. The Coleoptera of Kansas and Eastern New Mexico. Smithsonian Contributions
to Knowledge, no. 11, pp. 1-58.
1859b. The complete writings of Thomas Say on the entomology of North America. 2
vols. New York.
1873a. New species of North American Coleoptera. Part II. Smithsonian Miscel-
laneous Collections, No. 264, pp. 169-240.
1873b. Classification of the Coleoptera of North America. Part II. Smithsonian
Miscellaneous Collections, No. 265, pp. 279-348.
LEConTE, J. L., anp G. H. Horn
1883. Classification of the Coleoptera of North America. Smithsonian Miscellaneous
Collections, no. 507, xxxviii +- 567 pp.
Lenc, C. W. ano J. HAMILTON
1896. Synopsis of the Cerambycidae of North America. Part III, the Lamiinae. Trans-
actions of the American Entomological Society, vol. 23, pp. 101-178.
Linsey, E. G.
1962. Synopsis of the genus Elytroleptus Dugés. Folia Entomologica Mexicana, no. 3,
pp. 1-13.
Linstey, E. G., J. N. Knut, Ano M. STATHAM
1961. A list of Cerambycidae from the Chiricahua Mountain area, Cochise County,
Arizona. American Museum Novitates, no. 2050, pp. 1-34, 24 figs.
Martin, J. O.
1924. Studies in the genus Mecas. Entomological News, vol. 35, pp. 244-245.
NEwmMan, E.
1840. Entomological notes. Entomologist, vol. 1, pp. 1-16.
Norp, J. C., AND F. B. KnicHT
1970. Saperda inornata Say, 1824 (Insecta, Coleoptera): Proposed use of the plenary
powers to designate a neotype to stabilize the nomenclature. Bulletin of Zoo-
logical Nomenclature, vol. 27, pp. 123-128.
Rey, C. V.
1880. Food habits of the longicorn beetles or wood borers. American Entomologist,
vol. 3, pp. 237-239, 270-271.

VoL. XXXIX] CHEMSAK AND LINSLEY: MECAS 183

Save) Tr.

1824. Descriptions of coleopterous insects collected in the late expedition to the Rocky
Mountains, performed by order of Mr. Calhoun, Secretary of War, under the
command of Major Long. Journal of the Academy of Natural Sciences of
Philadelphia, vol. 3, pp. 139-216.

STRIDE, G. O., AND E. P. WArRwIckK

1962. Ovipositional girdling in a North American cerambycid beetle, Mecas saturnina.

Animal Behaviour, vol. 10, pp. 112-117.
THomson, J.

1864. Systema cerambycidarum on expose de tous les genres compris dans la famille
des Cerambycides et familles limitrophes. Memoires des Société Royale des
Sciences de Liege, vol. 19, pp. 1-540.

1868. Physis recueil d’Histoire Naturelle, vol. 2, pp. 6-208.

TOWNSEND, C. H. T.
1884. Notes on some Coleoptera taken in South Louisiana. Psyche, vol. 4, pp. 219-222.
Voct, G. B.

1949. Notes on Cerambycidae from the lower Rio Grande Valley, Texas. The Pan-

Pacific Entomologist, vol. 25, pp. 137-144, 175-184.
Witson, F.

1960. A review of the biological control of insects and weeds in Australia and Australian
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munication no. 1, pp. 1-102.

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PROCEEDINGS

OF THE Weoatls Hale, wage,
CALIFORNIA ACADEMY OF §S

FOURTH SERIES

Vol. XXXIX, No. 13, pp. 185-256; 26 figs.; 7 tables. July 9, 1973

COMPARATIVE REVISION OF SCOMBEROIDES,
OLIGOPLITES, PARONA, AND HYPACANTHAUS
WITH COMMENTS ON THE PHYLOGENETIC

POSITION OF CAMPOGRAMMA
(PISCES: CARANGIDAE)'

=
William F. Smith-Vaniz? and Jon C. Staiger

Rosenstiel School of Marine and Atmospheric Science, University of Miami
10 Rickenbacker Causeway, Miami, Florida 33149

TABLE OF CONTENTS

LVDS CYST ce SS ee RE ner eee WO OE 186
ee een aE DEY Hoe rants ek ei Oe oe Te Tae ee 186
Mrrseercrseanur MeCnOUS . te a ee 187
Semmsmscumucrores Lacepede = 2 toe oe ee eee 190
Mevarorspecies\GL scomberoides — 193
Neomberoides commersonianus Lacépede ——-__-___ 194
ReaMICROIGeS COld ((CUVIEL)" 2225. ee 199
RUCHOIOeS VSN |(CHOrSKal) Se a ee 205
tie ROTC SHLOUM\ OC UNIGE)) oe A. 2 ee a ae eee eee 209
DamuUerOres. SPINOSUS (SMITROV )) 22 ee ee ee eee Dil
ene AOU pTILes Gili o= Se Sa 2 2 AUS Vie a ee 213
evarospecies. 0: Oligoplitess =! ee 217
Oicoplites réjuigens Gilbert: and Starks — 27
Mapes SGiICNS (EOC) ey F2RN0 i) Fes 3 ee ee 218

1 Contribution number 1624 from the Rosenstiel School of Marine and Atmospheric Science, University
of Miami.

2Present address: The Philadelphia Academy of Natural Sciences, Nineteenth and the Parkway,
Philadelphia, Pa. 19103.

[185]

186 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Olicoplites-saurus (Block and Schneider). Eee 221
Oligopliesraitus,( Gunther), ..2 4 32% 2". 5 J ee eee 222
Oligoplites pelometa (Cuvier) .....<-. eee 225
Genus Parone Bete. 2 tS ee 225
Porona signata (Jenyns) ...- Eee 227
Genus Aypacanthus Ratinesque\._..__.__ Eee 228
Hy pacanthus ana (Linnaeus) Eee 230
Relationships and Zoogeography .....___.._.___.._ ____.__ _ = aaa Zou
Phylogenetic position of Campogramma Regan __.---._--------_--------------- 244
SUMMMATY = 32s ee ee Se i A Oe ee ee 247
Nominal species of Scomberoides and Oligoplites ____----_________ 248
Literature Cited. 2-2 ee eee 250
INTRODUCTION

This study evolved as a result of our efforts to unscramble the confused
synonymies of Indo-Pacific carangids of the genus Scomberoides. The initial
effort led us to review the species of Oligoplites, a New World genus closely
related to Scomberoides, last treated by Schultz (1945). It soon became obvious
that our paper would not be complete unless we included coverage of several
monotypic Atlantic genera that are superficially similar to the ‘leatherjackets’
of American waters. Comparison of these three carangid genera is especially
desirable because they, together with Oligoplites, are the only ones, of the 18
Atlantic genera that we currently recognize, which do not occur on both sides of
the Atlantic. The combinations of characters exhibited by these genera have
impressed us with the artificiality of the present subfamilial partitioning of the
Carangidae. In discussing relationships of these genera it is necessary to em-
phasize some of the inadequacies of the present subfamilial classification of
the Carangidae.

ACKNOWLEDGMENTS

We wish to thank the following individuals who assisted us in obtaining data
for our study, either through hospitality at their institutions or by providing
data on, or loans of, specimens (abbreviations refer to institutions and are ex-
plained in the methods section): Donn E. Rosen, AMNH; John R. Paxton,
AMS; N. B. Marshall and Alwyne C. Wheeler, BMNH; William N. Eschmeyer
and staff, CAS; M. L. Bauchot, MNHN; Carl L. Hubbs and Richard H. Rosen-
blatt, SIO; Warren C. Freihofer, SU; Frederick H. Berry, TABL; Boyd W.
Walker, UCLA; C. Richard Robins, UMML; Victor G. Springer, Martin F.
Gomon and Edgar E. Gramblin, USNM; Jérgen G. Nielsen, ZMC; R. J. McKay,
WAM.

Vor. XXXIX] SMITH-VANIZ & STAIGER: CARANGIDAE 187

Special thanks are due J. R. Paxton and R. J. McKay for providing identi-
fications and locality data for collections of Scomberoides housed in their re-
spective institutions, thus confirming our tentative zoogeographical conclusions.
Tomio Iwamoto examined MNHWN types of Scomberoides (subsequently re-
examined by us) that provided much of the impetus for initiation of the present
paper.

Additional help came from various sources. James E. Bohlke and William
N. Eschmeyer facilitated our study by making literature available. Phillip C.
Heemstra called to our attention several pertinent references. William H. Hulet
and Francisco J. Palacio assisted in translating Italian and Portuguese literature.
Grady W. Reinert illustrated the specimens shown in figure 1. The manuscript
was typed by Mrs. Kathie Jeffries.

Work space at the Southeast Fisheries Center Miami Laboratory, National
Marine Fishery Service, where much of the material cited in this paper is
housed, was generously provided by the Director, Mr. Harvey R. Bullis.

Frederick H. Berry and C. Richard Robins reviewed the manuscript and
offered suggestions for improvement.

Major collections of Scomberoides critical to our study were obtained in
Ceylon under Smithsonian Foreign Currency Grant 3818, C. Richard Robins
and Frederick H. Berry, principal investigators. Field work in Ceylon was
greatly facilitated by the cooperation of Mr. A. S. Mendis, Deputy Director
(Research), Department of Fisheries, Ceylon. The second author was able to
pursue this study through support of the National Geographic Society-Univer-
sity of Miami Deep-Sea Biology Program (University of Miami research account
8852R), Gilbert L. Voss, Principal Investigator. Partial support was also pro-
vided by a grant from the Tai Ping Foundation to Smith-Vaniz for study of
Indian Ocean Carangidae.

CHARACTERS AND METHODS

Osteological terminology generally follows that of Suzuki (1962); a specific
exception is the “postmaxillary process” which is here more appropriately
termed a premaxillary spur since it is a posterior projection on the premaxilla
that braces the maxilla. A basibranchial is defined (Nelson, 1969, p. 480) as
the median part of the gill arch endoskeleton that lies between successively
paired arch-elements. The number of the basibranchial refers to the paired
arch-element behind it.

Rayless predorsal interneurals are termed predorsal bones (Smith and
Bailey, 1961). All carangids have at least 3 predorsal bones; counting poste-
tiorly, the fourth element, whether supporting a dorsal-fin spine or not, has a well
developed spinelike projection on its anterodorsal margin. This structure,
commonly termed a procumbent spine, may occasionally protrude through the

188 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

skin. In the genera treated in this study, all anterior dorsal pterygiophores
support a single fin spine. Thus, a shortening of the dorsal-fin base through
loss of anterior spines results in a concomitant increase in number of predorsal
bones. The first fin-ray (spine) is often greatly reduced, depressed posteriorly,
and can easily be overlooked. In carangids there is a tendency for the number
of external spines to decrease in large specimens due to absorption, as evidenced
from examination of radiographs. In this study large specimens were not x-
rayed or dissected; thus the frequency of specimens with the least number of
dorsal spines may be slightly lower than indicated in tables 2 and 4.

The last dorsal- and anal-fin rays are split to the base, have a single pteryg-
iophore, and were counted as one ray.

The first caudal vertebra has a well developed haemal spine which artic-
ulates with the first anal pterygiophore; the urostyle is regarded as the terminal
caudal vertebra.

Counts of outer row dentary teeth in Scomberoides and Oligoplites are lateral
counts (one side only). Counts were made on the dentary that appeared to
have the most teeth. Broken teeth were counted but empty tooth sockets and
teeth that had not attained a functional position in the outer row were excluded.
Symphyseal dentary canines were also excluded in the case of Scomberoides.

Pectoral-fin ray counts are lateral counts (one side only). The dorsalmost
fin-ray element is spinelike and was counted separately.

Gillraker counts are lateral counts and were usually made on the right side
of each fish. Only gillrakers on the lateral side of the first arch were counted.
The gillraker at the angle of the upper and lower limb has the major portion
of its base on the ceratobranchial bone and was included in the count of the
lower limb. Rudimentary gillrakers (with the width of raker greater than its
length) were not included in the counts.

All measurements are straight line (point to point) measurements. Mea-
surements smaller than 120 mm. were made with dial calipers. Measurements
given in the material examined sections are fork length (FL), unless otherwise
stated, and have been rounded off to the nearest millimeter. In large fishes
with relatively rigid caudal fins, such as carangids, the end of the hypural plate
is often difficult to determine precisely. In addition to being easier to determine
than standard length, fork length is more familiar to fishery biologists, who
should have the greatest need to identify carangids. The following measurements
were used in this study:

Fork length (FL). Snout tip to tip of shortest median caudal-fin ray.

Standard length (SL). Snout tip to posterior end of hypural bones (caudal
base).

Head length (HL). Snout tip to posterior margin of fleshy opercular flap.

Snout-postorbit length. Snout tip to anterior margin of posterior adipose eyelid.

VoL. XXXIX] SMITH-VANIZ & STAIGER: CARANGIDAE 189

Snout length. Snout tip to posterior margin of anterior adipose eyelid.

Upper jaw length. Snout tip to posterior margin of maxilla.

Body depth Dz to Ag. Origin of terminal dorsal spine to origin of terminal anal
spine.

Height of fin lobes. Origin of fin to tip of longest segmented ray.

Pectoral-fin length. Origin of spine to tip of longest ray.

Abbreviations of institutions cited are as follows:

AMNH American Museum of Natural History, New York City

AMS Australian Museum, Sydney

BMNH British Museum (Natural History), London

CAS California Academy of Sciences, San Francisco

FMNH Field Museum of Natural History, Chicago

MNHN Muséum National d’Histoire Naturelle, Paris

NFIS Natur-Museum und Forschungs-Institut Senckenberg, Frankfurt

SIO Scripps Institution of Oceanography, University of California, La Jolla

SU Stanford University, Division of Systematic Biology (collection trans-
ferred to CAS)

TABL _ Southeast Fisheries Research Center, National Marine Fishery Service,
Miami (formerly Tropical Atlantic Biological Laboratory)

UCLA University of California at Los Angeles

UMML University of Miami, Rosenstiel School of Marine and Atmospheric
Science, Miami

USNM National Museum of Natural History, Division of Fishes, Washington,
DPC.

ZMC Zoological Museum, Copenhagen

WAM Western Australian Museum, Perth.

Uncataloged California Academy of Sciences specimens collected under the
auspices of the George Vanderbilt Foundation (GVF) are reported under their
register or station numbers. Most collections from Ceylon are uncataloged and
listed by station numbers. Abbreviations refer to the following collectors listed
in chronological order of their visits to Ceylon: F. J. Schwartz; W. F. Smith-
Vaniz; P. C. Heemstra; C. C. Koenig.

Localities in material examined sections are abbreviated. Summary state-
ments of material examined include only specimens examined by the authors.

190 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Scomberoides Lacépéde, 1802

Scomberoides LACEPEDE, 1802, p. 50 (type-species: Scomberoides commersonianus LACEPEDE,
1802, by subsequent designation of JORDAN, 1917, p. 60).

Orcynus RAFINESQUE, 1815, p. 84 (nomen nudum; substitute name for Scomberoides LACEPEDE,
1802).

Chorinemus Cuvier in Cuvier and Valenciennes, 1831, p. 367 (type-species: Scomberoides
commersonianus LACEPEDE, 1802, by subsequent designation of JORDAN, 1917, p. 137).

Rhaphiolepis FOWLER, 1905, p. 59 (as a subgenus of Scomberoides; type-species: Chorinemus
tol Cuvier in Cuvier and Valenciennes, 1831, by original designation).

Eleria JORDAN and SEALE, 1905, p. 774 (type-species: Eleria philippina JoRDAN and SEALE,
1905 = Chorinemus tala Cuvier in Cuvier and Valenciennes, 1831, by monotypy).

Palaeoscomber SmirRNov, 1936, p. 49-59 (type-species: Palaeoscomber spinosus SMIRNOV,
1936, by original designation).

The carangid genus Scomberoides comprises four recent and one fossil species
of tropical and subtropical marine fishes restricted to the Indian and western
and central Pacific oceans. All recent species attain at least 45 cm. fork length.
Scomberoides commersonianus, the largest species, occasionally exceeds 100 cm.
and is a valued game fish. In some areas S. commersonianus supports a seasonal
fishery of considerable importance. Major catches are obtained from drift nets
set at depths of 15 to 18 m. in waters surrounding offshore islands (James,
1967). Depending on the region, scomberoides are considered excellent food
fishes (Smith, 1949, p. 224) or they are mostly salted and not generally esteemed
(Chan, 1968, p. 82).

Halstead et al. (1972) give a detailed description of the venom apparatus
of Scomberoides lysan. Venom glands are associated with the first seven dorsal-
fin spines and the first two anal spines. The anal spines, rather than the dorsal
spines, are the most venomous. The anal spines are equipped with a frictional
locking device which resembles that found in the fin spines of catfishes and
balistid triggerfishes. Scomberoides lysan is capable of inflicting painful stings.
The nearly identical morphology of these spines in the other species of Scom-
beroides and Oligoplites (see comments under O. saurus) suggest that all species
of both genera are venomous.

The osteology of Scomberoides has received inadequate attention. The only
species studied in detail (under the name Chorinemus orientalis) is Scomberoides
lysan (Suzuki, 1962). The only known fossil species of Scomberoides was orig-
inally described as belonging to a new genus, Palaeoscomber, of Scombridae
(Smirnov, 1936) and was subsequently redescribed as a species of Oligoplites
(Danil’chenko, 1960).

The nomenclature of Scomberoides species has been greatly confused by
a plethora of nominal species inadequately described by early workers. Differ-
ences ascribed to nominal species have frequently been based on unrecognized
ontogenetic changes. Changes in dentition associated with growth led Jordan

VoL. XXXIX] SMITH-VANIZ & STAIGER: CARANGIDAE 191

TasleE 1. Nominal species of Scomberoides recognized in selected publications with our
present allocation.

Authors Species
Present Study commersonianus tala lysan tol
Smith (1970) lysan — tolooparah tala
Williams (1958) lysan — sanctipetri —
Roxas and Agco (1941) lysan tala tolooparah tol
Weber and lysan tala tolooparah tol
De Beaufort (1931) sanctipetri
Wakiya (1924) lysan _ sanctipetri formosanus
orientalis
moadetta
Day (1878) lysan tala sanctipetri moadetta
toloo

and Seale (1905, p. 774) to describe a new genus and species based on juveniles
of S. tala. The failure of recent workers to critically read original descriptions
and examine types adequately has resulted in the consistent misapplication of
the name Scomberoides lysan (Forskal). Because of the confused nomenclature
of Scomberoides species, we present in table 1 a list of the species recognized in
several major papers with our present allocation of them. An annotated list of
the nominal species of Scomberoides and Oligoplites is given in the appendix.

NOMENCLATURE. Considerable confusion exists as to whether Scomberoides
Lacépéde, 1802 was validly described and thereby has priority over Chorinemus
Cuvier, 1831. Williams (1958) briefly discussed the problem and rejected
Scomberoides, as did Weber and De Beaufort (1931) and Smith (1970). Re-
jection of Scomberoides as a valid generic name by several recent workers is
probably attributable to the comments of Barnard (1927, p. 562) who credits
the rejected name to Lacépéde (1800), in which only a figure is given with the
caption ‘“Scomberoide commersonnien.” Jordan (1917, p. 60) rightly credits
Scomberoides to Lacépéde (1802), in which ‘‘Les Scomberoides” was latinized
to accommodate the description of three new species. Caranx and Trachinotus,
widely accepted carangid generic names, originate from the same work and were
described in the same manner as Scomberoides.

Williams (1958, p. 416) states that “the use of Chorinemus has been common
and widespread since 1833 whilst Scomberoides has been restricted (except
Wakiya, 1924) to American authors of the last 75 years.”’ He further states
“should Scomberoides be proved to have priority over Chorinemus C.V..... :
then a case exists for submission to the International Commission of Zoological
Nomenclature for a suspension of the Rules, in that strict application of the
Rules will clearly result in greater confusion than uniformity.” In their review

192 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

of the Philippine Carangidae Roxas and Agco (1941) used Scomberoides as did
Oshima (1925) and McCulloch (1929). Except for Rofen (1963), American
authors of the last 75 years, the most recent of which are Gosline and Brock
(1965), have consistently used Scomberoides instead of Chorinemus. Although
Williams apparently attaches little importance to this, it should be noted that
American authors are responsible for a large portion of the literature treating
these fishes.

Confusion alse exists as to what constitutes the type-species of the genus.
Several workers have recognized Scomberoides noelii as the type-species.
Lacépede (1802) described three species in his new genus, listing them on page
50 with brief diagnoses (descriptions expanded on following pages) in the fol-
lowing order: (1) Scomberoides noelii; (2) S. commersonianus; (3) S. saltator.
He did not, however, specifically designate a type-species. The first author to
subsequently designate a type-species for Scomberoides in accordance with
Article 69a of the International Code of Zoological Nomenclature was Jordan
(1917, p. 60), who selected S. commersonianus. The published illustration of
S. commersonianus (as “Scomberoide commersonnien”) (Lacépéde, 1800 pl.
20, Fig. 3) is sufficiently accurate for positive identification.

The possession of needlelike scales does distinguish the type-species of Rhaph-
iolepis from its congeners. However, the transition from oval-shaped to needle-
like scales is obvious in Scomberoides (see figure 3), and we do not believe that
recognition of subgenera in such a small group of closely related species is
justified. The sole character mentioned in the original description to distinguish
Eleria from Scomberoides was based on unrecognized ontogenetic changes in
dentition (see discussion under S. tala). Reasons for referral of Palaeoscomber
to the synonymy of Scomberoides are discussed under S. spinosus.

DeEscrIPTION. Body moderately to strongly compressed. Adipose eyelids mod-
erately to poorly developed. Body, except head, covered with embedded scales,
broadly lanceolate to needlelike. Lateral-line scales not developed into scutes.
Lateral line without side branches, straight except for slight curve over pectoral
fin. Anterior rim of shoulder girdle smooth, without fleshy papillae or deep
groove near isthmus. Premaxillae not protractile, connected anteriorly to snout
at midline by a wide fleshy bridge. Swimbladder strongly bifid posteriorly,
length of bifurcated portion of swimbladder equal to or greater than undivided
part. Posterior dorsal- and anal-fin rays consisting of semidetached finlets,
distal fourth to half of ray not connected by interradial membrane (unattached
portion of rays increasing with growth); ultimate and penultimate rays more
closely spaced than adjoining rays, fully connected by interradial membrane;
none of dorsal- or anal-fin rays produced into long filaments. Pelvic fins rel-
atively short, depressible into shallow abdominal grooves. Pectoral fins short,
not falcate, 53 to 71 percent head length in adults. Maxilla extends posteriorly

Vout. XXXIX] SMITH-VANIZ & STAIGER: CARANGIDAE 193

from middle of eye to well behind orbital rim, 43 to 65 percent head length,
in adults.

Predorsal bones 3, rarely 4 (4 in one of 60 specimens); dorsal-fin rays VI
or VII + I, 19-21; total dorsal-fin rays 27-29. Anal-fin rays II + I, 16-20.
Pectoral-fin rays I, 16-18. Pelvic-fin rays I, 5. Vertebrae 10 + 16 = 26;
epipleural ribs 9 or 10. Branchiostegal rays 3 + 5 (epihyal + ceratohyal).
Upper gillrakers 0-8; lower gillrakers 7-20; total gillrakers 8-27.

Fronto-supraoccipital crest low; temporal crest extending forward to edge
of cranium slightly in front of middle of orbit; preorbital region short. Epiotics
broadly united along midline of cranium internally. Zygapophyses of exoccipital
joined beneath foramen magnum. Parasphenoid not expanded into a broad,
flattened plate posteriorly. Myodome with broad, posterior opening. Basi-
occipital without a pair of lateral processes on ventral surface. Ascending process
of premaxilla short and triangular; maxilla long and slender, rather closely
attached to premaxilla. Premaxillary spur absent. Small supramaxilla present.
Interosseous space between dorsal arm of dentary and upper margin of articular
greatly reduced. Well developed teeth on dentary, premaxilla, palatine, and
vomer; two distinct rows of dentary teeth (see discussion under Zoogeography
and Relationships Section); symphyseal premaxillary and dentary teeth of
juveniles (except S. /ysan) robust and caniniform. Mesopterygoids covered with
firmly ankylosed, minute, granular teeth. Pharyngeals not noticeably enlarged,
covered with sharp, pointed teeth.

Well developed suborbital shelf present; lower and posterior suborbitals
not greatly expanded posteriorly. Basibranchials 1-2 with a pair of tooth
plates, third basibranchial with two or more irregular tooth plates. Lower limb
of posttemporal not noticeably short or thickened. Postcoracoid process absent.
Interpelvic keel well developed; post pelvic process of each side coalesced entire
length, not forming an apical fork. Inferior vertebral foramina absent. First
anal pterygiophore and haemal spine of first caudal vertebra firmly attached
to form a strong strut. Anal pterygiophore expanded anterolaterally to form
roof over first two anal spines; first two anal spines articulated proximally
with each other; second anal spine asymmetrical, with deep groove on one side
of anterior face. Caudal skeleton with 2 epurals and 2 pairs of uroneurals.

Key To Recent Spectres oF Scomberoides

Color pattern will readily separate most specimens of the four species. In large individuals
and especially in small specimens that are not freshly caught the spots may fade or completely
disappear; this is particularly true of fish market specimens.

aeeGalirakerses) tom is dorsal-tine lobe) unttonmly plemented = 2
1b. Gillrakers 21 to 27; distal half of dorsal-fin lobe abruptly and heavily pigmented __ 3
2a. Large oval blotches above or touching lateral line; upper jaw extends well beyond

posterior margin of eye, especially in adults (fig. 2a); snout 21 to 26 (mean 23.4)

194 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

percent HL; dentary teeth of inner and outer rows subequal in adults; specimen fig-
UL ell yp oS cat ven ae ee 2 Et ie ee ee S. commersonianus
2b. Vertically elongate blotches intersecting lateral line; upper jaw extends slightly beyond
posterior margin of eye (fig. 2b); snout 26 to 30 (mean 28.2) percent HL; inner row
dentary teeth distinctly larger than outer row teeth in adults; specimen figure 1b —__ S. tala
3a. Double series of 6 to 8 dusky, roundish blotches above and below lateral line, occasionally
connected by narrow isthmus; scales lanceolate (fig. 3b) ; specimen figure 1c ___ S. lysan

¢

3b. Oval or vertically oblong blotches, the first 4 or 5 intersecting lateral line; scales slender,
needlelike -(fig) Sd); specimen figure Id) 22 == Eee Ss Hal!

Scomberoides commersonianus Lacépéde.
(Figures la, 2a, 3a, 4a, 5—7, 14c, 16b, 19, 23b, 25b; tables 2-3.)
Scomberoides commersonianus LACEPEDE, 1802, p. 50 (type locality Madagascar).

Scomber madagascariensis SHAW, 1803, p. 590, pl. 85 (type locality Madagascar).
Chorinemus exoletus EHRENBERG in Cuvier and Valenciennes, 1831, p. 379 (278) (based
solely on an unpublished drawing by Ehrenberg of a fish from Lohaia, Red Sea).
Chorinemus delicatulus RICHARDSON, 1846, p. 269 (based solely on a drawing in the Reeves

Collection of Chinese fish drawings).
Chorinemus leucopthalmus RicHARDSON, 1846, p. 269 (based solely on a drawing in the
Reeves Collection of Chinese fish drawings).

NOMENCLATURE. The figure of S. commersonianus (as “Scombéroide com-
mersonnien”) and original description are sufficiently accurate for positive
identification, and the name has consistently been applied to the correct species,
although usually in synonymy (see nomenclature section under S. lysan). Al-
though in the original description S. commersonianus was spelled with a double
“n,”’ we prefer the emended spelling with a single “‘n,’’ which is correct.

The description of Chorinemus exoletus refers to a deep-bodied species with
a long upper jaw. This together with the stated type locality is sufficient to
confidently identify it as conspecific with S. commersonianus.

Whitehead (1969) has published the original drawings upon which the de-
scriptions of Chorinemus delicatulus and C. leucopthalmus are based. These
drawings and that of Scomber madagascariensis are reasonably diagnostic and
are undoubtedly based on S. commersonianus. Although the large oval blotches
characteristic of S. commersonianus are not shown on the sides of Chorinemus
delicatulus and C. leucopthalmus, they rapidly fade and disappear in dead
specimens.

DescripTION. Characters given in the generic description are not repeated
here. Frequency distributions of selected meristic characters are given in table
2. Dorsal-fin rays VI-VII + I, 19-21; anal-fin rays II + I, 16-19; pectoral-
fin rays, I, 17-19. Gillrakers 0-3 + 7-12 = 8-15. Upper jaw extending well
beyond posterior margin of eye, especially in adults (fig. 2a). Scales broadly
lanceolate (fig. 3a). Inner and outer row of dentary teeth subequal in adults
(fig. 4a); dentary typically with one or two pair of symphyseal canines, at least

VoL. XXXIX] SMITH-VANIZ & STAIGER: CARANGIDAE 195

Ficure 1. Scomberoides species: a, S. commersonianus, TABL 107366, 468 mm. FL;
b, S. tala, TABL 107365, 464 mm. FL; c, S. lysan, TABL 107364, 523 mm. IGS Ch, ual
TABL 107363, 450 mm. FL.

[Proc. 4TH SER.

CALIFORNIA ACADEMY OF SCIENCES

196

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VoL. XXXIX] SMITH-VANIZ & STAIGER: CARANGIDAE 197

Cc d

Ficure 2. Upper jaw development of adult Scomberoides species: a, S. commersonianus,
TABL 107192, 355 mm. FL; b, S. tala, TABL 107712, 351 mm. FL; c, S. lysan, TABL 107364,
412 mm. FL; d, S. tol, TABL 107714, 410 mm. FL. Maxilla shown in black.

in young; young with dentary teeth of outer row numerous and considerably
more closely spaced than inner row teeth.

Proportional measurements apply only to specimens longer than 150 mm.
FL, and are expressed as percent FL unless otherwise stated. Depth (origin
Dz to Ag) 25.7-36.2; height dorsal-fin lobe 14.4-19.8; height anal-fin lobe 13.5-
18.5; upper jaw 55.9-64.6 percent head length; snout 20.6—26.0 percent head
length.

PIGMENTATION. Sides with 5 to 8 large oval blotches above or touching
lateral line, first two may intersect lateral line. Dorsal and anal fins uniformly
pigmented. Pectoral fin of adults frequently with a dusky blotch ventrally.
In life, body gray to silvery white ventrally, dusky green or bluish dorsally;

198 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

TABLE 3. Comparison of several proportional measurements in Scomberoides species
expressed as hundredths of head length.

Upper Jaw Length Snout Length
<20 cm. = =

Species Range FL N Range x Range %
commersonianus 82-198 19 59-64 60.1 21-26 23.6
tala 71-199 23 53-61 58.2 26-30 28.3
lysan 45-189 20 51-56 53.9 28-31 29.0
tol 44-181 17 47-51 48.6 30-33 Sue3

Upper Jaw Length Snout Length

re pper J g g

Species Range FL N Range X Range x
commersonianus 224-943 64 56-65 60.6 21-26 23.4
tala 201-571 58 54-61 57.8 26-30 28.2
lysan 201-585 65 50-54 51.6 28-32 30.0
tol 209-468 70 43-49 46.3 29-33 Silks

lateral blotches plumbeous gray. Large individuals often golden, especially
ventrally. Annunziata and Cedrone (1972, p. 61) present a color photograph
of a large specimen from Madagascar.

DISTRIBUTION. Widely distributed throughout the Indian Ocean and Indo-
Australian Archipelago. The species has also been reliably reported from the
Red Sea and Persian Gulf. Scomberoides commersonianus appears to be re-
stricted to neritic waters and does not occur east of the New Guinea-Solomon
Islands region. Records of the species (as S. /ysan) from Tahiti and other
central Pacific islands are based on misidentifications.

MATERIAL (96 specimens, 82—943 mm. FL, from 53 collections). KENYA:
Mombasa Fish Market, TABL uncataloged Anton Bruun cruise 9 station, FT-2
(1: 398). WEST PAKISTAN: off Karachi, TABL uncataloged (3: 236-295).
INDIA: Gulf of Kutch, TABL uncataloged (1: 203); Bombay, TABL 107190
(1: 384), 107350 (1: 158); Palk Strait, TABL 107292 (4: 127-288), TABL
uncataloged FHB 66-36 (1: 766); Porto Novo, TABL 107303 (1: 198), TABL
uncataloged (1: 190). CEYLON: TABL 107192 FJS 69-41 (9: 327-422),
FJS 69-45 (6: 195-244), FJS 69-64 (1: 474), FJS 69-50 (1: 400), TABL
107366 S-V 69-77 (3: 303-476), S-V 69-86 (2: 224-254), S-V 69-126 (1: 335),
S-V 69-151 (2: 754-762), PCH 69-183 (1: 617), PCH 69-185 (7: 123-158),
TABL 107715 PCH 69-201 (3: 398-443), PCH 69-218 (10: 130-168), PCH
69-267 (1: 503), PCH 69-305 (1: 913), CCK 69-1 (1: 843), CCK 69-44
(1: 797), CCK 69-53 (1: 475), CCK 69-60 (7: 234-349), CCK 69-97 (2: 160-
218). GULF OF THAILAND: GVF station 57-45 (2: 355-391), GVF station
57-92 (1: 520), GVF station 57-83 (2: 559-579), GVF station 60-113 (2:

VoL. XXXIX] SMITH-VANIZ & STAIGER: CARANGIDAE 199

849-943), GVF station 60-146 (2: 727-729), GVF-1548 (1: 133), GVF-1557
(2: 186-197), GVF-2195 (1: 93, cleared and stained), GVF-2513 (2: 727-739),
GVF-2546 (1: 291), GVF-2547 (6: 243-612). HONG KONG: GVF station
HiK-111 (1: 829). PHILIPPINES: Luzon, USNM 72190 (1: 172), 83447
(1: 241), 168322 (1: 309), SU 20341 (2: 82-116); Samar, USNM 149810
(1: 129); Iloilo, USNM 72189 (1: 129), 149812 (1: 152); Palawan, USNM
149813 (1: 128), 149815 (1: 151). NORTH BORNEO: Sandakan, USNM
149814 (1: 172), 168323 (1: 234). JAVA: Djakarta, TABL 107300 (1: 266).
AUSTRALIA: Western Australia, Exmouth Gulf, Onslow, WAM P.2772 (1:
302), P.15487 (1: 175); Port Hedland, WAM P.180 (1: 329); Broome, WAM
P.555 (1: 95); Northern Territory, Charles Point, AMS IA.7781 (1: 164);
Port Darwin, AMS 1.9778 (1: 300), IA.4375 (1: 280); Darwin, AMS 1B.3168
(1: 420), IB.3171 (1: 85); Bedwell Point, AMS IA.7678 (1: 356); Gulf of
Carpentaria, Groote Eylandt, USNM 173954 (4 of 7: 240-345); 17°22’S.,
149°45’E., AMS T.15557-118 (5: 87-172) ; Queensland, Burdekin River, USNM
47835 (1: 275), AMS A.18299-300 (2: 285-615); Lindeman Island, AMS
IA.6279 (1: 297); Hayman Island, AMS IA.6104 (1: 460); New South Wales,
Port Jackson, USNM 47910 (1: 357). NEW GUINEA: AMS A.12694 (1:
300); Port Moresby, AMS 1.13357 (1: 565).

Scomberoides tala (Cuvier).

(Figures 1b, 2b, 3c, 4b, 5-7, 15b, 17, 19, 24b; tables 2-3.)

Chorinemus tala CUVIER in Cuvier and Valenciennes, 1831, p. 377 (type locality Malabar).

?Chorinemus toloo Cuvier in Cuvier and Valenciennes, 1831, p. 377 (description based on
“toloo parah” RUSSELL, 1803, p. 29, pl. 137, Vizagapatam).

Eleria philippina JorpAN and SEALE, 1905, p. 744, fig. 1 (type locality Negros Island,
Philippines).

Chorinemus hainanensis Cuu and CHENG, 1958, p. 317, fig. 2 (type localities Sanya, Kanchium,
Kwonghoi, and Chinglan, China).

NOMENCLATURE. In the Muséum National d’Histoire Naturelle there are
two collections (A. 6587 and A. 6588) from Malabar, collected by Belanger,
which are labeled syntypes of Chorinemus tala Cuvier. A. 6588, here designated
lectotype of C. tala, is a single specimen 298 mm. SL. A. 6587 consists of two
specimens of S. tol Cuvier. In his treatment of the western Indian Ocean species
of Scomberoides, Smith (1970, p. 221) referred S. tol to the synonymy of S. tala.
This erroneous action was based on his examination of one of the specimens of
A.6587. Smith apparently was unaware of the existence of other material
labeled as types of S. tala, and fortunately did not designate a lectotype.

Although Weber and De Beaufort (1931, p. 282) and Smith (1970, p. 219)
considered the species insufficiently described for positive identification, the
following statements given in the original description of Chorinemus tala leave
no doubt as to its specific identity: (1) “Chorinemus tala differs [from the

200 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Ficure 3. Scales of Scomberoides species (from area between anal-fin origin and lateral
line): a, S. commersonianus, TABL 107715, 398 mm. FL; b, S. lysan, TABL 107364, 412 mm.
FL: c, S. tala, TABL 107712, 370 mm. FL: d, S. tol, TABL 107714, 410 emma Hees seale:
20.0 mm.

preceding species, C. commersonianus | by a smaller gape, a maxillary that hardly
extends beyond the posterior border of the orbit. (2) The maxillary is, in
addition, covered by the suborbital and only shows its posterior third; it is
enlarged more posteriorly and is cut off more squarely. (3) The teeth are
stronger in proportion. (4) The curve of the snout is a little less straight .. . ,
but the shape of the body, the scales, the lateral line and the number of rays
are as in the preceding species.”

Statement 4 and the fact that S. tala was compared with S. commersonianus,
suggest that the name S. tala applies to a deep-bodied species with high fin
lobes. Scomberoides tol and S. lysan are both relatively slender species (fig. 6)
and have shorter fin lobes (fig. 7) than do the other two species of Scomberoides.
Statements 1 and 4 exclude S. tol, in which the maxilla does not reach the pos-
terior border of the orbit and the scales are needle-shaped (fig. 3d). Statement
3 excludes S. lysan and S. tol, in which the teeth in both jaws are relatively
small in adults and applies best to the species here recognized as S. tala, which
has the largest teeth (fig. 4b). Statements 1 and 2 are sufficient to exclude
S. commersonianus (compare figs. 2a and 2b).

The fact that Cuvier recognized S. commersonianus (and several of its
synonyms) and described as new, from Malabar specimens, examples of the
three remaining valid species of Scomberoides leaves no doubt that he was able

VoL. XXXIX] SMITH-VANIZ & STAIGER: CARANGIDAE 201

10 mm

Ficure 4. Dentary teeth of adult Scomberoides species: a, S. commersonianus; b,
S. tala, c, S. lysan, d, S. tol. Data as in figure 2. Teeth of inner row shown in black.

to distinguish them. It is obvious from his description of S. fala that he would
not have confused it with S. tol. At some time type labels or MNHN catalog
numbers must have become inadvertently mixed. Since the description of S. tol
was based in part on specimens from Malabar, the two specimens of that species
labeled as types of S. tala are here designated putative paralectotypes of Scom-
beroides tol (Cuvier).

Russell’s poor figure of ‘“toloo parah,” upon which the original description
of Chorinemus toloo is based, cannot be positively identified as S. tala, although
it most closely resembles that species. Weber and De Beaufort (1931) included
S. toloo in their synonymy of Scomberoides tala; the figure of Chorinemus toloo
published in Day’s Fishes of India is clearly referable to S. tala.

Jordan and Seale distinguished their new genus and species, Eleria philip pina,
from Scomberoides solely on the basis of strong symphyseal canines on the lower
jaw. They apparently were unaware that such canines are typically present in
the young of S. tala.

The description and figure of Chorinemus hainanensis leave no doubt as to
the identity of this nominal species. In their description, Chu and Cheng em-
phasized the absence of spots on side of body. Like a number of authors, they
were not cognizant of the fact that these spots frequently fade completely.

DEScRIPTION. Characters given in the generic description are not repeated
here. Frequency distributions of selected meristic characters are given in table
2. Dorsal-fin rays VI-VII + I, 19-21; anal-fin rays IJ + I, 17-19; pectoral-

202 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

110

100

COMMERSONIANUS
90 TALA
LYSAN
TOL

80

=>
=> 70
==
S60
za
LJ
—J
ae
<q
=
o 40
lJ
QO
oO
=)

30

20

) 10 20 30 40 50 60 70 80
SNOUT- POSTORBIT LENGTH (MM)

Ficure 5. Relationship of upper jaw length to snout-postorbit length in Scomberoides
species. Dashed lines represent upper jaw length as percent snout-postorbit length.

VoL. XXXIX] SMITH-VANIZ & STAIGER: CARANGIDAE 203

260
s
240
s
s
| s
220 .
COMMERSONIANUS a
TALA .
ai LYSAN =n
= TOL .
= 180 .
of = 2
N Oo
aq 160 .
o
o
2 2 7 26%
4
w 140 . ay a4
fa) eat RI ®
Oo s.
Oo nA
ie e
Z 120 "ome “Ns
— ie) = ef e
© ote “fae ee
= - .? “es,
= 100 stig of Pes ?
ro) 7108 So
Care ave
i ZA Eo)
s y »
sid of a eer@on es
oO q™ s ge 0585 5°
fo}
lu ae Ve %
a dig
60 af 5
BF epeasoo
rs ete
Fo
40 fo)
Past
ot
°
20 “EP
ff
° l
° 100 200 300 400 500 600 700 800 900

FORK LENGTH ~ (MM)

Ficure 6. Relationship of body depth to fork length in Scomberoides species. Dashed
line represents depth as percent fork length.

fin rays I, 16-17. Gillrakers 1-3 + 7-11 = 11-15. Upper jaw extending slightly
beyond posterior margin of eye in adults (fig. 2b). Scales lanceolate (fig. 3c).
Inner row of dentary teeth distinctly larger than outer row teeth in adults (fig.
4b); dentary typically with one or two pair of symphyseal canines in young;
young with dentary teeth of outer row numerous and considerably more closely
spaced than inner row teeth.

Proportional measurements apply only to specimens more than 150 mm.
fork length (FL), and are in percent FL unless otherwise stated. Depth (origin
Ds to As) 29.0-34.9; height dorsal-fin lobe 14.6-18.0; height anal-fin lobe
13.1-16.2; upper jaw 54.3-61.6 percent head length; snout 26.1—30.1 percent
head length.

204 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

160

140

(MM)

COMMERSONIANUS
TALA

LYSAN
120 TOL

LOBE

100

80

60

40

20

HEIGHT SECOND DORSAL-FIN

° 100 200 300 400 500 600 700 800 900

FORK LENGTH (MM)

Ficure 7. Relationship of dorsal-fin lobe height to fork length in Scomberoides species.
Dashed line represents dorsal-fin lobe height as percent fork length.

PIGMENTATION. Sides with 4 to 8 vertically elongate blotches, most of them
intersecting lateral line. Dorsal and anal fins uniformly pigmented. Life color-
ation not observed.

DISTRIBUTION. Widely distributed throughout the Indo-Australian Archi-
pelago, and the Bay of Bengal in the Indian Ocean. Although more collections
are needed, the apparent absence of the species from the western Indian Ocean,
Red Sea, and Persian Gulf appears to be real. Scomberoides tala is apparently
restricted to neritic waters, and does not occur east of the New Guinea-Solomon
Islands region.

MATERIAL (110 specimens, 31—615 mm. FL from 45 collections). CEYLON:
TABL 107365 FJS 69-27 (2: 295-464), S-V 69-77 (1: 560), S-V 69-156 (1:
116, cleared and stained), TABL 107712 CCK 69-43 (12: 244-490), CCK
69-45 (3: 517-567), CCK 69-53 (4: 265-474), CCK 69-54 (2: 197-565),
CCK 69-89 (8: 366-483), CCK 69-94 (1: 505), CCK 69-104 (2: 529-551),
CCK 69-111 (1: 568). INDIA: Malabar, MNHN A. 6588 (315, lectotype

Vor. XXXIX] SMITH-VANIZ & STAIGER: CARANGIDAE 205

of Chorinemus tala); Madras, AMS B. 8124 (1: 197); Calcutta, AMS B. 8093
(1: 115). BURMA: Maungmagan Island, SU 33635 (2: 91-95). ANDAMAN
ISLAND: AMS I. 68 (1: 57). GULF OF THAILAND: GVF station 57-90
(1: 87), GVF station 57-99 (1: 167), GVF-1470 (4: 185-214), GVF-1503
(1: 176), TABL 107040 GVF-1512 (5: 190-219), GVF-2195 (1: 45), GVF-
meio: 31-71), GVE-2203 (1: 131), GVF-2222 (3: 183-198), GVE-2497
(2: 159-161), GVF-2540 (1: 345), GVF-2546 (12: 219-275). MALAYSIA:
Penang, TABL 107266 FJS 69-79 m. (3: 168-183). PHILIPPINES: Luzon,
USNM 65339 (1: 265), 149803 (3: 110-132), 149804 (1: 80); Batangas
Prov., CAS uncataloged (6: 135-172); Manila Bay, CAS uncataloged (1: 165);
Leyete, USNM 149805 (1: 178), 149811 (1: 178); Mindanao, USNM 55990
(1: 271); Matnog Bay, USNM 194913 (1: 218); Balabac Island, AMS I. 10478
(1: 220); Southern Negros, USNM 51945 (98, holotype of Eleria philippina),
zon ho 112), SU 9131 (2: 103-111). JAVA: USNM 72616 (2: 157=161),
72617 (1: 118), 72618 (1: 109), 72619 (1: 114). AUSTRALIA: Northern
Territory, Port Darwin, AMS I. 5291-2 (2: 76-100), IA. 3639 (1: 67); Gulf
of Carpentaria, Groote Eylandt, USNM 173957 (5: 317-339); Queensland, Cape
York, BMNH 1879.5.14.282 (1: 381); Fitzory River Estuary, AMS IB. 1249
(io )e NEW BRITAIN: 4°19’S. 152°1’E., GVF station HK-130 (1: 615).
SOLOMON ISLANDS: Bougainville, AMS IB. 1458 (1: 219).

Scomberoides lysan (Forskal).
(Figures 1c, 2c, 3b, 4c, 5—7, 19; tables 2-3.)

Scomber lysan ForsKAL, 1775, p. 54 (type locality Djeddae and Lohajae, Red Sea).

Scomber forsteri BLocH and SCHNEIDER, 1801, p. 26 (type locality Pacific Ocean [based on
Scomber maculatus Forster MS]).

Lichia tolooparah RUPPELL, 1828, p. 91 (type locality Massawa, Red Sea).

Chorinemus sanctipetri Cuvier in Cuvier and Valenciennes, 1831, p. 379 (279) (type locality
Malabar coast).

Chorinemus moadetta EHRENBERG in Cuvier and Valenciennes, 1831, p. 382 (282) (based
on an unpublished drawing by Ehrenberg of a fish from the Red Sea).

Chorinemus mauritianus Cuvier in Cuvier and Valenciennes, 1831, p. 382 (282) (type locality
l’Isle-de-France [=Mauritius Island]).

Scomber maculatus Forster, 1844, p. 195 (type locality Tahiti).

Chorinemus orientalis TEMMINCK and SCHLEGEL, 1844, p. 106, pl. 57, fig. 1 (type locality
Nagasaki, Japan).

Thynnus moluccensis GRoNoviuS in Gray, 1854, p. 121 (type locality Insulas Moluccenses).

Scomberoides formosanus OSHIMA (non Wakiya), 1925, p. 349, pl. 1, fig. 1 (type locality
Toko, Formosa).

Scomberoides oshimae WHITLEY, 1951, p. 65 (replacement name for Scomberoides formosanus
OsHIMA, 1925, preoccupied).

NOMENCLATURE. The description of S. /ysan is inadequate for positive identi-
fication, but the following statements given by Forskal suggest that he did not
have S. commersonianus (= ‘lysan’ of recent workers): (1) scales lanceolate

206 CALIFORNIA ACADEMY OF SCIENCES [Proc, 4TH SER.

(in “SS. lysan” they are usually referred to as ovate or broadly lanceolate) ;
(2) “about the lateral line there are obsolete | ?indistinct|] dark spots”; (3)
dentes [teeth] numerous, small. Ruppell (1828), who set out especially to
clarify the identities of Forskal’s Red Sea fishes, applied the name ‘lysan’ to
S. commersonianus. His application of the name, however, was based on the
fact that native fishermen referred to that species as “lysan.” Smith (1968,
p. 174) has clearly shown that native names of Red Sea fishes were often not
specific. Cuvier and Valenciennes (1831) did not believe the description of
S. lysan was sufficiently diagnostic for positive identification. They did, how-
ever, state that since Forskal’s description contained no mention of a dark
blotch in the second dorsal fin, they were hesitant to refer it to their Chorinemus
moadetta. Ginther (1860) and all subsequent workers have regarded S. lysan
as a senior synonym of S. commersonianus.

Klausewitz and Nielsen (1965, pl. 25) have published an excellent photograph
and radiograph of the 216 mm. SL lectotype (ZMC 44) of Scomber lysan, re-
produced without comment in Smith (1970). The most distinctive feature of
the lectotype is the position of the maxilla, which ends below the middle of
the eye. It is also apparent from the radiograph that the snout has been pushed
in on the dried skin, giving the snout an atypically blunt appearance. Since the
maxilla extended well beyond the posterior margin of the orbit in all our spec-
imens, we suspected that the ‘/ysan’ of recent authors was not the same species
as the lectotype. Accordingly we wrote Dr. Jérgen Nielsen and asked him to
examine the lectotype for us. The following information, largely provided by
Dr. Nielsen, is sufficient to identify the lectotype as conspecific with S. tolo-
oparah of recent workers: (1) Although there are no teeth left in the outer
dentary row, judging from the sockets the missing teeth were the same size and
number as those of the inner row, which excludes S. tala (see fig. 4); (2) com-
parison of the premaxillary teeth also excludes S. tala; (3) there are no sym-
physeal canines left. Symphyseal canines are never present in S. lysan, and
usually are still retained in specimens of S. commersonianus and S. tala at this
size; (4) the length of the upper jaw is 21 mm.; the dentary is 30 mm. The
1:1.5 ratio between the length of the upper jaw and dentary agrees fairly well
with similar sized specimens of S. tolooparah and is sufficiently different to ex-
clude S. commersonianus; (5) the scales of the paralectotype and the lectotype
are the same size and form. Comparison of a small patch of skin (scales included)
removed from the paralectotype just anterior to the spinous dorsal fin with
scales from the same area of the four species of Scomberoides is sufficient to
exclude S. tol, S. commersonianus, and, with less confidence, S. fala. The most
diagnostic feature of these scales is their strongly lanceolate posterior margins
(see fig. 3).

Unfortunately, the name S. /ysan (applied to S. commersonianus by recent

VoL. XXXIX] SMITH-VANIZ & STAIGER: CARANGIDAE 207

authors) must be retained as a senior synonym of the widely used S. tolooparah.
It would be difficult to justify suppression of the name, however, since it is
not simply an overlooked synonym of S. tolooparah. Scomberoides forsteri is
also a senior synonym of S. tolooparah; and S. sanctipetri, a junior synonym,
has frequently been recognized as a valid species. Although carangids are com-
mercially important fishes, most genera have not been adequately revised, and
consequently many of the names currently in use may eventually prove to be
synonyms.

Although the descriptions of Scomber forstert and S. maculatus are poor,
they apply best to S. lysan. Since the type locality of S. maculatus (and thus
that of S. forsteri as well) is Tahiti, an area from which only S. lysan is known,
there can be little doubt as to the specific identity of either nominal species.
The characters given for Chorinemus moadetta, and comparison of it in the
original description with Forskal’s S. lysan, leaves no doubt that it too is a
junior synonym of S. lysan.

Smith (1970) presents photographs of the holotypes of Lichia tolooparah
(NFIS 386) and Chorinemus sanctipetri (MNHN A. 6587). They are both
clearly synonyms of S. lysan. Likewise, the original descriptions and figures of
Chorinemus orientalis and Scomberoides formosanus agree well with S. lysan.
The description and insular type locality of Chorinemus mauritianus are also
sufficient to identify it as conspecific with S. lysan.

We have not examined the type of Thynnus moluccensis, but according to
Wheeler (1958, p. 220) it is a synonym of Chorinemus sanctipetri (= S. lysan).
The 142 mm. SL holotype bears the catalog number BMNH 1853. 11.12.29.

DEscRIPTION. Characters given in the generic description are not repeated
here. Frequency distribution of selected meristic characters are given in table
2. Dorsal-fin rays VI-VII+I, 19-21, anal-fin rays II-I, 17-19; pectoral-fin
rays I, 16-17. Gillrakers 3-8 + 15-20 = 21-27. Upper jaw extending to or
slightly beyond (in large adults) posterior margin of eye (fig. 2c). Scales lan-
ceolate (fig. 3b). Inner and outer row of dentary teeth subequal in adults (fig.
4c); dentary without symphyseal canines; young with dentary teeth of outer
row numerous and considerably more closely spaced than inner row teeth.

Proportional measurements apply only to specimens more than 150 mm.
fork length (FL), and are in percent FL unless otherwise stated. Depth (origin
Ds to Ag) 20.9-27.3; height dorsal-fin lobe 9.1-14.2; height anal-fin lobe 7.7—
11.6; upper jaw 50.3-54.6 percent head length; snout 27.8-31.8 percent head
length.

PIGMENTATION. Sides of adults with a double series of 6 to 8 dusky round-
ish blotches above and below lateral line, occasionally connected by narrow
isthmus. Distal half of dorsal-fin lobe abruptly and heavily pigmented; anal-
fin lobe white or pale yellow, often with interradial membranes of several

208 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

anterior rays black, especially in large individuals. In life, body gray-green
above, silvery gray to midline, silvery white below.

DISTRIBUTION. Widely distributed throughout the Red Sea, Indian Ocean,
Indo-Australian Archipelago and central Pacific Ocean including Hawaii and
the Marquesas Islands. Scomberoides lysan is the only species of Scomberoides
that occurs east of the Solomon Islands and frequents entirely oceanic islands
in the Indian Ocean, e.g., Seychelles and Chagos Archipelago. The other species
of Scomberoides are generally restricted to neritic habitats. This suggests a
fundamental difference in the biology of S. lysan.

MATERIAL (276 specimens, 20-585 mm. FL from 109 collections). TAN-
ZANIA: Zanzibar, TABL 105602 (1: 281). MADAGASCAR: USNM 171034
(1: 336), TABL uncataloged station JR-68 (1: 54). MAURITIUS ISLAND:
USNM 118407 (13: 256-403). COMORO ISLANDS: TABL uncataloged
station FT-9 (2: 45-46), TABL uncataloged station HA-8 (1: 123), HA-10
(1: 52). ALDABRA ISLAND: TABL uncataloged station HA 67-56 (1: 385).
CHAGOS ARCHIPELAGO: Diego Garcia Atoll, TABL uncataloged station
HA 67-42 (2: 83-113), HA 67-43 (3: 146-154). INDIA: Malabar, MNHN
A. 5893 (480, holotype of Chorinemus sanctipetri); Madras, AMS B. 8091 (3:
110-128). CEYLON: TABL 107364 FJS 69-27 (2: 412-523), S-V 69-123
(8: 177-303), S-V 69-128 (2: 123-161, cleared and stained), S-V 69-152 (9:
278-355), PCH 69-201 (1: 541), PCH 69-224 (1: 279), PCH 69=225 (1: 381),
PCH 69-228 (1: 585), CCK 69-9 (1: 525), CCK 69-49 (4: 467-530), CCK
69-53 (1: 339), CCK 69-110 (1: 343), CCK 69-113 (2: 370-535), CCK 69—
130 (1: 555). SOUTH CHINA SEA: Hainan Island, USNM 94780 (1: 178).
TAIWAN: USNM 192563 (1: 194). OKINAWA ISLAND: USNM 72093
(2: 125-211). PHILIPPINES: Luzon, USNM 139573 (2: 33=38)pei49omo7
(1: 47), 149800 (1:31), 168276 (1: 215), 168277 (12 220), 1OlSS5a( 2 aera
191888 (11: 45-73), 191889 (11: 83-103), 191891 (6: 76-116), 191894 (4:
44-100), 191896 (2: 200-213), 191907 (1: 76), 191913 (1: 54); Burias Island,
USNM 194916 (1: 232); Busanga Island, USNM 191884 (3: 42-56); Lina-
capan Island, USNM 191893 (2: 66-92); Pandanon Island, USNM 191910
(2: 89-97); Mindoro, USNM 149799 (3: 66-79), 191902 (1: 73), 191908
(1: 96); Samar, USNM 191859 (4: 190-196); Leyete, USNM 191886 (11:
64-188); Palawan, USNM 191887 (19: 26-187), 191897 (4: 81-185), 191905
(1: 39); Cuyo Island, USNM 191911 (2: 67-81). BORNEO: Sandakan Bay,
USNM 191901 (3: 83-109). AUSTRALIA: Northern Territory, Bedwell Point,
AMS IA. 7679-80 (2: 136-200); Gulf of Carpentaria, Sir Edward Pellew Group,
AMS IA. 1488 (1: 129), TA. 2559 (1: 144); Queensland, AMS A. 12495 (1:
141); Cape York, AMS TI. 9712-4 (3: 206-217); Barrier Reef, USNM 176914 (6:
129-194); Townsville, AMS IA. 2324-5 (2: 150-164); Lindeman Island, AMS
TA. 6288 (1: 290), TA: 6289-90 (2: 112=158), TA. 6647 (22 TIO=128) Rei

VoL. XXXIX] SMITH-VANIZ & STAIGER: CARANGIDAE 209

7498-7501 (4: 51-75), IA. 7887 (1: 88); Hayman Island, AMS IA. 6126-7
(4: 35-132); Burdekin River, I. 5314-5 (2: 175-187), A. 18302 (1: 157); New
South Wales, Sydney, MKT AMS IB. 4232 (1: 290), USNM uncataloged sta-
tion BBC 1464 (1: 295); Gafton, AMS I. 15594—001 (1: 65). NEW GUINEA:
Aroma, AMS IA. 5738 (3: 33-69); Hula Village, AMS IB. 1706 (1: 160);
Bat Island, Purdy Archipelago, AMS IB. 1523 (1: 82). NEW BRITAIN: GVF
station HK-29 (1: 297), GVF station HK-130 (1: 418). SOLOMON ISLANDS:
Bougainville, TABL uncataloged Te Vega Expedition, Cruise 6, station 247 (1:
155); Ontonga Java Island, AMS IA. 3696 (1: 355). PALAU ISLAND: GVF-
546 (1: 81), GVF-591 (1: 83), GVF-1971 (2: 189-205), GVF station 59—708
(11: 373-442), GVF station 59-710 (2: 369-410), GVF station 59-724 (2: 365—
379). MARIANA ISLAND: Saipan, USNM 154647 (1: 54). MARSHALL IS-
LANDS: USNM 66022 (1: 40), 66024 (1: 37), 66025 (1: 30), 142040 (1: 580),
142041 (2: 30), 142042 (2: 28-32), 163134 (1: 26), 165648 (2: 78-80), 179433
(1: 63). GILBERT ISLANDS: USNM 66021 (1: 60); Onotoa, USNM 167428
(2: 380-415); Arorae, AMS IB. 6618 (1: 25); Nauru, AMS TA. 6999 (1: 375).
ELLICE ISLAND: AMS TIA. 5531 (1: 215); Funafuti, USNM 66020 (1: 96),
AMS I. 3562 (1: 495). PHOENIX ISLANDS: Canton Island, USNM 115330
(t= 456). SAMOA ISLANDS: Apia, USNM 52361 (3: 219-240). NEW
HEBRIDES ISLANDS: Malehuka, AMS I. 11294 (1: 162). FIJI ISLANDS:
Suva, USNM 66023 (2: 176-177). TONGA ISLANDS: Vavau, USNM 66019
(1: 103); Tonga, TABL uncataloged, Te Vega Expedition, Cruise 8, station
g06m(2-9215-227). SOCIETY ISLANDS: Tahiti, GVF-1155 (1: 324),GVF
station 57-6 (1: 296). TUAMOTU ARCHIPELAGO: Rangiroa, USNM 66018
(2: 84-135); Takaroa, GVF station 56—?? (1: 299); Raroia, GVF-87 (3: 502-
555). MARQUESAS ISLANDS: Nukahiva, USNM 66016 (1: 416), 66017
(3: 184-215); Hiva Oa, USNM 186329 (2: 33-38). LINE ISLANDS: Pal-
myra Island, GVF station 51-41 (5: 350-419), GVF station 51-44 (4: 140-
157); Christmas Island, USNM 19238 (3: 402-415). JOHNSTON ISLAND:
USNM 26825 (1: 440). HAWAIIAN ISLANDS: Hawaii, USNM 78073
(et 26-69) 982818 (1:2 279), 83429 (1: 227); Oahu, SU 3270 (52 172-220),
figaie(o !03—=190), 53345 (1: 20), USNM 55082 (1: 357), 55371 (1: 141),
88161 (1: 166), 88162 (3: 172-255), 126691 (1: 174), 151615 (3: 191-260).

Scomberoides tol (Cuvier).
(Figures 1d, 2d, 3d, 4d, 5—7, 14d, 19, 20b, 21c, 22c; tables 2-3.)
Chorinemus tol Cuvier in Cuvier and Valenciennes, 1831, p. 385 (type localities Pondichery ;

Malabar coast; Amboine; Ile de Bourou).
Scomberoides formosanus WAxktIya, 1924, p. 236, pl. 38, fig. 3 (type locality Kii, Formosa).

NoMENCLATURE. The original description of Chorinemus tol is sufficiently
diagnostic for positive identification (see also discussion under S. tala). Likewise

210 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

the statement “scales linear, pointed at both ends” given in the description of
S. formosanus could apply only to S. tol Cuvier. F. H. Berry has examined the
130 mm. SL holotype (FMNH 59499), and made his data available to us.

DEscripTION. Characters given in the generic description are not repeated
here. Frequency distributions of selected meristic characters are given in table
2. Dorsal-fin rays VI-VII + I, 19-21; anal-fin rays II + I, 18-20; pectoral-
fin rays I, 15-18. Gillrakers 4-7 + 17-20 = 21-26. Upper jaw does not extend
to posterior margin of eye (fig. 2d). Scales slender, needlelike (fig. 3d). Inner
and outer row of dentary teeth subequal in adults (fig. 4d); dentary typically
with one or two pairs of symphyseal canines in young; young with dentary teeth
of outer row only slightly more numerous and closely spaced than inner row teeth.

Proportional measurements apply only to specimens more than 150 mm.
fork length (FL), and are in percent FL unless otherwise stated. Depth (origin
Dp to Ay) 20.0-24.7; height dorsal-fin lobe 7.7-11.2; height anal-fin lobe 6.3—
8.8; upper jaw 42.6-49.3 percent head length; snout 29.1-32.8 percent head
length.

PIGMENTATION. Sides of adults with 5 to 8 oval or vertically oblong blotches,
the first 4 or 5 of which intersect lateral line. Distal half of dorsal-fin lobe abruptly
and heavily pigmented; anal-fin lobe usually immaculate, white in life. In life,
body white ventrally, bluish dorsally; lateral blotches black, faint or absent
in young.

DistrRIBuTION. Widely distributed throughout the Indo-Australian Archi-
pelago and Indian Ocean. Although we have not examined specimens, the species
has been reliably reported from the Red Sea. Scomberoides tol is apparently
restricted to neritic waters, and does not occur east of the New Guinea-Solomon
Islands region.

Type MatertaL. MNHN A. 6605 (150, here designated lectotype of Chor-
inemus tol) Malabar. Paralectotypes as follows: MNHN B. 2650 (2: 135-147,
removed from A. 6605), B. 5542 (dried skin, not measured) Pondichery, A. 6585
(1: 170) Bourou Island, A. 6620 (1: 119) Amboine. MNHN A. 6587 (2: 296—
302, putative paralectotypes) Malabar (see discussion under S. tala). FMNH
59499 (130 mm. SL, holotype of S. formosanus) Kii, Formosa.

OTHER MATERIAL (142 specimens, 20-468 mm. FL from 66 collections).
SOUTH AFRICA: Durban, TABL uncataloged FJS 69-1 (7: 224-267). MO-
ZAMBIQUE: Ponta Mabonl, TABL uncataloged FJS 69-5 (1: 173); Polana,
TABL uncataloged FJS 69-12 (1: 208); Mecaneta, TABL uncataloged FJS
69-14 (3: 225-232). KENYA: Mombasa, TABL uncataloged Anton Bruun
cruise 9, station FT-2 (2: 271-276). MADAGASCAR: USNM 171055 (1: 185),
TABL uncataloged JR-27 (3: 171-173), JR-70 (1: 120), TABL uncataloged
Anton Bruun cruise 7, Tulear Harbor, (1: 124). PERSIAN GULF: USNM
147819 (1: 20), 148074 (1: 323). INDIA: Bombay, TABL uncataloged FHB

VoL, XXXIX] SMITH-VANIZ & STAIGER: CARANGIDAE 211

66-1 (4: 37-55); Kerala State, Cochin, TABL uncataloged FHB 66-53 (1:
127); Madras State, AMS B. 8045 (1: 171); Cape Comorin, TABL uncataloged
FHB 66-46 (1: 222), FHB 66-49 (2: 264-330); Porto Nova, TABL un-
cataloged FHB 66-17 (2: 282-284). CEYLON: FJS 69-14 (3: 225-232),
TABL 107707 S-V 69-86 (4: 262-284), S-V 69-123 (8: 240-390), TABL
107363 S-V 69-126 (8: 268-450), S-V 69-128 (6: 107—-131.5, 2 cleared and
stained), PCH 69-180 (1: 326), PCH 69-185 (10: 102-107), PCH 69-202
(2105-126), PCH 69-237 (1: 310), PCH 69-266 (4: 358-373), PCH 69-292
(3: 146-154), CCK 69-9 (1: 340), CCK 69-43 (1: 181), CCK 69-73 (1: 395),
CCK 69-91 (1: 242), CCK 69-94 (1: 468), CCK 69-113 (3: 410-426), CCK
69-129 (8: 351-460). MALAYSIA: Penang, TABL uncataloged FJS 69-74
m. (2: 209-219). GULF OF THAILAND: GVF station 13B-C (6: 209-246)
GVF-1541 (7: 141-203), GVF-2546 (8: 269-320), GVF-2655 (2: 117-119).
TAIWAN: USNM 76607 (8: 41-149), SU 7317 (1: 169), 20997 (2: 203-218).
JAPAN: Kagoshima, SU 23774 (3: 86-117). PHILIPPINES: Luzon, USNM
aoe oe lOS—224),, 168275. (1: .207),, 168281 (1: 372), 168282 (12-331),
USNM uncataloged (out of USNM 191896) (1: 115), 191899 (1: 156), 191906
(1: 77), 191909 (1: 89), 191912 (1: 89), AMST. 10539 (1: 215); Leyte, USNM
149796 (1: 38); Cebu, USNM 191900 (2: 123-134); Panay, USNM 149794
(1: 30), 191898 (1: 105); Mindanao, USNM 55995 (1: 244), 58011 (2: 143-
144), 59004 (1: 149), 168283 (1: 348). JAVA: Batavia, USNM 726211 (2:
22-040) 726012 (1: 171), 72613 (1: 145), 72614 (1: 130), 72615 (2: 162=
208). AUSTRALIA: Western Australia, Onslow, WAM P. 15488—-90 (3: 110—
174); Port Hedland, AMS IB. 1638 (1: 128); Gulf of Carpentaria, 16°53’S.,
139°21’E., AMS I. 15557-118 (5: 87-172); Queensland, Barrier Reef, USNM
uncataloged (out of USNM 176914) (1: 171); Palmers Island, AMS I. 2962-3
(2: 90-91) ; Hayman Island, AMS IA. 6373 (1: 221). NEW GUINEA: Sumarai
District, AMS IA. 5740 (1: 235). SOLOMON ISLANDS: Bougainville, TABL
uncataloged Te Vega Expedition, cruise 6, station 246 (2: 43-44); Malaita
Island, AMS I. 15360—056 (1: 350).

Scomberoides spinosus (Smirnov).
(Figure 8.)

Palaeoscomber spinosus SMIRNOV, 1936, p. 49-59, pl. 7, figs. 31-32 (type locality Zuramakent
Horizon of the Upper Maikop deposits, Northeastern Caucasus).

Danil’chenko (1960) has given a detailed redescription of Palaeoscomber
spinosus and convincingly demonstrated that it is a species of carangid, although
originally believed to represent a new genus and subfamily of Scombridae
(Smirnov, 1936). In reassigning P. spinosus to Oligoplites, Danil’chenko em-
phasized the peculiar needlelike scales which he regarded as a “characteristic
sign” of the genus. He did not compare it with Scomberoides tol, apparently

212 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

in (ite: |

Ficure 8. Diagrammatic sketch of Palaeoscomber spinosus Smirnov (after Danil’chenko,
1960).

unaware that its scales are identical to those of Oligoplites. Although we have
not examined specimens of P. spinosus, and therefore do not include a description,
we believe it is a species of Scomberoides closely related to S. tol. The number
of dorsal-fin rays (VI-VII + I, 20) and predorsal bones (3) reported for P.
spinosus apply best to a species of Scomberoides. Likewise, the relatively short
upper jaw (47 to 48 percent HL) and general configuration of the maxilla
suggest S. tol rather than a species of Oligoplites (compare figs. 2d and 8).

The four fossils upon which the description is based are from the Zuramakent
Horizon of the Upper Maikop deposits (Lower Miocene) in the Northeastern
Caucasus. According to Danil’chenko (1960), only 3 of 18 genera examined
from this deposit do not represent recent genera. The recent species Scomber
japonicus Houttuyn is also present. The characters of Oligoplites and its present
distribution (western Atlantic and eastern Pacific) suggest that it evolved
from a Scomberoides-like ancestor that reached the New World via the Pacific
(see discussion under Relationships and Zoogeography Section). It seems more
logical in view of the geographical location and recent nature of the Upper
Maikop fauna that P. spinosus represents a genus present in the Indian Ocean
rather than one that is restricted to the western Atlantic and eastern Pacific
oceans.

Two characters mentioned in the redescription of P. spinosus require com-
ment. (1) The supramaxilla is said to be absent. The small supramaxilla of
Scomberoides could easily be lost or overlooked in a fossil. (2) The vertebral
count is given as 10 + 15. In his discussion of methods Danil’chenko (1960)
stated that the vertebrae were counted as in recent fishes (the urostyle is treated
as a separate vertebra), yet his diagrammatic drawing of P. spinosus (here re-
produced as fig. 8) clearly shows 10 + 16 vertebrae. We believe the count of
10 + 15 is an error. Berry (1968, p. 148) stated that the number of vertebrae
is intraspecifically very constant in all Carangidae. One exception that he noted
was one specimen of Oligoplites saurus inornatus Gill with 10 + 17 vertebrae;

VoL. XXXIX] SMITH-VANIZ & STAIGER: CARANGIDAE 213

19 others had 10 + 16. We have made numerous vertebral counts of all species
of Oligoplites and Scomberoides and except for the specimen mentioned above,
all have 10 + 16 vertebrae.

The available material of Scomberoides spinosus should be carefully com-
pared with similar sized specimens of S. tol. We tentatively regard them as
specifically distinct because several of the proportional measurements given
for S. spinosus do not agree with our specimens of S. tol.

Oligoplites Gill, 1863

Oligoplites GILL, 1863, p. 166 (type species: Scomber saurus BLtocH and SCHNEIDER, 1801,

by subsequent designation of JorDAN, 1917, p. 324).

Leptoligoplites Fowurr, 1944, p. 226 (as a subgenus of Oligoplites; type-species: Oligoplites
refulgens GILBERT and STARKs, 1904, by original designation).

The carangid genus Oligoplites comprises five species of tropical and sub-
tropical marine fishes restricted to the eastern Pacific and western Atlantic
oceans. Only one species, O. saurus, occurs in both oceans. Parenthetically,
it should be noted that in his excellent review of the zoogeographic relation-
ships of tropical American marine shore fishes, Rosenblatt (1967, p. 590) has
perpetuated a misconception by reporting Oligoplites from the eastern Atlantic.
The genus is largely confined to inshore, neritic waters, and the young are
sporadic invaders of coastal freshwater habitats (Gunter and Hall, 1963; Miller,
1966; Gilbert and Kelso, 1971). Breder (1942) has made the interesting ob-
servation that the young of Oligoplites saurus 27 to 34 mm. SL, which appear
on the west coast of Florida during June and July, frequently drift passively
at the surface head-down or twisted in a most unfishlike manner. He noted that
their behavior and general appearance almost perfectly mimicked a floating
dead leaf. The young of Scomberoides lysan have been observed exhibiting
similar behavior (Major, 1973). The largest species of Oligoplites seldom, if
ever, exceed 40 cm. fork length. In Central and South America they are com-
monly sold in fish markets. F. H. Berry im McClane (1965, p. 477) has reported
Oligoplites altus (as O. mundus) to be an exceptionally strong and stubborn
fighter on light tackle.

Oligoplites shows a striking external resemblance to the allopatric Indo-
Pacific genus Scomberoides, and together they constitute the subfamily Scom-
beroidinae (=Chorineminae) of recent authors. Starks (1911) gives a detailed
osteological comparison of these two genera. The adults of Scomberoides have
a series of prominent spots or bars on the sides that is absent in Oligoplites,
although faint bars may occasionally be present in O. saurus. The genus was
last reviewed by Schultz (1945). Several junior synonyms overlooked in that
work are included in our synonymies. The species that we recognize may be

[Proc. 4TH SER.

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VoL. XXXIX] SMITH-VANIZ & STAIGER: CARANGIDAE 215

TABLE 5. Comparison of upper jaw length (expressed as percent HL) and body depth
(expressed as percent FL) in Oligoplites species.

PCO. Upper Jaw Length Depth
Species Range FL N Range x N Range x
altus 35-149 13 59-66 63.4 10 27-32 29.8
palometa 92-147 12 58-63 59.7 12 24-31 27.3
saliens 88-146 7 57-67 62.3 8 26-29 27.0
refulgens 44-148 35 41-46 43.6 36 19-29) @ e199
Saurus saurus 31-144 11 52-57 54.7 10 23-26 23.9
saurus inornatus 53-132 20 52-57 55.3 20 23-25 24.0
$5450 mn: Upper Jaw Length Depth
Species Range FL N Range X N Range x
altus 154-379 65 58-70 64.3 47 27-38 32.0
palometa 160-392 26 59-64 61.6 26 26-34 29.2
saliens 165-390 34 62-68 65.3 33 27-31 28.8
refulgens 150-230 22 42-46 43.5 22 19-22 20.0
Saurus saurus 165-285 26 52-57 54.3 26 24-27 DSP)
saurus inornatus 200-257 7 52-57 54.6 il 25-28 26.8

identified by reference to the key and tables 4-5. We have not included seg-
mented dorsal- and anal-fin ray counts because, as in Scomberoides, they are
not useful in defining species.

NOMENCLATURE. Schultz (1945) considered O. inornatus Gill the “genotype”
of Oligoplites, however, no type-species designation was given in the original
description. We believe that the action of Jordan (1917, p. 324) qualifies Scom-
ber saurus Bloch and Schneider, 1801, as the type-species in accordance with
provision (iv) of Article 68a of the International Code of Zoological Nomen-
clature (1964).

We concur with Schultz (1945, p. 330) that recognition of the subgenus
Leptoligoplites on the basis of a few more gill rakers and elongate body is un-
justified. Differences in dentition would be a much better basis upon which
to establish subgenera if such were needed.

DEscrRIPTION. Body moderately to strongly compressed. Adipose eye-
lids moderately to poorly developed. Body, except head, covered with needle-
like embedded scales. Lateral-line scales not developed into scutes. Lateral line
without side branches, straight except for slight curve over pectoral fin. Anterior
rim of shoulder girdle smooth, without fleshy papillae or deep groove near
isthmus. Premaxillae not protractile, connected anteriorly to snout at midline
by a wide fleshy bridge. Swimbladder strongly bifid posteriorly, length of bi-

216 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

furcated portion of swimbladder equal to or greater than undivided part. Pos-
terior dorsal- and anal-fin rays consisting of semidetached finlets, distal fourth
to half of ray not connected by interradial membrane (unattached portion of
rays increasing with growth); ultimate and penultimate rays more closely
spaced than adjoining rays, fully connected by interradial membrane; none of
dorsal- or anal-fin rays produced into long filaments. Pelvic fins relatively short,
depressible into shallow abdominal grooves. Pectoral fins short, not falcate,
57 to 80 percent head length in adults. Upper jaw extends posteriorly from
middle of eye to well behind orbital rim, 42 to 70 percent head length in adults.

Predorsal bones 4 to 6, rarely 4; dorsal-fin rays IV-VI (rarely VI) + I,
18 to 21; total dorsal-fin rays 23 to 27. Anal-fin rays II + I, 19 to 21. Pectoral-
fin rays I, 15 to 17. Pelvic-fin rays I, 5. Vertebrae 10 + 16 to 17 (rarely 17)
= 26; epipleural ribs 8-10. Branchiostegal rays 3 + 4 or 2% + 4% (epihyal
+ ceratohyal). Upper gillrakers 2 to 8; lower gillrakers 8 to 22; total gillrakers
11 to 29.

Fronto-supraoccipital crest low; temporal crest poorly developed, not ex-
tending forward to posterior rim of orbit and not reaching edge of cranium;
preorbital region short. Epiotics broadly united along midline of cranium in-
ternally. Zygapophyses of exoccipital widely separated beneath foramen mag-
num. Parasphenoid not expanded into a broad, flattened plate posteriorly.
Myodome with broad, posterior opening. Basioccipital without a pair of lateral
processes on ventral surface. Ascending process of premaxilla short and tri-
angular; maxilla long and slender, rather closely attached to premaxilla. Pre-
maxillary spur absent. Supramaxilla absent. Interosseous space between dorsal
arm of dentary and upper margin of articular minute or absent. Well developed
teeth on dentary, premaxilla, palatines, and vomer; two distinct rows of dentary
teeth (see discussion under Relationships and Zoogeography Section); sym-
physeal premaxillary and dentary teeth of juveniles not robust or caniniform.
Mesopterygoids edentate. Pharyngeals not noticeably enlarged, covered with
sharp, pointed teeth.

Suborbital shelf absent; lower and posterior suborbitals expanded poste-
riorly, especially in large specimens, thus covering a large portion of cheek.
Basibranchials 1-3 usually each with a pair of large tooth plates. Lower limb
of posttemporal not noticeably short or thickened. Postcoracoid process absent.
Interpelvic keel well developed; postpelvic process of each side coalesced entire
length, not forming an apical fork. Inferior vertebral foramina absent. First
anal pterygiophore and haemal spine of first caudal vertebra firmly attached to
form a strong strut. Anal pterygiophore expanded anterolaterally to form roof
over first two anal spines; first two anal spines articulated proximally with each
other; second anal spine asymmetrical, with deep groove on one side of anterior
face. Caudal skeleton with 2 epurals and 2 pairs of uroneurals.

VoL. XXXIX] SMITH-VANIZ & STAIGER: CARANGIDAE 217

Key To Spectres oF Oligoplites

la. Upper jaw 41 to 46 percent HL; body depth 18 to 22 (mean 20.0) percent FL; head
length usually equal to or greater than body depth; specimen figure 10a
Pan Sra Nn eles a O. refulgens (Pacific)
1b. Upper jaw 52 to 70 percent HL; body depth 23 to 38 percent FL; head length less
(Haver |oyaya hy “YoU Ey Oy 1 See se ao ee ede Do oe ne Roh Oe a YE al oe 2
2a. Ventral profile of dentary strongly convex (fig. 9c); premaxillary teeth in a single row,
somewhat irregular in young; gillrakers of lower limb of first gill arch 17 to 20 (mean
18.8) ; first dorsal spines consistently 4; specimen figure 9c O. saliens (Atlantic)
2b. Ventral profile of dentary moderately convex; premaxilla with 2 distinct rows of teeth
or a band of villiform teeth; gillrakers on lower limb of first gill arch 8 to 18 (11 to
1S tin A eiM(e Kpoyeees)) 8 aanaste GloveseNl Goynavess 2 (0) @ 3
3a. Upper jaw 52 to 57 (mean 54.7) percent HL; body depth 23 to 28 percent FL; gillrakers
on lower limb of first gill arch 13 to 18; premaxillary teeth essentially in 2 distinct rows,
an additional row of minute teeth may occur anteriorly; first dorsal spines typically
SeEspecimen, tieures 9aand, (Ob) 22 O. saurus (Atlantic and Pacific)
3b. Upper jaw 58 to 70 percent HL; body depth 24 to 38 percent FL; gillrakers on lower
limb of first gill arch 8 to 14; premaxilla with a band of villiform teeth, wide anteriorly;
HITS PEG OLSAIESDINCSRAmONG OM eee tL a a oe ee ee 4
4a. Top of head with numerous sensory pores, which are connected by network of tubes
wisible) beneath skin specimen) figures lOc, d@ 2 O. altus (Pacific)
4b. Sensory pores on top of head sparse, never aS numerous as in specimens of O. altus;
SHECIMENMMC MTCRG Due ee = mes ee ee O. palometa (Atlantic)

Oligoplites refulgens Gilbert and Starks.
(Figures 10a, 11, 19; tables 4-5.)

Oligoplites refulgens GILBERT and STARKS, 1904, p. 73, pl. 11, fig. 19 (type locality Panama
Bay).

This Eastern Pacific species is easily distinguished from its congeners by
its short upper jaw and slender body (see table 5). From those species with
which it occurs sympatrically, it is also distinguished by its higher number of
gill rakers. On the basis of its premaxillary dentition, relationship of predorsal
bones to first dorsal pterygiophores, and upper jaw development, it appears to
be most closely related to O. saurus.

Oligoplites refulgens is the smallest species of the genus, rarely exceeding
250 mm. FL. It occurs in the Gulf of California north at least to Kino Bay,
Sonora, Mexico and south to the Gulf of Guayaquil, Ecuador.

MatTERIAL (61 specimens, 38-230 mm. FL, from 14 collections). GULF OF
CALIFORNIA: TABL 103470 (1: 148.5), 103475 (2: 113-131, cleared and
stained), 103477 (2: 162), SIO 59-245 (1: 170). GULF OF PANAMA: TABL
103473 (1: 117), 103474 (2: 186-189), 103476 (4: 116-132), 107711 (4: 164—
175), USNM 81986 (10 of 36: 38-82). COLOMBIA: TABL 103472 (6: 89-
OA 107708 (1-230). ECUADOR: TABL 103471 (12209), TABLE 103478
Cismotss 15-157): PERU: Zorritos, TABL 107710 @4=z 150=196);

218 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Ficure 9. Western Atlantic Oligoplites: a, O. s. sauwrus, TABL 101582, 225 mm. FL;
b, O. palometa, USNM 121804, 245 mm. FL; c, O. saliens, TABL 101351, 204 mm. FL.

Oligoplites saliens (Bloch).
(Figures 9c, 11, 14f, 19, 24c; tables 4-5.)
Scomber saliens BirocH, 1793, p. 49, pl. 335 (type locality Antilles).

?Scomber calcar BLocH, 1793, p. 55, pl. 336, fig. 2 (type locality Accra, coast of Guinée).
Scomberoides saltator LACEPEDE, 1802, p. 50 (based on unpublished manuscript of Plumier).

Schultz (1945) did not include Scomber calcar in his synonymy of O. saliens.
Bloch’s illustration shows what appears to be a species of Oligoplites, with a
relatively deep body and 3 first dorsal spines. Since Bloch’s description was
based on a small specimen, Regan (1903, p. 349) is probably correct in stating
that it is ‘“‘without much doubt, based on a young example of Scomberoides
| =Oligoplites| saliens, B\.”. The eastern Atlantic type locality would seem to

VoL. XXXIX] SMITH-VANIZ & STAIGER: CARANGIDAE 219

See,

Ficure 10. Eastern Pacific Oligoplites: a, O. refulgens, SIO 59-245, 170 mm. FL;
b, O. saurus inornatus, TABL 103480, 200 mm. FL; c, O. altus UCLA w58-304, 195 mm. FL;
d, O. altus (‘mundus’ form), UCLA w58-304, 180 mm. FL.

220 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

ALTUS
PALOMETA

SAURUS

o
e
SALIENS wx
°
REFULGENS #8

UPPER JAW LENGTH (MM)

19) 10 20 30 40 50 60 70 80 90 100

HEAD LENGTH (MM)

Ficure 11. Relationship of upper jaw length to head length in Oligoplites species. Dashed
lines represent upper jaw length as percent head length.

exclude S. calcar representing a species of Oligoplites, but the type locality is
possibly in error. C. L. Smith has documented a number of such errors (Smith,
1971, pp. 165 and 208) involving other species described by Bloch.

The most diagnostic character of O. saliens is the strongly convex ventral
profile of its massive lower jaw (see fig. 9c). It is the most specialized species
of Oligoplites and does not appear to be closely related to any other species.
It differs in having a 5 + 4 arrangement of predorsal bones and first dorsal
pterygiophores (see table 4), a single row of enlarged premaxillary teeth, and
a relatively constant number of outer dentary teeth (see fig. 19) that do not
exhibit any recognizable ontogenetic changes. Oligoplites saliens can also be
distinguished from the other two Atlantic species of Oligoplites by its higher
number of gillrakers on the lower limb of the first gill arch.

Oligoplites saliens is a large species, attaining at least 390 mm. FL. In the
western Atlantic it ranges north at least to Cartasca Lagoon, Honduras, and south
to Rio de Janeiro, Brazil.

MATERIAL (42 specimens, 88-390 mm. FL, from 25 collections). HON-
DURAS: Caratasca Lagoon, TABL 101351 (11: 185-219), 101353 (2: 183.5—
207), 104795 (5: 195-208). COLOMBIA: UMML 27265 (1: 122). GULF

VoL. XXXIX] SMITH-VANIZ & STAIGER: CARANGIDAE 221

OF VENEZUELA: TABL 103213 (1: 390), USNM 123073 (1: 88), 123075
(1: 94). TRINIDAD: TABL 104796 (1: 203). GUYANA: TABL 104330 (1:
meme 04378 (1: 235), 107716 (1: 165), 107717 (1: 177). SURINAM: USNM
uncataloged R/V Oregon station 4175 (1: 169). BRAZIL: Rio Para, SU 53055
(i201), 53062 (1: 249), 53063 (1: 315), 53064 (1: 165); Vitoria, SU 66973
(3[2 cleared and stained]: 95-133), 66975 (1: 146), 66976 (1: 145); Rio de
Janeiro, SU 53056 (1: 215), 53057 (1: 270), USNM 76330 (1: 194), 100824
(#7503), 100825 (1: 295).

Oligoplites saurus (Bloch and Schneider).

(Figures 9a, 10b, 11, 14e, 18, 19, 20c, 21d, 22d; tables 4-5.)

Scomber saurus BLocu and SCHNEIDER, 1801, p. 32 (type locality Jamaica).

?Lichia quiebra Quoy and GAIMARD, 1824, p. 365 (type locality Brazil).

Chorinemus saltans Cuvier in Cuvier and Valenciennes, 1831, p. 393 (289) (type locality
Martinique).

Chorinemus lanceolatus Girard, 1858, p. 168 (type locality Joseph’s Island, Texas).

Chorinemus occidentalis GUNTHER, 1860, p. 475 (type localities Jamaica; San Domingo;
Trinidad; Puerto Cabello; Bahia).

Oligoplites inornatus GILL, 1863, p. 166 (type locality west coast of Panama).

Oligoplites rathbuni MiranpA-RiBeEtRO, 1915, p. 8 (type locality Bahia Fish Market, Brazil).

Schultz (1945) apparently overlooked the descriptions of Chorinemus saltans
and C. lanceolatus. The descriptions of both species and their type localities are
sufficient to identify them positively as synonyms of O. saurus. We have
examined the 102 mm. holotype (USNM 710) of C. lanceolatus.

Centronotus argenteus Lacépéde (1802, p. 316) was questionably included
by Schultz in the svnonymy of O. s. saurus. The name, however, does not appear
to be available nomenclaturally, and we regard it as a nomen nudum. No de-
scription was given and species belonging to two different genera were listed as
synonyms.

Oligoplites inornatus was not compared with O. saurus in the original de-
scription. Despite the fact that these two nominal species were described from
different oceans, recent workers have regarded them as only subspecifically
distinct at best. The only characters that appear to distinguish them are a
slightly higher number of gillrakers and a few more premaxillary teeth ante-
tiorly in Pacific specimens. Certainly the slight differences shown between these
two nominal species are not of sufficient magnitude to justify specific separation.
However, until large series of specimens from throughout the range of both
populations are compared, we believe it is best to maintain them as subspe-
cifically distinct.

The best characters by which to distinguish O. saurus are given in the key
to Oligoplites species. Berry in McClane (1965, p. 476) reported that the two
detached anal-fin spines of O. s. inornatus apparently are connected with toxic

bo
bo
bo

CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

glands. In south Florida O. saurus is commonly referred to by commercial fisher-
men as stinging jack. We know of no report of ‘stinging’ characteristics in
any other species of Oligoplites.

Although O. saurus attains at least 285 mm. FL, it seldom exceeds 250 mm.
In the western Atlantic it occurs as far north as the Gulf of Main (Bigelow
and Schroeder, 1953, p. 381) and south at least to Recife, Brazil. It is the only
species of Oligo plites that occurs along the coast of the United States and through-
out most of the Caribbean. It has not, however, been reported from the Bahama
Islands, perhaps because of the absence of suitable estuarine habitats. In the
eastern Pacific O. saurus inornatus occurs in the Gulf of California at least to
Concepcion Bay, along the outer coast of Baja California to Magdalena Bay,
and south to Esmeraldas, Ecuador, and the Galapagos Islands.

MATERIAL. The specimens of O. saurus which we have studied are listed
under the two subspecies as follows:

Oligoplites saurus saurus (71 specimens, 31-285 mm. FL, from 30 col-
lections). GEORGIA: TABL 102643 (1: 191), 103197 (1: 23G)R@77en
(2: 92-98, cleared and stained). FLORIDA: UMML 39 (1: 67), 3144 (5:
51-93), 4273 (2: 47-139), 4436 (2: 90-92), 5114 (1: 116), 5628 (1: 60),
5975 (1: 73), 9597 (2: 165-185), 13123 (1: 253), 29468 (4: 211-250), BABE
102682 (1: 188), 104730 (9: 101-128), 105386 (3: 196-285), TABL uncata-
loged station 67-258 (4: 126-168), station 67-280 (3: 131-152). MISSIS-
SIPPI: TABL 102644 (1: 202). LOUISIANA: TABL) 1026835 G=saoie
TEXAS: USNM 710 (102, holotype of Chorinemus lanceolatus). JAMAICA:
TABL 101582 (4: 208-225). MARTINIQUE: TABL 103469 (1: 206),
105791 (3: 130-179). GRENADINE ISLAND: TABL 104420 (2: 84-98).
TRINIDAD: TABL 103466 (4: 31-144), 104372 (1: 259), 105364 (4: 230-
269). VENEZUELA: Carupano, TABL 103207 (3: 199-280). BRAZIL:
Recife, SU 51844 (2: 192-196).

Oligoplites saurus inornatus (27 specimens, 53-257 mm. FL from 12 col-
lections). GULF OF CALIFORNIA: TABL 103481 (6: 112-132), 103483
(7: 53-93), SIO 60-298 (2: 110-125), USNM 101037 (1: 246). BAJA CAL-
IFORNIA: Magdalena Bay, SIO-304 (3: 104-109). MEXICO: Banderas
Bay, TABL 103479 (1: 220), 103482 (1: 104). COSTA RICA: SIO H52-93
(2: 253-257). PANAMA: USNM 30959 (248, holotype of Oligoplites in-
ornatus), TABL 105800 (1: 127). ECUADOR: Esmeraldas, TABL 103480
(1: 200). GALAPAGOS ISLANDS: AMNH 8308 (2: 71-75).

Oligoplites altus (Giinther).
(Figures 10c, 10d, 11; tables 4-6.)

?Lichia albacora GUICHENOT, 1848, p. 231 (type locality Valparaiso, Chile).

VoL. XXXIX] SMITH-VANIZ & STAIGER: CARANGIDAE 223

Chorinemus altus GUNTHER, 1868, p. 433, unnumbered fig. (of head) (type locality west
coast of Panama).

Oligoplites mundus JorDAN and Starks in Jordan and Evermann, 1898, p. 2844 (type locality
San Juan Lagoon, Mexico, mouth of Ahome River).

In the original description Lichia albacora was compared only with Scomber
calcar Bloch, a probable synonym of Oligoplites saliens. The description, al-
though poor, applies best to a species of Oligoplites. The statement “each jaw
has two rows of velvet teeth” is a good description of the teeth in a young
individual of Oligoplites, but the single type was reported to be 45 inches total
length. However the following statements, also given in the description, suggest
that the type was small, probably only 4.5 inches TL: “The fishermen of Val-
paraiso catch it on rare occasions. It seems that it is usually much larger than
the one we have observed.” Lichia albacora may have been based on a young
specimen of O. altus. This species is known to occur farther south than the
other eastern Pacific members of Oligoplites, and the first dorsal spine count
of three may have been the result of overlooking the small, frequently depressed,
first spine. The type of Lichia albacora could not be located in the Museum
National d’Histoire Naturelle, Paris, where, if extant, it should be deposited.

Schultz (1945) regarded Oligoplites mundus as a distinct species, distin-
guished from O. altus on the basis of a slightly longer upper jaw. Both species
were otherwise very similar, sharing a number of characters not found in other
Pacific species of Oligoplites. Schultz had only two examples which he con-
sidered conspecific with O. altus, the same two specimens on which the account
of the species in Meek and Hildebrand (1925, p. 388) is based. We have ex-
amined 80 specimens, including the type of O. mundus and Schultz’s two examples
of O. altus. However, more specimens are needed before it will be possible
to resolve the systematic status of the two forms. Because we are unable to
confidently assign a large number of specimens to one or the other form, we
tentatively regard them as conspecific.

The species, as here defined, exhibits a more extreme range of variation in
several characters than is found in other species of Oligoplites. Particularly
striking is the range of variation in upper jaw length (see table 6). This vari-
ation is not due to sexual dimorphism; nor is it due to population differences,
because specimens of approximately the same size, from the same station, exhibit
the extremes in variation for several characters. Individuals with long jaws
generally have deeper bodies, higher fin lobes, smaller eyes, fewer gillrakers,
dorsal spines, and segmented dorsal- and anal-fin rays than do their short jawed
counterparts. When any of these characters are plotted against upper jaw length
(expressed as percent head length) an obvious negative or positive correlation
with jaw length is apparent. It is however, impossible to separate specimens
into two groups on the basis of these characters. Both forms are distinguished
from their congeners in having the dorsum of the head covered with numerous

224 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

TaBLe 6. Comparison of fork length and upper jaw length (as percent head length)
in Oligoplites altus.

7 Length
Fork Length hel yee ae

Class (mm.) 58-59.9 60-61.9 62-63.9 64-65.9 66-67.9 68-70 N
100-150 at 1 3 6 =e = 10
151-199 3 5 6 4 1 3 22
200-249 2 1 3 2 3 2 13
250-299 ae 1 5 3 2 3 14
300-349 = 2 = 4 1 2 9
350-380 = _ 1 3 1 1 6

sensory pores, which are connected by a network of tubes visible beneath the
skin. They may also be distinguished from those species with which they occur
sympatrically by their premaxillary dentition, lower number of gillrakers, and
longer upper jaw (see table 5). An explanation for the wide range of variation
in upper jaw length and strong correlation with several other characters in O.
altus is a subject that requires investigation by future workers. One possibility
that should be considered is that this variation is due to introgressive hybrid-
ization (see Anderson, 1949). In many of its characters the short jawed form
of O. altus is intermediate between the long jawed form and O. saurus inornatus.

Oligoplites altus reaches at least 379 mm. FL and attains the greatest body
depth of any Oligoplites species. On the basis of our material, O. altus occurs
in the northern Gulf of California, north at least to Punta Estrella, along the
outer coast of Baja California to Magdalena Bay, and south to the Gulf of
Guayaquil. It has also been reported by several authors from Peru, apparently
on the authority of Tortonese (1939, p. 338), who listed a single specimen from
Callao.

MATERIAL (81 specimens, 27-379 mm. FL, from 36 collections). BAJA CAL-
IFORNIA: Magdalena Bay, TABL 105541 (2: 354-372), 105544 (1: 348),
SIO 62-718 (2: 259-299), 62-725 (7 of 11: 281-300). GULF OF CAL-
IFORNIA: TABL uncataloged, Almejas Bay (2: 236-317), TABL 105534 (1:
315), 105542 (1: 355), 105543 (2: 332-354), SIO 61-180 (8: 171-199), 61—
182 (1: 257), UCLA W50-46 (2: 195-209), W52-132 (1: 168), W58-48 (2:
147-149). MEXICO: San Juan Lagoon, USNM 48876 (231, holotype of O.
mundus); Tepic, USNM 130860 (2: 122-137.5); Banderas Bay, TABL un-
cataloged (2: 112-114). EL SALVADOR: USNM 87338 (U3572);eAbe
104616 (1: 231). COSTA RICA: Gulf of Nicoya, TABL uncataloged (2:
292-314), UCLA W54-12 (1: 130), W54-171 (1: 138.5). GULF OF PAN-
AMA: SIO 58-402 (3: 250-310), UCLA W58-303 (2: 189-212), W58-304
(8: 162-202), W58-305 (3: 137-155), USNM 80063 (1: 198), 82043 (1:215),

VoL. XXXIX] SMITH-VANIZ & STAIGER: CARANGIDAE 225

128698 (3: 27-59), 128699 (1: 83); San Miguel, TABL uncataloged (3: 178-
298). COLOMBIA: Buenaventura Market, TABL uncataloged (1: 241).
BeRADOR: Esmeraldas, TABL 105528 (1: 207), 105529 (2: 195-201),
TABL uncataloged (1: 220); Gulf of Guayaquil, TABL 107260 (2: 379),
TABL uncataloged (6: 222-337).

Oligoplites palometa (Cuvier).

(Figures 9b, 11, 19; tables 4-5.)

Chorinemus palometa Cuvier in Cuvier and Valenciennes, 1831, p. 392 (288) (type locality
Lake Maracaibo, Venezuela).

?Chorinemus guaribira Cuvier in Cuvier and Valenciennes, 1831, p. 393 (289) (type locality
Brazil) .

This western Atlantic species is very closely related to the eastern Pacific
O. altus. Both species are separable from other forms of Oligoplites on the
basis of their premaxillary dentition, which consists of a band of villiform teeth
along entire length of premaxilla, wide anteriorly, becoming narrow posteriorly.
Superficially they are very similar, although O. altus is generally a more deep-
bodied fish. The only character that will separate these two species is the
relative development of cephalic pores. Reduction of cephalic pores in O. pal-
ometa suggest that it is the more specialized of the two species.

Oligoplites palometa is a large species, attaining at least 392 mm. FL. It
occurs from Lake Yzabal, Guatemala, south to Sao Paulo, Brazil.

MATERIAL (42 specimens, 84-392 mm. FL, from 22 collections). GUATE-
MALA: Lake Yzabal, USNM 13480 (1: 139). NICARAGUA: AMNH
19027 (18: 84.5-160), USNM 44205 (3: 329-268), 45373 (1: 348), 45374
(1: 317), 45375 (1: 304). GULF OF VENEZUELA: USNM 121803 (1: 344),
121804 (1: 245), 121805 (2: 241-303), 121806 (1: 133.5). BRAZIL: USNM
100817 (1: 342), 100847 (1: 392), 100851 (1: 297), SU 66978 (1: 141); Rio
Para, SU 53059 (1: 248), 53060 (1: 219), 53061 (1: 222); Salvador (Bahia), SU
j20o4( 125535): Recife, SU 53058 (1: 305), 67028 (1: 334); Sao Paulo, SU
67007 (1: 171), 67014 (1: 228).

Parona Berg, 1895

Paropsis JENYNS, 1842, p. 65 (type-species: Paropsis signatus Jenyns, 1842, by monotypy).
Parona BeErc, 1895, p. 39 (substitute name for Paropsis Jenyns, 1842, preoccupied, and,
therefore, taking the same type species: Paropsis signatus Jenyns, 1842).

This monotypic genus is restricted to subtropical and temperate waters of
the south Atlantic from Brazil to southern Argentina. Almost nothing has been
published on the biology of Parona, although the genus is reported (Evermann
and Kendall, 1906, p. 99) to be very common along the whole coast of Argentina,
and caught at Montevideo, Uruguay, during certain times of the year in enormous

226 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

quantities. The unusual lateral line with 5—9 dorsal branches, the deep, com-
pressed body, and the lack of pelvic fins suggest that their ecology differs from
that of most carangids. Norman (1937, p. 59) reported six specimens, 465-—
600 mm. SL, caught by otter trawl in 51-56 m.; these specimens are the largest
examples known to us. Nothing on the osteology of Parona has been published
previously.

DEscRIPTION. Body strongly compressed. Adipose eyelids poorly developed.
Body, except head, covered with oval-shaped scales. Lateral-line scales not
developed into scutes. Lateral line with 5—9 dorsal branches; horizontal portion
of lateral line straight except for moderate curve over pectoral fin. Anterior
rim of shoulder girdle smooth, without fleshy papillae or deep groove near
isthmus. Premaxillae not protractile, connected anteriorly to snout at midline
by a narrow fleshy bridge. Swimbladder simple, not bifid posteriorly. Posterior
dorsal- and anal-fin rays not consisting of semidetached finlets; ultimate and
penultimate rays no more closely spaced than adjoining rays; none of dorsal-
or anal-fin rays produced into long filaments. Pelvic fins absent. Pectoral fins
short, not falcate, 60.0 to 67.0 percent head length in adults. Upper jaw ex-
tending posteriorly from slightly behind (young) to well behind (adult) orbital
rim, 52.6 to 57.0 percent head length in adults. Depth (origin Dz to Ag) 37.4
to 44.0 percent fork length in adults.

Predorsal bones 4; dorsal-fin rays VI-VII (rarely VII) + I, 32 to 37;
total dorsal-fin rays 39-44, Anal-fin rays II + I, 34 to 38. Pectoral-fin rays
I, 19-21. Pelvic fins absent. Vertebrae 10 + 17 = 27, epipleural ribs 10 or 11.
Branchiostegal rays 3 + 6 (epihyal + ceratohyal). Upper gillrakers 4 to 8;
lower gillrakers 15 to 17; total gillrakers 20 to 24.

Fronto-supraoccipital crest high; temporal crest extending nearly straight
forward to front of the frontal; preorbital region short. Epiotics broadly united
along midline of cranium internally. Zygapophyses of exoccipital united beneath
foramen magnum. Parasphenoid not expanded into a broad, flattened plate
posteriorly. Myodome with broad, posterior opening. Basioccipital without
a pair of lateral processes on ventral surface. Ascending process of premaxilla
short and triangular; maxilla and premaxilla relatively broad, not closely
attached to premaxilla. Premaxillary spur very reduced, consisting of only a
slight bulge. Large supramaxilla present. Interosseous space between dorsal
arm of dentary and upper margin of articular well developed. Well developed
teeth on dentary, premaxilla, palatines, and vomer; dentary and premaxillary
teeth conical, arranged in a narrow band anteriorly becoming a single row poste-
riorly; symphyseal premaxillary and dentary teeth of juveniles slender and
caniniform. Mesopterygoids partially covered by a patch of free-floating gran-
ular teeth. Pharyngeals not noticeably enlarged, covered with sharp, pointed

teeth.

VoL. XXXIX] SMITH-VANIZ & STAIGER: CARANGIDAE 227

Suborbital shelf present; lower and posterior suborbitals not expanded
posteriorly to cover a large portion of cheek. Basibranchials 1-3, each with a
large, median tooth plate. Lower limb of posttemporal not noticeably short
or thickened. Postcoracoid process well developed. Inter-pelvic keel absent;
postpelvic process of each side coalesced entire length, not forming an apical
fork. Inferior vertebral foramina absent. First anal pterygiophore and haemal
spine of first caudal vertebra firmly attached to form a strong strut. Anal
pterygiophore not expanded anterolaterally to form roof over first two anal
spines; first two anal spines not articulated proximally with each other; second
anal spine symmetrical, without deep groove on one side of anterior face. Caudal
skeleton with 2 epurals and 2 pairs of uroneurals.

Parona signata (Jenyns).
(Figures 12, 14a, 15a, 16a, 20a, 21a, 22a, 23a, 24d, 25a.)

Paropsis signatus JENYNS, 1842, p. 66, pl. 13 (type locality Bahia Blanca, coast of northern
Patagonia).

DEscriPTION. Since Parona is monotypic, refer to generic description.

PIGMENTATION. Most specimens of Parona signata have a conspicuous
elongate, horizontal, black blotch on sides beneath pectoral fins, which may also
extend onto base of ventral third of pectoral fin. In a few specimens this blotch
is short and almost completely hidden by the pectoral fin; it is completely
absent in 7 small specimens (TABL 102908) from Argentina. A small round
spot at the dorsal origin of the pectoral fin is present on all specimens. Also
present on all specimens is a dusky blotch on dorsal! third of opercle, heavily
pigmented areas, slightly larger than pupil, on either side of snout that are
separated at dorsum by paler region, and concentration of pigment at tip of
lower jaws. Jenyns (1842, p. 67) reported the life colors as “uniform bright

228 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

silvery, the ridge of back bluish: a patch of black on the gill-cover, and another
under the pectoral fin.”

DISTRIBUTION. Southern Brazil to southern Argentina. Apparently re-
stricted to western Atlantic Ocean. Reported from Chile “‘Blacino del Rio Santa
Cruz” (Fowler, 1945, p. 70), but this record needs verification. The same locality
is generally listed as being in Argentina.

MATERIAL (20 specimens, 75-368 mm. FL, from 11 collections). BRAZIL:
TABL 107713 (2: 358-368); Florianopolis, SU 53052 (1: 264), 53053 (1:
360); Port of Rio Grande do Sul, SU 51868 (2: 171-192). URUGUAY: USNM
87730 (2: 312-335), 86715 (1: 148 mm. SL, cleared and stained) ; Montevideo,
USNM 77375 (1: 135), 86689 (1: 298). ARGENTINA: USNM 53441 (1:
289); Buenos Aires, TABL 101156 (1: 270); Mar del Plata, TABL 102908
(7: 75-84, 3 cleared and stained).

Hypacanthus Rafinesque, 1810

Hypacanthus RA¥FINESQUE, 1810, p. 43 (type-species: Centronotus vadigo Lacépede, 1802 =
Scomber amia Linnaeus, 1758, by present designation).

Lichia Cuvier, 1817, p. 321 (type-species: Scomber amia Linnaeus, 1758, by monotypy).

Porthmeus VALENCIENNES im Cuvier and Valenciennes, 1833, p. 255 (190) (type-species:
Porthmeus argenteus Valenciennes, 1833, by original designation).

This monotypic genus is restricted to subtropical and temperate waters of
the Mediterranean, eastern Atlantic, and southern Indian Ocean. The habitat
of Hypacanthus is littoral (Poll, 1954, p. 160) “. . . judging by the origins of
specimens—collections and total absence of this species from catches of trawls.”
Along the west African Gold Coast small specimens of Hypacanthus are caught
in seines, but uncommonly, and they are taken with lines from January to April
(Norman and Irvine, 1947, p. 144). Smith (1949, p. 222) states that it “grows
to at least 6 feet and is one of the finest game fishes, fighting fiercely to the
end; prefers live or moving bait, enters estuaries, and may often be seen in
pursuit of mullet on the surface.” Superficially Hypacanthus most closely
resembles Scomberoides but differs in having scaled cheeks and a lateral line
that is very irregular and sinuous, dropping below midline of body between pelvic
fins and origin of anal fin. Nothing on the osteology of Hypacanthus has been
published previously.

NoMENCLATURE. In his paper on the nomenclature of the European fishes
of the subfamily Trachinotinae, Wheeler (1963) clearly demonstrated that
Hypacanthus is a senior synonym of Lichia but chose to regard the older name
as a nomen oblitum and advocated suppression of the name. We do not believe
that this action is justified and continue to recognize Hypacanthus. In so doing
we have taken special note of the fact that the name Hypacanthus has always
been correctly applied, while fishes belonging to at least four different carangid
genera have been described as new species of Lichia.

VoL. XXXIX] SMITH-VANIZ & STAIGER: CARANGIDAE 229

In the generic diagnosis of Hypacanthus, Rafinesque referred to only two
species. One of these species, Centronotus vadigo Lacépéde, is clearly a synonym
of Scomber amia Linnaeus and is here designated the type-species of Hypa-
canthus. The other reference was to “Scomber aculeatus di Linnaeus.” As
noted by Wheeler (1963, p. 535), the species does not seem to exist in Linnaeus’s
works and it might be argued that Rafinesque wrote “di Linnaeus” in error for
S. aculeatus Bloch (1793). Bloch’s illustration strongly suggests a species of
Oligoplites or Scomberoides, and several workers have followed Cuvier and
Valenciennes (1831, p. 384) in questionably regarding S. aculeatus Bloch as a
species of Chorinemus (=Scomberoides). We concur with Wheeler (1963, p.
537) that it is a junior synonym of Hypacanthus amia (Linnaeus). Bloch’s
illustration shows a fish with 7 first dorsal spines (which does not agree with
Oligoplites), and the species is reported from the Mediterranean where neither
Oligo plites or Scomberoides occurs.

The illustration of Porthmeus argenteus Valenciennes (see fig. 13) is un-
mistakably based on a young individual of Hypacanthus amia.

DEscrIPTION. Body moderately to strongly compressed. Adipose eyelids
developed. Body, including cheeks, covered with narrow, oval-shaped embedded
scales, becoming needlelike on breast. Lateral-line scales not developed into
scutes. Lateral line without side branches, very irregular and sinuous, dropping
below midline of body between pelvic fins and origin of anal fin. Anterior rim
of shoulder girdle smooth, without fleshy papillae or deep groove near isthmus.
Premaxillae protractile, separated anteriorly from snout at midline by a complete
groove. Swimbladder moderately bifid (in adults), bifurcated portion of swim-
bladder extending posteriorly to beneath caudal vertebra 6 or 7. Posterior
dorsal- and anal-fin rays not consisting of semidetached finlets; ultimate and
penultimate rays no more closely spaced than adjoining rays; none of dorsal-
or anal-fin rays produced into long filments. Pelvic fins relatively short, not
depressible into shallow abdominal groove. Pectoral fins short, not falcate, 51.2
to 63.4 percent head length. Maxilla extends posteriorly to slightly behind
orbital rim, 50.5 to 54.2 percent head length. Body depth (origin D» to Ag)
31.0 to 39.4 percent fork length.

Predorsal bones 3; dorsal fin rays VII + I, 19-21; total dorsal rays 27-29.
Anal-fin rays II-I, 17-20. Pectoral-fin rays I, 18-20. Pelvic-fin rays I, 5.
Vertebrae 10 + 14 = 24; epipleural ribs 10 or 11. Branchiostegal rays 2% +
5% (epihyal + ceratohyal). Upper gillrakers 1 or 2; lower gillrakers 8 or 9;
total gillrakers 9 to 11.

Fronto-supraoccipital crest low; temporal crest poorly developed, not ex-
tending forward to posterior rim of orbit and not reaching edge of cranium;
preorbital region short. Epiotics not united along midline of cranium internally.
Zygapophyses of exoccipital united beneath foramen magnum. Parasphenoid

230 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

not expanded into a broad, flattened plate posteriorly. Myodome with broad,
posterior opening. Basioccipital without a pair of lateral processes on ventral
surface. Ascending process of premaxilla short and triangular; maxilla moder-
ately slender, not closely applied to premaxilla. Premaxillary spur absent. Small
supramaxilla present. Interosseous space between dorsal arm of dentary and
upper margin of articular moderate. Well developed teeth on dentary, premaxilla,
palatines, and vomer; dentary and premaxillary teeth slender and pointed,
arranged in a broad band anteriorly becoming narrower posteriorly; juveniles
without symphyseal premaxillary and dentary canines. Mesopterygoids edentate.
Pharyngeals not noticeably enlarged, covered with sharp, pointed teeth.

Suborbital shelf present; lower and posterior suborbitals not expanded poste-
riorly so as to cover a large portion of cheek. First basibranchial without tooth
plates, basibranchials 2—3 each with a pair of small widely separated tooth
plates. Lower limb of posttemporal not noticeably short or thickened. Post-
coracoid process moderate. Interpelvic keel well developed; postpelvic process
on each side not coalesced entire length, forming an apical fork. Inferior
vertebral foramina present. First anal pterygiophore and haemal spine of
first caudal vertebra firmly attached to form a strong strut. Anal pterygiophore
not expanded anterolaterally to form roof over first two anal spines; first two
anal spines not articulated proximally with each other; second anal spine slightly
asymmetrical, with shallow depression on one side of anterior face. Caudal
skeleton with 3 epurals and two pairs of uroneurals.

Hypacanthus amia (Linnaeus).

(Figures 13, 14b, 21b, 22b, 24a.)

Scomber amia LINNAEUS, 1758, p. 299 (based solely on references to Artedi, 1738).

Scomber aculeatus Biocy, 1793, p. 51, pl. 336, fig. 1 (type locality: no specific locality
given; reported from coast of Mediterranean and several localities in Caribbean).

Centronotus vadigo LACEPEDE, 1802, p. 318 (type locality Mediterranean).

Porthmeus argenteus VALENCIENNES in Cuvier and Valenciennes, 1833, p. 256 (191), pl. 264
(type locality Cape of Good Hope).

NOMENCLATURE. Wheeler (1963, pp. 536-7) gives a detailed discussion
of the specific synonymy of Hypacanthus. Some additional comments are given
in the preceding generic account.

DeEscriPTION. Since Hypacanthus is monotypic, refer to generic description.

PIGMENTATION. The young, to at least 120 mm. FL, have prominent vertical
bands (fig. 13). Judging from a color illustration of a 2'2 inch specimen (Smith,
1949, pl. 25, fig. 539), the bands are brownish-black and the background color-
ation is yellow-orange. Adults are brown dorsally, silvery white below the lateral
line, and have pale brown fins.

DISTRIBUTION. Mediterranean, eastern Atlantic, and Indian Ocean north

VoL. XXXIX] SMITH-VANIZ & STAIGER: CARANGIDAE 231

Ficure 13. Hypacanthus amia: above, TABL 104703, young adult, 200 mm. FL; below,
drawing of holotype of Porthmeus argenteus = juvenile H. amia (after Cuvier and Valen-
ciennes, 1833, pl. 264).

to Delago Bay, Mozambique. The littoral habitat, general occurrence in sub-
tropical and temperate waters and absence of Hypacanthus from most of the
Indian Ocean indicates that the Indian Ocean distribution of Hypacanthus is
the result of migration around the Cape of Good Hope.

MATERIAL (24 specimens, 62—202 mm. FL, from 5 collections). MEDI-
TERRANEAN: Lebanon, St. George Bay, TABL 104703 (5: 62-200, 1 cleared
and stained), 104714 (15 of 30: 79-110, 2 cleared and stained). LIBERIA:
Mesurado River, USNM 193783 (1: 202), 193813 (1: 82). SOUTH AFRICA:
Durban Harbor, TABL 107250 (2: 190).

RELATIONSHIPS AND ZOOGEOGRAPHY
GENERA Scomberoides AND Oligoplites.

The strong external resemblance and large number of characters shared
by Scomberoides and Oligoplites clearly reflect their intimate relationship, and
they are usually placed together in their own subfamily—the Scomberoidinae
(=Chorineminae). Some of the more notable shared characters are the following:
scales lanceolate or needlelike (fig. 3); the posterior segmented dorsal- and
anal-fin rays consisting of semi-detached finlets; two distinct types of dentary

iS)
Ww
iS)

CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

PAPA =e
POT TAT AANA WV A Te /

AAUANDN ADNAN CTA amg

om,

WOMAN UAW Vnitr 4

a

— FE AA

=

hy,

WR pe mipctte ere yy mnt aT HET ula Ua eva
fi RE if] I

Ficure 14. Jaw bones: a, Parona signata, USNM 86715, 148 mm. SL; b, Hypacanthus
amia, TABL 104714, 100 mm. SL, c, Scomberoides commersonianus, GVF 2195, 86 mm. SL;
d, S. tol, TABL uncataloged S-V 69-128, 107 mm. SL; e, Oligoplites s. saurus, TABL 107781,
91.5 mm. SL; f, O. saliens, SU 66973, 122 mm. SL. DN, dentary; AR, articular; AN, angular;
PMX, premaxilla; MX, maxilla; SMX, supramaxilla. Scale 5.0 mm.

teeth during different stages of their ontogeny (see discussion below) ; slender,
nearly straight maxillae that are closely applied to the premaxillae; a broad
premaxillary frenum; interosseous space between dorsal arm of dentary and
upper margin of articular minute or absent (fig. 14); the first anal pterygi-
ophore expanded anterolaterally, forming a roof over anal spines (fig. 15b);
the first two anal spines articulated proximally with each other, and the second
anal spine asymmetrical with a deep groove on one side of anterior face, so that
when anal spines are depressed the first spine folds into the groove in scissor-
like fashion; caudal skeleton with 2 epurals (fig. 16b); swimbladder strongly

VoL. XXXIX] SMITH-VANIZ & STAIGER: CARANGIDAE 233

Ficure 15. First anal pterygiophore: a, Parona signata, TABL 102908, 80 mm. SL; b,
Scomberoides tala, TABL S-V 69-156, 109 mm. SL.

bifid, even in smallest specimens (fig. 17). All of these conditions probably
represent specializations.

Each dentary has two rows of teeth in Scomberoides and Oligoplites. In
most species (exceptions are S. tol and O. saliens) the outer dentary teeth of
juveniles and young adults have spatulate tips, and are closely spaced and
strongly hooked outward (fig. 18). This arrangement produces a trough be-
tween the inner and outer rows of teeth into which the premaxillary teeth fit
when the mouth is closed. In adults these outer ‘juvenile teeth’ are shed and
replaced with more robust, wider spaced teeth that are not strongly hooked and
have pointed tips (figs. 18-19). The inner dentary teeth, consisting of a single
type, apparently are permanent. We know of no other carangid genus in which
there are two distinct types of dentary teeth associated with ontogeny. Since
this unusual adaptive feature is shared by both Scomberoides and Oligoplites,
we believe the character was possessed by the progenitor of both genera and
its loss in S. tol and O. saliens must be interpreted as a subsequent specialization.
Major (1973) has correlated this change in dentition in Scomberoides lysan, Oligo-
plites refulgens, and O. saurus inornatus with a shift in feeding habits. Juveniles
and young feed primarily on larval crustaceans, and on scales and epidermis of
schooling fishes. He suggested that the outer spatulate dentary teeth are inserted
under the scales, which are then ripped off. With growth and loss of ‘juvenile’

234 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

E1 Eo UN

Ficure 16. Caudal skeleton: a, Parona signata, USNM 86715, 148 mm. SL; b, Scom-
beroides commersonianus, GVF 2195, 86 mm. SL. E, epural; UN, uroneural.

teeth there is a gradual shift in diet until these species become fully piscivorous,
ingesting whole fish. In juveniles of all species of Scomberoides except S. lysan
the symphyseal dentary and premaxillary teeth are robust and caninelike (figs.
14c-d). Such teeth are greatly reduced or absent in the adults; they are never
present in Oligoplites.

Carr and Adams (1972) found that ectoparasites, primarily caligoid co-

50 mm

Ficure 17. Lateral and ventral view of swimbladder of Scomberoides tala, TABL 107712,
264 mm. FL.

Vor. XXXIX] SMITH-VANIZ & STAIGER: CARANGIDAE 235

b 0.5 mm :

Ficure 18. Lateral and frontal view of outer row, dentary tooth of Oligoplites s. saurus:
a, juvenile 90 mm. FL; b, adult 185 mm. FL.

236 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

160
SCOMBEROIDES
fe}
140 COMMERSONIANUS e
° tier Che TALA
fe) Oo
° 2 ° <) LYSAN
120
° é r TOL
s ° ° e CS
100 A
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Ce mle x * Ce ou “eo
[a °
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= 3
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+ x *
0 e ae Hers ov e ox = Ba On 5
2 * 2a se x Ge *& & e
== x ke RE Oe Ok 0 OE a # xO * °
20 Pig Why Fw * we : x
Le le aes
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- 1 L l l l L l l Nl ! ]
LL
Lu 200 x
180 Oh °
> oer oe he, C OLIGOPLITES
ce &° 2 e
q 160 SAURUS
ee
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<. A ; REFULGENS
uj 14° oe: QD ¢ A
e LIENS
a eee
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e e ° °
LL os ee ° o 9°90 <
e ° OX
oO ° fo) x °
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oc 100 ion ay
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60 80 100 120 140 160 180 200 220 240 260 280

FORK LENGTH (mm)

Ficure 19. Scatter diagram of outer row teeth on dentary (one side) plotted against
fork length for species of Scomberoides and Oligoplites (O. altus and eastern Pacific specimens
of O. saurus were not plotted).

VoL. XXXIX] SMITH-VANIZ & STAIGER: CARANGIDAE 237

mo)66hCUlMh ee SC

Ficure 20. Upper dentition: a, Parona signata, TABL 101156, 271 mm. FL; b, Scom-
beroides tol, TABL 107707, 284 mm. FL; c, Oligoplites s. saurus, TABL 105364, 230 mm. FL.
PMX, premaxilla; V, vomer; PA, palatine; MSP, mesopterygoid.

pepods, accounted for approximately 50 percent of the stomach contents of ju-
veniles of Oligoplites s. saurus 26 to 40 mm. SL. In contrast, ectoparasites con-
stituted only 2 to 4 percent of the stomach contents in fish less than 26 mm. or
greater than 60 mm. SL. Although no feeding observations were made, they
postulated that this carangid passes through a stage in its development in which
intentional removal of ectoparasites from other fishes is an important activity.
Most of the fish skin, flesh, and scales found in the stomachs of juveniles was
presumed to have come from cleaning activities. Major (1973) believed that
scales and/or epidermal tissue was the primary food source sought by juveniles
of Scomberoides lysan in Hawaii. His analysis of the stomach contents of Oligo-
plites juveniles is inconclusive, since the smallest specimen examined was 48
mm. SL. If the conclusions regarding the primary food source sought by ju-
veniles of Oligoplites and Scomberoides are valid, they may provide valuable
insight into the evolution of cleaning symbiosis in fishes. Ectoparasite removal
seems to be a more specialized type of feeding than is aggressive scale ingestion,
although one that might easily have evolved from the latter activity.

The following characters of Oligoplites are interpreted as specializations of
the condition found in Scomberoides and suggest that the amphi-American
endemic distribution of Oligoplites is a relatively recent evolutionary develop-
ment (this does not necessarily imply that the genus evolved elsewhere): 1)
reduced number of dorsal spines with a concomitant increase in number of pre-
dorsal bones (see discussion under Characters and Methods Section) ; 2) loss of
mesopterygoid teeth (fig. 20c); 3) loss of supramaxilla; 4) loss of suborbital

238 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

FicureE 21. Lateral and dorsal view of suborbital bones: a, Parona signata TABL 102908,
83 mm. SL; b, Hypacanthus amia, TABL 104714, 100 mm. SL; c, Scomberoides tol, TABL
S-V 69-128, 122 mm. SL; d, Oligoplites s. saurus, TABL 107781, 91.5 mm. SL. Third suborbital
bone shaded.

shelf (Smith and Bailey, 1962) and expansion of suborbitals 2—4 posteriorly
across cheek (fig. 21d), a character best developed in large specimens; 5) re-
duction in number of branchiostegal rays (fig. 22d); and 6) presence of needle-
like scales in all species. Steps in the evolution from broadly lanceolate to
needlelike scales can be seen in figure 3.

We believe that Oligoplites evolved from a Scomberoides-like ancestor that
reached the New World via the Pacific. Although the East Pacific Barrier re-
stricts the distributions of many groups of tropical shorefishes (Briggs, 1961),
it is not such a formidable obstacle to carangids as evidenced by the relatively

>

Ficure 22. Left hyoid arch; branchiostegal rays shown in black: a, Parona signata,
USNM 86715, 148 mm. SL; b, Hypacanthus amia, TABL 104714, 100 mm. SL; c, Scom-
beroides tol, TABL S-V 69-128, 122 mm. SL; d, Oligoplites s. saurus, TABL 107781, 91.5
mm. SL.

VoL. XXXIX]

SMITH-VANIZ & STAIGER: CARANGIDAE

240 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

]

Ficure 23. Fifth and sixth precaudal vertebrae (pleural ribs are shown in _ black):
a, Parona signata, TABL 102908, 80 mm. SL; b, Scomberoides commersonianus, GVF
2195, 86 mm. SL.

large number of circumtropical genera and species. One widely distributed
Indo-West Pacific carangid, Caranx (Gnathanodon) speciosus (Forskal), that
has become established in the eastern Pacific appears to be a recent migrant
across this several thousand mile stretch of open ocean. The wide distribution
of Scomberoides lysan in the central Pacific demonstrates that this group of
carangids also has the ability to cross large stretches of oceanic water. Given
this potential, it is not difficult to envision successful bridging of the East Pacific
Barrier by a Scomberoides-like carangid. As previously noted, the nature of the
characters that differentiate Oligoplites from Scomberoides indicates a relatively
recent evolutionary separation. This suggests that by the time the progenitor
of Oligoplites reached the new world it possessed most of the characters present
in Scomberoides.

Although inconclusive, the known fossil carangid record supports the above
hypothesis. Middle Eocene deposits from Monte Bolca near Verona, Italy con-
tain a fairly rich carangid fauna (Blot, 1969) but no Oligoplites-like fossil has

VoL. XXXIX] SMITH-VANIZ & STAIGER: CARANGIDAE 241

Ficure 24. Basibranchial tooth plates (shown in black) and basihyal teeth (stippled):
a, Hypacanthus amia, TABL 104714, 100 mm. SL; b, Scomberoides tala, TABL S-V 69-
156, 109 mm. SL; c, Oligoplites saliens, SU 66973, 122 mm. SL; d, Parona signata, TABL
102908, 80 mm. SL.

been found. The only record of a fossil Oligoplites outside the new world ap-
pears to have been based on a misidentification (see discussion of Scomberoides
spinosus).

GENUS Parona.

Parona parallels Scomberoides (and Oligoplites) in several of its specialized
characters, has others not found elsewhere in the Carangidae, and shares with
Hypacanthus and Trachinotus a number of generalized (primitive) characters
that have become specialized in the Scomberoidinae. Characters that suggest
affinity with Scomberoides are the broadly united epiotics, high number of caudal
vertebrae, fusion of the basibranchial tooth plates and presence of a premaxillary
frenum. In having the epiotics broadly united internally, Parona possesses a
character utilized by Starks (1911) and Suzuki (1962) in defining the subfamily
Scomberoidinae. The shape of the precaudal vertebrae and the position of attach-
ment of the pleural ribs is very different from the condition in Scomberoides (fig.
23). This is probably due to differences in body shape however, because Parona
and Tvrachinotus ovatus (Linnaeus), which have considerably deeper bodies
than do the species of Scomberoides, have precaudal vertebrae that are nearly
identical.

242 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Ficure 25. Posttemporal, clavicle, hypercoracoid, hypocoracoid, and radials (arrow points
to postcoracoid process): a, Parona signata, TABL 102908, 80 mm. SL; b, Scomberoides
commersonianus, GVF 2195, 86 mm. SL.

Nelson (1969, p. 500) noted that there are three pairs of basibranchial tooth
plates in Chorinemus, and the median plates of Parona (fig. 24d) must have
arisen through fusion of such paired elements (his ‘““Chorinemus occidentalis” =
Oligoplites s. saurus). He also stated (p. 502) that ‘‘jaw protrusion and pharyn-
geal-bone mobility seem to be improvements, and shearing and grinding jaw
dentitions are specializations, inversely correlated with basibranchial dentition.
The large paired plates (and their median derivatives, e.g., in Parona) that in
some cases occur over the basibranchials of teleosts could represent a reversal
in the general trend toward loss of basibranchial dentition. Functionally, this
reversal would mean that the basibranchial area in some cases is secondarily
involved in seizing and holding large prey.” Since the species of Parona, Scom-
beroides, and Oligoplites have the best developed basibranchial dentition in the
family, comprise the only carangid genera that have a premaxillary frenum
(which restricts upper jaw protrusability), and feed on large prey as adults, we
believe that the well developed basibranchial plates of these genera represent a
secondary specialization associated with mode of feeding.

The mutual occurrence in these genera of several derived character states
that are not found in any other carangids is here interpreted as evidence that
Parona and Scomberoides evolved from a common ancestor. Although some of
the shared specializations, especially those associated with trophic ecology, might
be due to parallelism, this does not seem likely in the case of united epiotics.

Parona differs from all other carangids in having typically 9 branchiostegal

Vout. XXXIX] SMITH-VANIZ & STAIGER: CARANGIDAE 243

rays and in lacking pelvic fins (a small pelvic girdle is retained). The pelvic
fins are very reduced, however, in Trachinoius stilbe (jordan and McGregor),
and in fossil trachinotid genus Paratrachinotus (Blot, 1969, p. 285). The lat-
eral line of Parona is unusual in having 5—9 dorsal branches. The mesopterygoid
teeth are firmly ankylosed to the mesopterygoid in Scomberoides. In Parona the
‘mesopterygoid’ teeth are free-floating and thus may not be homologous. The first
anal pterygiophore and articulation of the first two anal spines is very general-
ized compared to the complex arrangement in Scombcroides. Other generalized
characters of Parona are the relatively broad, curved maxillae; well developed
interosseous space between dorsal arm of dentary and upper margin of articular;
large supramaxillae; caudal skeleton with 3 epurals; and broad, oval-shaped
scales. The shoulder girdle of Parona differs from that of all other carangids,
except those of the subfamily Naucratinae, in having a well developed postcora-
coid process (fig. 25a).

The large number of differences between Parona and Scomberoides suggest
that they have had a long history of evolutionary separation. The restriction
of Parona to subtropical and temperate waters of the South Atlantic Ocean prob-
ably reflects an extended period of isolation in that region.

Genus Hypacanthus.

Hypacanthus differs from the previous genera in having inferior vertebral
foramina; post-pelvic processes forming an apical fork; the lateral line very
irregular and sinuous, dropping below the midline of body between the pelvics
and origin of anal fin; poorly developed basibranchial dentition; young with
strong vertical bars (young of the other genera lack such bars); and completely
scaled cheeks. The most notable thing about Hypacanthus is the lack of any
obvious specialized characters (except perhaps the lateral line). The eastern
Atlantic-Mediterranean distribution of Hypacanthus and its general occurrence
in subtropical and temperate waters supports the hypothesis that it evolved in
the Mediterranean region.

The exact phyletic position of Hypacanthus is obscure owing to the lack of
specialized characters which might be useful in determining relationships. The
greatly reduced basibranchial dentition of Hypacanthus (fig. 24a), suggests,
however, that it did not share a common ancestor with Parona-Scomberoides.

Trachinotus, a circumtropical genus consisting of about 19 species (F. H.
Berry, personal communication) that is at least superficially similar to Hypa-
canthus, differs from Hypacanthus and the other genera most notably in having
dentary and premaxillary teeth that are poorly developed or absent; upper pha-
ryngeals that are noticeably enlarged and covered with blunt, molariform teeth;
the parasphenoid expanded into a broad, flattened plate posteriorly; and the
basioccipital with a pair of lateral processes on its ventral surface. The complete

244 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

absence of basibranchial dentition in Trachinotus represents the end product of
a trend toward specialization in the opposite direction from that of Parona-
Scomberoides. All of the above characters are probably correlated and constitute
a single character complex associated with trophic ecology. Although the species
of Trachinotus comprise a well defined group, there are no characters that are
useful in establishing from what group of carangids they evolved.

SUBFAMILIAL RECOGNITION.

In comparing the Carangidae to the Scombridae, Starks (1911) made the
following observations which illustrate some of the problems encountered in
subfamilial partitioning of the Carangidae: ‘“‘The family Carangidae is a much
more compact group than the Scombridae. The gaps between the genera are
much smaller, and though the osteological characters, like the form of the body,
differ much in the extremes of variation, there are no sudden or complete
changes.” Although he recognized four subfamilies (Scomberoidinae, Trachin-
otinae, Naucratinae, and Caranginae), Starks (1911, p. 30) acknowledged the
largely artificial nature of these subfamilies by stating: “though Selene is not
nearly so closely related to Trachyurops |= Selar|—for instance—as some
forms are to each other that are here arranged in different subfamilies, there is no
place to draw a dividing line.” Parona and Scomberoides are probably no more
divergent phylogenetically than are some genera presently placed together in
the subfamily Caranginae. Within the present classification of the family, Parona
might be recognized as a monotypic tribe in the subfamily Scomberoidinae.
Hypacanthus would require its own subfamily, since its relationship to other
carangid genera is obscure. Until the relationships of the other genera are better
understood, we prefer not to recognize any carangid subfamilies.

The following characters shared, in combination, by Oligoplites, Scomberoides,
Parona, Trachinotus, and Hy pacanthus distinguish them as a group from all other
carangids: lateral line without scutes; body moderately to strongly compressed;
second dorsal-fin base approximately equal in length to anal-fin base; and haemal
spine of first caudal vertebra and first anal pterygiophore firmly attached to form
a strong strut. These characters may, however, have little or no phylogenetic
significance. Other characters possessed by these five genera are summarized in
table 7.

PHYLOGENETIC POSITION OF Campogramma REGAN, 1903
(Figure 26)

Wheeler (1963) illustrated this poorly known eastern Atlantic-Mediterranean
carangid and discussed its nomenclatural history. We agree with Wheeler (1963,
p. 538) that the description of Centronotus glaycos Lacépéde, 1803 is sufficiently
diagnostic to determine its specific identity, thereby establishing it as a senior
synonym of the widely used but nomenclaturally invalid name Campogramma

VoL. XXXIX]

TABLE 7.

SMITH-VANIZ & STAIGER: CARANGIDAE

Summary of selected characters used to define genera.

245

Character

epiotics united
internally

fronto-supraoccipital
crest

suborbital shelf

suborbitals 2—4
expanded posteriorly

interosseous space
between dentary and
articular

supramaxilla
premaxillary frenum

dentary teeth

‘mesopterygoid’
teeth

branchiostegal rays

basibranchial tooth
plates

caudal vertebrae

inferior
vertebral foramina

epural bones
predorsal bones

dorsal-fin rays

posterior dorsal-
and anal-fin rays

pelvic fins

postpelvic
processes

postcoracoid
process

first anal
pterygiophore
expanded antero-
laterally, forming roof
over anal spines

body scales

cheek scales

lateral line

swimbladder

Trachinotust Hypacanthus Parona Scomberoides Oligoplites
no no yes yes yes
high low high low low
present present present present absent
no no no no yes
well developed moderate well developed minute minute or
absent
absent small large small absent
absent absent present present present
(narrow ) (wide) (wide)
villiform band wide band narrow band 2 rows 2 rows
(often absent (single row
in adults) posteriorly )
absent absent present (free- present (ankylosed absent
floating) to bone)
7-8 8 9 8 7
absent poorly well developed well developed well developed
developed (large median (paired plates) (paired plates)
plate)
14 14 17 16 16
absent present absent absent absent
3 3 2 2
4or 5 4 3 or 4 4 to 6
(rarely 5) (rarely 4) (rarely 4)
VI-VII+ VII+ VI-VII+ VI-VII+ IV-VI
I, 16-29 I, 19-21 I, 32-38 I, 19-21 (rarely VI)+
1 ail
normal normal normal semidetached semidetached
finlets finlets
present present absent present present
united forming apical united united united
distally fork distally distally distally
moderate moderate well developed absent absent
no no no yes yes
(fig. 15a) (fig. 15b) like Scom-
beroides
oval-shaped oval-shaped, oval-shaped broadly lanceo- needlelike
becoming strongly late to needlelike
lanceolate on (fig. 3)
breast
present or present absent absent absent
absent
normal irregular and 5—9 dorsal normal normal
sinuous dropping — branches
below midline of
body between pel-
vics and anal fin
simple (young) simple (young) simple strongly bifid strongly bifid

to moderately
bifid (adults)

to moderately
bifid (adults)

(all sizes)

(all sizes)

(all sizes)

1 Several unique characters of Trachinotus not included here are discussed in the text under Relationships

and Zoogeography.

246 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

FIGURE 26. Campogramma glaycos, BMNH 1892.4.20.3, 377 mm. SL. Mediterranean.
Snout appears unnaturally short due to bent condition of specimen.

vadigo (Risso, 1810). We do not agree with Wheeler who treated C. glaycos as
a nomen oblitum because past workers either overlooked the name or considered
it unavailable because of homonymy with Lichia glauca of authors. In fact,
C. glaycos can not be considered a homonym of the differently spelled L. glauca.
Campogramma liro proposed by Dollfus (1955, p. 58) as a replacement name
for C. vadigo Risso is unnecessary. A junior synonym of Campogramma glaycos
(Lacépéde) overlooked by Wheeler is Oligoplites africana Delsman (1941).
Bauchot and Blanc (1963, p. 53) proposed Solagmedens with Oligoplites africana
as the type species. They apparently were unaware of the description of
Cam pogramma.

Recent authors have assigned the monotypic Cam pogramma to the subfamily
Trachinotinae, but without presenting substantiating data. Characters used to
define the Trachinotinae that are associated with trophic ecology, such as en-
larged pharyngeals and expansion of the parasphenoid posteriorly into a broad
flattened plate, are absent in Campogramma. Three external characters of Cam-
pogramma that are shared, in combination, exclusively with naucratinine (=
Seriolinae) genera are: lateral line without scutes; body terete, only slightly
compressed; anal-fin base shorter than second dorsal-fin base.

The osteology of Campogramma suggests that it evolved from a form close
to Seriola. The haemal spine of the first caudal vertebra is weakly articulated
with the first anal pterygiophore, not enlarged and indistinguishable from the
haemal spines of other caudal vertebrae. In lateral view the anterior margin of
the first anal pterygiophore is distinctly concave, with a free distal tip, as in
most species of Serviola (Mather, 1971, p. 186, fig. 3b). The last three pairs of
parapophyses are connected dorsally by a bony bridge which encircles the haemal
canal, and the last two are united distally to form a single haemal spine. As
noted by Starks (1911, p. 29), in the other carangid subfamilies the posterior
parapophyses are distinct, and not united as a single haemal spine, though a
bony bridge connects their bases. Other characters shared with Seriola are:

VoL. XXXIX] SMITH-VANIZ & STAIGER: CARANGIDAE 247

zygapophyses of exoccipital joined beneath foramen magnum; myodome with a
large posterior opening; large supramaxilla; premaxillary spur present; branchios-
tegal rays 3 + 4 (epihyal + ceratohyal); vertebrae 10 + 14 = 24; caudal skel-
eton with 3 epurals; cluster of free-floating, irregularly shaped patches of minute
granular teeth overlying mesopterygoids; and swimbladder not bifid.

Campogramma is distinguished from all other genera of Naucratinae in
having a single row of caniniform teeth in both jaws, naked cheeks, partially
naked breast, and slightly falcate pectoral fins.

MATERIAL EXAMINED. BMNH 1893.2.24.3, 112 mm. SL, cleared and
stained; BMNH 1892.4.20.3, 377 mm. SL, x-rayed.

SUMMARY

The carangid fish genera Scomberoides, Oligoplites, Parona, and Hypacanthus
are revised. These four genera, along with Trachinotus, are distinguishable as
a group from all other genera of Carangidae. Campogramma, formerly placed
in the subfamily Trachinotinae, is considered to be more closely related to
Seriola.

Scomberoides consists of four recent species, S. commersonianus Lacépéde,
S. tala (Cuvier), S. lysan (Forskal), and S. tol (Cuvier). Scomberoides spinosus
(Smirnov), described from Lower Miocene deposits, is believed to be closely
related to S. tol. Scomberoides lysan, consistently treated as a senior synonym
of S. commersonianus, is instead a senior synonym of the widely used S. tolo-
oparah (Riippell). The species of Scomberoides range from the Indian Ocean
and Red Sea into the central Pacific; only one species, S. /ysan, occurs east of
the Solomon Islands and frequents entirely oceanic habitats.

Oligoplites consists of five species, O. saurus (Bloch and Schneider), O.
saliens (Bloch), O. palometa (Cuvier), O. refulgens Gilbert and Starks, and
O. altus (Ginther). Oligoplites mundus Jordan and Starks is placed in the
synonymy of O. altus. The genus is restricted to the eastern Pacific and west-
ern Atlantic oceans; only O. saurus occurs in both oceans.

Parona, a monotypic genus restricted to the western South Atlantic Ocean,
is believed to be most closely related to Scomberoides and Oligoplites. Parona
signata (Jenyns) is unique among carangids in that it lacks pelvic fins.

Hypacanthus, also monotypic, has a littoral distribution in the Mediterranean
Sea, along the Atlantic coast of Africa, and the southern Indian Ocean. Hypa-
canthus ania (Linnaeus) is the least specialized of the carangids studied.

Osteological descriptions are given for Scomberoides, Oligoplites, Parona,
and Hypacanthus. Of the genera treated, Scomberoides and Oligoplites are the
most closely related, with Oligoplites the most specialized. Within the frame-
work of the present classification of the family Carangidae the two monotypic
genera would require their own subfamilies. The phyletic relationship of all
of these genera is such that we do not recognize any subfamilies.

248 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

APPENDIX

List oF NoMINAL SPECIES OF Scomberoides AND Oligoplites, INCLUDING
SomME NAMES ORIGINALLY ASSIGNED TO ONE OF THESE GENERA (OR THEIR
SYNONYMS) THAT WERE MISAPPLIED OR ARE UNIDENTIFIABLE.

The following list gives in order: (1) the scientific name in alphabetical order
by species as it originally appeared (capitalized specific names in lower case,
however); (2) the author or authors (Cuvier and Valenciennes, 1831 is ab-
breviated to C. & V., 1831, and Jordan and Evermann, 1898 is abbreviated to
J. & E. 1898); (3) date of publication; (4) page reference (see literature cited
section for complete reference); the genus and species to which we currently
assign the species, if such is possible. Included are several nominal species
described in Scomberoides or Chorinemus that are unidentifiable. The basis for
our assignment of nominal species of Scomberoides is discussed in the text under
their senior synonyms. Unless stated otherwise in the text, synonymy of Oligo-
plites species follows that of Schultz (1945).

Species, author, publication date, and page

reference Present allocation

Oligoplites africana Delsman, 1941, p. 52 Campogramma glaycos
Lichia albacora Guichenot, 1848, p. 231 ?Oligo plites altus

Chorinemus altus Giinther, 1868, p. 433 Oligo plites altus

Scomber calcar Bloch, 1793, p. 46 2Oligoplites saliens
Scomberoides commersonnianus Lacépeéde,

1802, p. 50 Scomberoides commersonianus
Chorinemus delicatulus Richardson, 1846,

p. 269 Scomberoides commersonianus
Chorinemus exoletus Ehrenberg in C. & V.,

1831, p.379 (278) Scomberoides commersonianus
Chorinemus farkharrii Cuvier in C. & V.,

1831, p. 388 (285) unidentifiable (?Scombridae)
Scomberoides formosanus Wakiya, 1924,

p. 236 Scomberoides tol
Scomberoides formosanus Oshima [non

Wakiya], 1925, p. 349 Scomberoides lysan

Scomber forsteri Bloch & Schneider, 1801,

p. 26 Scomberoides lysan
Chorinemus guaribira Cuvier in C. & V.,

1831, p. 393 (289) Oligo plites palometa
Chorinemus hainanensis Chu and Cheng,

1958, pool? Scomberoides tala

Oligo plites inornatus Gill, 1863, p. 166 Oligo plites saurus inornatus

VoL. XXXIX]

Chorinemus lanceolatus Girard, 1858,

p. 168

Chorinemus leucopthalmus Richardson, 1846,
p. 269

Scomber lysan Forskal, 1775, p. 54
Scomber maculatus Forster, 1844, p. 195
Chorinemus maculosus Saville-Kent, 1893,
p. 369

Scomber madagascariensis Shaw, 1803,

p. 590

Chorinemus mauritianus Cuvier in C. & V.,
1831, p. 382 (281)

Chorinemus moadetta Ehrenberg in C. & V.,
1831, p. 382 (280)

Thynnus moluccensis Gronovius in Gray,
1854, p. 121

Oligoplites mundus Jordan & Starks in

J. &E., 1898, p. 2844

Scomberoides noelii Lacépede, 1802, p. 50
Chorinemus occidentalis Gunther, 1860,
p. 475

Chorinemus orientalis Temminck & Schlegel,
1844, p. 106

Scomberoides oshimae Whitley, 1951, p. 65
Chorinemus palometa Cuvier in C. & V.,
1831, p. 392 (288)

Eleria philip pina Jordan & Seale, 1905, p. 744
Lichia quiebra Quoy & Gaimard, 1824, p. 365
Oligoplites rathbuni Miranda-Riberio,
1915, p. 8

Oligoplites refulgens Gilbert & Starks,

1904, p. 73

Scomber saliens Bloch, 1793, p. 41
Chorinemus saltans Cuvier in C.& V.,

1831, p. 393 (289)

Scomberoides saltator Lacépede, 1802, p. 50
Chorinemus sancti petri Cuvier in C. & V.,
1831, p.379 (279)

Scomber saurus Bloch & Schneider, 1801,
peo2

SMITH-VANIZ & STAIGER:

CARANGIDAE 249

Oligo plites s. saurus
Scomberoides commersonianus
Scomberoides lysan

Scomberoides lysan

nomen nudum (Scomberoides
species )

Scomberoides commersonianus
Scomberoides lysan
Scomberoides lysan
Scomberoides lysan

Oligo plites altus
unidentifiable (?Scombridae )

Oligoplites s. saurus

Scomberoides lysan
Scomberoides lysan

Oligo plites palometa
Scomberoides tala
Oligo plites (?s. saurus)

Oligo plites s. saurus

Oligo plites refulgens
Oligo plites saliens

Oligo plites s. saurus
Oligo plites saliens

Scomberoides lysan

Oligo plites s. saurus

250 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Palaeoscomber spinosus Smirnov, 1936,

p. 49-59 Scomberoides spinosus
Chorinemus tala Cuvier in C. & V., 1831,

pe ov? (277) Scomberoides tala
Chorinemus tol Cuvier in C. & V., 1831,

p. 385 (283) Scomberoides tol
Chorinemus toloo Cuvier in C. & V., 1831,

pest GATT) Scomberoides (?tala)
Lichia tolooparah Riippell, 1828, p. 91 Scomberoides lysan

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1828-1830. Atlas zu der Reise im nordlichen Afrika. Zoologie. Fische des Rothen Meeres.
Frankfurt am Main, 144 pp., 35 pls.
RUSSELL, PATRICK
1803. Descriptions and figures of two hundred fishes; collected at Vizagapatam on the
coast of Coromandel. London, vol. 1, pp. vii + 78, pls. 1-100; vol. 2, pp. 1-85,
pls. 101-198.
SAVILLE-KEnT, W.
1893. The Great Barrier Reef of Australia, its products and potentialities. London,
387 pp., 64 pls.
ScHULTZ, LEONARD P.
1945. The leatherjackets, Carangid fishes of the genus Oligoplites Gill, inhabiting Amer-
ican waters. Journal of the Washington Academy of Sciences, vol. 35, no. 10,
pp. 330-336.

Vor. XXXIX] SMITH-VANIZ & STAIGER: CARANGIDAE 255

SHAW, GEORGE

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pp. x1 + 187-632, pls. 26-92.
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SmitH, C. LAVETT

1971. A revision of the American groupers: Epinephelus and allied genera. Bulletin of

the American Museum of Natural History, vol. 146, art. 2, pp. 69-241, 41 figs.
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1962. The subocular shelf of fishes. Journal of Morphology, vol. 110, no. 1, pp. 1-10,
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SmiTH, J. L. B.

1949. The sea fishes of South Africa. Central News Agency Limited, Cape Town, 550 pp.,
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1968. Studies in carangid fishes No. 4. The identity of Scomber sansun Forskal, 1775.
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1970. Studies in carangid fishes no. 5. The genus Chorinemus Cuvier, 1831 in the
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Suzuxk1, KtyvosH1

1962. Anatomical and taxonomical studies on the carangid fishes of Japan. Report of
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1939. Risultati ittiologici del viaggio di circumnavigazione del globo della R. N.
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1963. The nomenclature of the European fishes of the subfamily Trachinotinae. Annals

GOs 1 ~ «CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
2
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1951. New fish names and records. Proceedings of the Royal Zoological Society of
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lological Laborztory
LIBRAPY

TH 4 8 1973
Weeds Acta, iviass.

PROCEEDING

OF THE

CALIFORNIA ACADEMY OF SCIENCES

FOURTH SERIES

Vol. XXXIX, No. 14, pp. 257-272; 4 figs. July 9, 1973

CTENACIS AND GOLLUM, TWO NEW
GENERA OF SHARKS
(SELACHIT; CARCHARHINIDAE)

iy
L. J. V. Compagno

Division of Systematic Biology, Stanford University, Stanford, California

ABSTRACT: Two new genera are described for two species of carcharhinid sharks
formerly placed in Triakis Miller and Henle. Ctenacis, new genus, is proposed for
Triakis fehlmanni Springer, 1968. Gollum, new genus, is proposed for Triakis
attenuata Garrick, 1954. Both genera are closest to Proscyllium Hilgendorf and
Eridacnis H. M. Smith within the Carcharhinidae. Ctenacis is most similar to
Eridacnis and somewhat less so to Proscyllium. Gollum is more distant from
Proscyllium, Ctenacis, and Eridacnis than the latter three genera are from each other.

Garrick (1954) described a peculiar new shark, Triakis attenuata, that in
many features differs strikingly from typical species of that genus. Subsequent
workers on Tyviakis, as Smith (1957), Kato (1968), and Springer (1968),
accepted Garrick’s generic placement of ‘attenuata.’ In addition, Springer (1968)
described another new species of Triakis, T. fehlmanni, that also differed greatly
from the type-species, 7. scvllium, and its closest relatives. Springer regarded
the generic position of ‘fek/manni’ as provisional pending a revision of Triakis.

Compagno (1970) proposed a preliminary reorganization of Triakis. He
removed T. leucoperiptera from Triakis and assigned it to the genus Hemitriakis.
Proscyllium was resurrected for T. habereri and T. venusta and Calliscyllium
were synonymized with it. Eridacnis and Neotriakis were previously synonymized
with Triakis by Kato (1968), but Compagno revived Eridacnis for E. radcliffe,
Neotriakis sinuans, and Triakis barbouri. Neotriakis was synonymized with
Eridacnis. Following several previous workers, Compagno transferred Triakis
henlei to Mustelus.

Triakis, as restricted by Compagno (1970), contained a group of closely

[257]

CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

bo
OL
Co

similar species, T. scvllium, T. semifasciata, T. maculata, and T. acutipinna,
that seem to grade into Mustelus through intermediate forms as M. migro-
punctatus, M. megalopterus, M. henlei, and M. dorselis. Triakis attenuata
was considered to be probably distinct from Triakis proper but inadequate data
on the species at the time precluded more definite generic assignment. Tviakis
fehlmanni was treated as a doubtful appendage to Eridacnis that might merit
generic or subgeneric distinction.

Reconsideration of available data on T. fehlmanni indicates that this species
should be separated from Eridacnis and Triakis in a new genus. Examination
of two specimens of 7. attenuata confirmed my earlier opinion that ‘attenuata’
was generically distinct from Tvriakis and required a new genus also.

The terminology used here for external morphology, vertebral counts, and
dentition is from Compagno (1970). Clasper terminology follows Leigh-Sharpe
(1921) and Compagno and Springer (1971). Terms for vertebral calcification
patterns are from Ridewood (1921). Cranial terms are modified from Gegenbaur
(1872), Allis (1923), and Holmgren (1941).

Ctenacis Compagno, new genus

Type spectes. Tviakis fehlmanni Springer, 1968.

DEFINITION. Head very broad, depressed, its length 23 percent of total
length in adult. Head length greater than distance between pectoral and pelvic
bases. Snout outline subparabolic in dorsoventral view, not bell-shaped. Snout
short, length 6.5 percent of total length. Eyes high on sides of head, above
level of nostrils by a space about equal to eye height. Subocular ridge very
strong, not indented on its dorsal surface. Eyes not visible in ventral view of
head. Eyes elongate-elliptical, about 2.5 times as long as high. Nictitating
lower eyelid rudimentary, with its edge horizontal. Subocular pouch very
shallow, with its interior surface covered with denticles.

Spiracles present, their length about 5 times in eye length. Shortest gill
opening over “4 as long as longest. Gill rakers present on gill arches.

Internarial space about 1.2 times nostril width. Anterior nasal flaps are
broad, low, triangular lobes with fringed posterior edges, not elongated, barbel-
like, or tubular. Posterior edges of anterior nasal flaps close to mouth but
separated from level of upper symphysis by distance equal to % of nostril
width. Posterior nasal flaps large, with fringed edge. No nasoral grooves.

Mouth distinctly angular in shape, with edges of lower jaw nearly straight.
Large papillae present in buccal cavity. Labial furrows extremely short, es-
sentially confined to the mouth corners.

Dignathic heterodonty weak, upper teeth with higher and narrower crowns
than lowers at symphysis, lower teeth more comblike than uppers at ends of
dental bands. No disjunct monognathic heterodonty. In both jaws teeth de-
crease in size and height relative to root width towards rictuses. In the lower

Vor. XXXIX] COMPAGNO: NEW SHARK GENERA 259

jaw premedial cusplets increase in size and number and the primary cusp
shifts postlaterally on the crown base towards the rictus, producing comblike
posterior teeth.

Tooth rows 86/88, series functional 4-5/5—7. Teeth not bladelike, without
serrations or sharp edge. Premedial cusplets present on all teeth except a
few in the region of the symphysis. Postlateral cusplets present on all teeth.
Primary cusp present on all teeth, narrow, erect or slightly oblique. Crown
foot with a strong basal ledge overlapping a deep basal groove. Transverse
ridges present on all teeth, extending on to the cusps of uppers, confined to
the bases in lowers. Roots of teeth broad, flat, low, platelike. No transverse
groove and notch on attachment surface of roots. Teeth not extending onto
sides and ventral surfaces of lower jaw.

No interdorsal ridge, lateral dermal keels on the caudal peduncle, or pre-
caudal pits. Length of head and trunk (from snout tip to vent) about 46 per-
cent of total length.

Denticles below first dorsal fin with crowns about as wide as long or
slightly longer than wide in adult. Denticle crowns with a strong medial cusp
and a pair of strong medial ridges; a pair of weaker lateral ridges are also
present and a pair of very weak lateral cusps may be present or not.

Pectoral fin skeleton projecting about %% of pectoral anterior margin length
into fin. Longest distal radials of pectoral much shorter than longest proximal
ones. Distal pectoral radials with parallel edges and truncate tips.

Pelvic bases over twice as far from second dorsal base as from first dorsal
base. Posterior margins, free rear tips, and posterior margins of pelvic fins
not attenuate. Clasper morphology unknown.

Midpoint of first dorsal base about twice as far from pectoral bases as
from pelvic bases. Free rear tip of first dorsal posterior to pelvic origins.

Second dorsal fin nearly as large as first, its height almost %o of height of
first, its base length 1.2 times base length of first dorsal. Posterior margin of
second dorsal weakly concave.

Anal fin much smaller than second dorsal, its height about half of second
dorsal height, its base length about * of second dorsal base length. Anal origin
posterior to second dorsal origin by distance about % of second dorsal base
length. Anal insertion under second dorsal insertion. Posterior margin of anal
straight.

Caudal fin without distinct ventral lobe, postventral caudal margin not
notched or otherwise differentiated. Subterminal caudal margin about 4 of
terminal caudal margin. Caudal short, not tapelike, dorsal caudal margin 23
percent of total length. No lateral undulations in dorsal caudal margin. Ter-
minal sector of caudal about 4 times in dorsal caudal margin. No caudal crest
of denticles.

Cranium with paired lateral and unpaired medial rostral cartilages, fused

260 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

PREF oN
FM oC OCN

Ficure 1. Ctenacis fehlmanni, U. S. National Museum 202969, 46 cm. adult female,
neurocranium in dorsal (left) and ventral (right) views. Abbreviations: AF, anterior
fontanelle; BP, basal plate; CF, internal carotid foramen; EC, ectethmoid chamber; FM,
foramen magnum; HF, hyomandibular facet; LR, lateral rostral cartilage; MR, medial
rostral cartilage; NA, nasal aperture; NC, nasal capsule; NF, nasal fontanelle; NP, notch
for orbital process of palatoquadrate; O, orbit; OC, occipital condyle; OCN, occipital
centrum; OR, opisthotic ridge; OT, otic capsule; PC, preorbital canal; PF, profundus
foramen; PR, preorbital process; PRF, parietal fossa; PT, postorbital process; SC,
supraorbital crest; SF, stapedial foramen; SR, sphenopterotic ridge; SS, suborbital shelf.

at their tips to form a tripod (fig. 1). Rostral node (conjoined tips of rostral
cartilages) not yokelike, not penetrated by a rostral fenestra. Rostrum fairly
short, length of medial rostral cartilage from its base on the internasal septum
to the anterior tip of the rostral node about three times in nasobasal length
(distance from base of medial rostral cartilage to the posterior edge of the
occipital centrum; here used as an independent variable for cranial propor-
tions). Distance between bases of lateral rostral cartilages about equal to length
of medial rostral cartilage. Lateral rostral cartilages with their bases far an-
terior to the anterior fontanelle and not connected to the dorsal edge of the
fontanelle by a ridge.

Nasal capsules oval-shaped, their long axis transverse to the longitudinal
axis of the cranium. Transverse width of capsule from internasal septum to
lateral edge of capsule about twice its length across its anterior and posterior
walls. Greatest transverse width across nasal capsules about 1.3 in nasobasal
length. Nasal aperture and nasal fontanelle apparently broadly continuous
with each other, the fontanelle not separated from the nasal aperture by a

VoL. XXXIX] COMPAGNO: NEW SHARK GENERA 261

LR

lr OC OCN

Ficure 2. Gollum attenuatus, DM-3636, 955 mm. adult male, neurocranium in dorsal
(left) and ventral (right) views. Abbreviations as in figure 1, except: BR, basirostral
fenestra; ECN, lateral ectethmoid condyle.

Cm OR

bridge of cartilage. Greatest width of nasal aperture about six times distance
between nasal apertures across internasal septum. Ectethmoid chamber opening
inside nasal capsule on its posteroventral wall, with its aperture just above
posterior edge of nasal fontanelle and not separated from the fontanelle by a
horizontal ledge beneath it. No ectethmoid condyles or subethmoid fossa. No
ectethmoid foramen communicating between the ventral surface of the nasal
capsule and the ectethmoid chamber.

Anterior fontanelle subcircular in shape, its greatest width about 4.6 in
nasobasal length. Cranial roof convex between orbits.

No deep notch separating anterior end of suborbital shelf from nasal capsule.
Basal plate expanded just posterior to nasal capsules into a broad suborbital
ledge, the least width across which is about 1.7 in nasobasal length. Arterial
foramina on ventral surface of basal plate include a pair of small internal
carotid foramina about half as far from each other as from each of the
stapedial foramina distal to them. The paired stapedial foramina, for the

262 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Yi MR

f

or Shane a -

=
Re rm FG 1m

Ficure 3. Triakis semifasciata, LJVC (writer’s personal collection) 0067, 937 mm.
immature female, neurocranium in dorsal (left) and ventral (right) views. Abbreviations
as in figure 1, except: FG, foramen for glossopharyngeal (IX) nerve; FV, foramen for
vagus (X) nerve.

stapedial or orbital arteries, are tiny pinhole apertures. Basal plate nearly flat,
without keels.

Supraorbital crest present, connecting low preorbital and postorbital processes.
Edge of supraorbital crest arcuate in dorsal view. Dorsal edge of crest not
extending above level of cranial roof between the orbits. Width across nar-
rowest part of supraorbital crest about 2.6 in nasobasal length. Postorbital
processes not extending from supraorbital crest and sphenopterotic ridge, with
tips not bifurcated. Suborbital shelf very wide, greatest width across it 1.3
in nasobasal length.

Otic capsules short, not greatly enlarged or inflated. Length of otic capsule
about 4 in nasobasal length. Sphenopterotic ridge extending slightly lateral
to side of otic capsule.

Occipital condyles very small, without dorsolateral and ventrolateral plates
that partially sheath base of vagus nerve.

Total vertebral count 136 (1). Monospondylous precaudal (MP) centra
28.6, diplospondylous precaudal (DP) centra 35.2, and diplospondylous caudal
(DC) centra 36 percent of total vertebral count. Ratio of DP/MP centra
counts 1.23, DC/MP centra 1.26. “A” ratio (length of penultimate MP centrum/
first DP centrum) 138, “B” ratio (length/width of penultimate MP centrum)
114. No ‘stutter zone’ of alternating long and short centra in DP region.
Last few MP centra not greatly enlarged.

VoL. XXXIX] COMPAGNO: NEW SHARK GENERA 263

Vertebral centra without wedgelike intermedialia. Diagonal calcified lamel-
lae present in form of knoblike projections from calcified double cone.
Notochordal canal rather large, not blocked at apices of double cone.

Spiral intestinal valve present, with about 10 turns.

Mode of development not known, though possibly oviparous (see Springer,
1968, for discussion).

A color pattern of dark saddles, blotches, bars, and spots on a light back-
ground present.

Size small, the one specimen of C. feklmanni known at present being an
adult female 46 cm. long.

COMPARISON WITH OTHER GENERA. Ctenacis is a member of Compagno’s
(1970) group of ‘scyliorhiniform triakoids, which are morphologically inter-
mediate between scyliorhinids and genera such as Tviakis, Mustelus, and
Hemitriakis. Ctenacis is most similar to Fvridacnis, somewhat less so to
Proscyllium, and yet more distant from Gollum, the other new genus described
here. All four genera show sufficient similarity in external morphology, denti-
tion, cranial anatomy, and vertebral calcification pattern to form an apparently
related group.

For brevity comparisons of both Ctenacis and Gollum with other carcharhinid
genera is here limited to the other ‘scyliorhiniform’ genera and to Tviakis-
Mustelus, the generic complex to which ‘feklmanni’ and ‘attenuata’ were
originally assigned. Both genera are not particularly close to carcharhinid
from these genera by many characters given in the generic definitions.
genera other than Proscyllium and Eridacnis and can be readily distinguished

Eridacnis differs from Ctenacis in having a narrower head; anterior and
posterior nasal flaps not fringed; tooth rows less numerous, 55—78/63—77; head
and trunk slightly shorter, 38—43 percent of total length, and rather slim; pelvic
posterior margins, free rear tips, and inner margins elongate, somewhat at-
tenuate; caudal very long, tapelike, length 25-30 percent of total length;
cranium narrower, width across nasal capsules 1.4—-1.6 in nasobasal length;
nasal capsules nearly spherical; no diagona! calcified lamellae in vertebral
centra; no striped and spotted color pattern on body; and size smaller, less
than 370 mm. total length.

Proscyllium differs from Ctenacis in having a narrower and shorter head,
with length only 16-18 percent of total length; fifth gill opening less than
half length of third; anterior nasal flaps very large, reaching nearly to upper
symphysis; internarial space very narrow, nostril width 1.7—2.2 times inter-
narial; anterior and posterior nasal flaps not fringed; tooth rows less numerous,
46—62/49-59; head and trunk very short, only 39-41 percent of total length, very
slender; first dorsal origin well posterior to free rear tip of pectoral; anal origin
slightly anterior to second dorsal origin; cranium narrower, width across nasal
capsules 1.5 in nasobasal length; nasal capsules nearly spherical; total vertebral
counts 146-168 (6); and DP/MP ratio 1.6-1.8.

264 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Ctenacis is compared with Gollum in the account of the latter genus below.

Triakis and Mustelus differ from Ctenacis in having external, transitional,
or internal nictitating lower eyelids; no gill rakers or buccal papillae; anterior
nasal flaps elongated and lobate; posterior nasal flaps absent or rudimentary;
labial furrows long and extending well onto upper and lower jaws; lower teeth
towards ends of dental band not comblike, with cusplets reduced or absent;
roots of teeth usually subdivided on their attachment surface by a transverse
groove and notch; interdorsal ridge present; longest distal radials of pectoral
fin skeleton about as long as longest proximal radials or slightly longer;
midpoint of first dorsal fin base about equidistant between pectoral and pelvic
bases or definitely closer to pectoral bases; anal insertion slightly posterior to
second dorsal insertion; anal posterior margin concave to deeply notched in
adults; cranium with a deep notch separating the anterior end of the sub-
orbital shelf from the nasal capsule (fig. 3); postorbital process exserted from
the supraorbital crest and bifurcated distally; and vertebral centra with wedge-
shaped intermedialia.

In addition Triakis (including only 7. scvllium, T. semifasciata, T. maculata,
T. acutipinna, and probably ‘Mustelus’ megalopterus and ‘M.’ nigropunctatus )
differs from Ctenacis in having an arcuate mouth (fig. 4C); fewer tooth rows,
44—-65/34—56; and a well developed ventral caudal lobe in adults. Mustelus (in-
cluding ‘Tviakis’ henlez) also differs from Ctenacis in having a longer snout, with a
narrowly parabolic to almost angular shape in dorsoventral view and narrowly
wedgelike shape in lateral view; teeth extending onto sides and ventral surface
of lower jaw; teeth forming a regular pavement, with primary cusp typically
reduced or absent in most species; a tooth peg present on the inner face of the
crown; and ectethmoid condyles present on the nasal capsules.

DERIVATION OF NAME. Ctenacis, from Greek ktenos, comb, and akis, point
(feminine), in allusion to the comblike posterior teeth of this genus.

SPECIES. Only one, Ctenacis fehlmanni (Springer, 1968).

STUDY MATERIAL. The holotype and only known specimen of Ctenacis
feklmanni, U. S. National Museum 202969.

Gollum Compagno, new genus

Type species. Tvriakis attenuata Garrick, 1954.
DEFINITION. Head very broad, depressed, its length about % (21.3-21.6 per-
cent) of total length in adults. Head length considerably shorter than distance

>

Ficure 4. A-B, Gollum attenuatus, DM-4841, 1015 mm. adult female. Prebranchial
head in ventral (A) and lateral (B) views. C—D, Triakis scyllium, University of Michigan
Museum of Zoology 179067, 700 mm. immature male. Prebranchial head in ventral (C)
and lateral (D) views.

265

NEW SHARK GENERA

COMPAGNO

VoL. XXXIX]

266 CALIFORNIA ACADEMY OF SCIENCES [Proc, 47TH SEr.

between pectoral and pelvic bases. Snout with peculiar bell-shaped outline in
dorsoventral view (fig. 44). Snout very long, preoral length 8.4—8.6 percent
of total length in adults. Eyes high on sides of head, above level of nostrils
by a space equal to about half eye height or slightly less. Subocular ridge very
strong, with a distinct depression on its dorsal surface (fig. 4B). Eyes not
visible in ventral view. Eyes elongate-elliptical, with their apertures about 2%
to 2% times as long as high. Nictitating lower eyelid rudimentary, with its
edge horizontal. Subocular pouch very shallow and covered with denticles.

Spiracles present, their length about 3.8 to 8 times in eye length. Shortest
gill opening “4 to 45 as long as longest one. No gill rakers.

Internarial space about 1.8 to 1.9 times nostril width. Anterior nasal flaps
short and subtriangular, not tubular or barbel-like, with edges entire and not
fringed. Posterior edges of anterior nasal flaps close to mouth but separated
from the upper symphysis by distance equal to % to “4 of nostril width.
Posterior nasal flaps large, with fringed edges. Nasoral grooves absent.

Mouth distinctly angular in shape, with edges of lower jaw straight. No
large papillae in buccal cavity. Labial furrows very short, essentially confined
to mouth corners.

Dignathic heterodonty weak, virtually absent at symphysis but increasing
towards ends of dental bands, where lower teeth are more comblike than uppers.
No disjunct monognathic heterodonty. In both jaws teeth decrease in size and
height relative to root width toward rictuses. In the lower jaw premedial cusplets
increase in number and size relative to the primary cusp and the cusp shifts
postlaterally on the crown in a direction towards the rictus, producing comblike
posterior teeth. Gynandric heterodonty absent.

Tooth rows 96—99/108-114 (3), series functional 3—6/3—7. Crown not
sharp, serrated, or bladelike. Premedial and postlateral cusplets present on all
teeth. Primary cusp present on all teeth, narrow and erect. Crown foot with
a deep basal ledge overlapping a deep basal groove. Transverse ridges present
on teeth, numerous and small, extending onto primary cusp and cusplets. Roots
of teeth moderately high and deep, not subdivided on their attachment surface
by a transverse groove and notch. Teeth not extending onto sides and ventral
surface of lower jaw.

Interdorsal ridge present. No lateral dermal keels or precaudal pits present
on the caudal peduncle. Length of head and trunk about equal to tail length.

Denticles from sides of body below first dorsal fin with crowns much
longer than wide in adults. Denticle crowns with a strong medial cusp, a strong
medial ridge or a pair of medial ridges, a pair of short but strong lateral cusps,
and a pair of lateral ridges.

Pectoral fin skeleton projecting slightly less than % of pectoral anterior
margin length into fin. Longest distal radials of pectoral skeleton somewhat

VoL. XXXIX] COMPAGNO: NEW SHARK GENERA 267

shorter than longest proximal ones. Distal radials with parallel edges and
truncate tips.

Pelvic fin bases slightly closer to second dorsal base than to first dorsal base.
Pelvic posterior margins, free rear tips, and inner margins not attenuate.

Dorsal edges of clasper groove not fused together between apopyle and
hypopyle. A soft-edged, fleshy rhipidion present, with its dorsal surface ex-
posed for most of its length and not concealed by the cover rhipidion. Cover
rhipidion present, a small, short, square, fleshy flap. Exorhipidion present,
a low, fleshy flap lateral to pseudopera. Exorhipidion without clasper hooks.
A large pseudosiphon and pseudopera present.

Midpoint of first dorsal base about 1.3 times as far from pelvic bases
as from pectoral bases. Free rear tip of first dorsal well anterior to pelvic
origins.

Second dorsal fin about as large as first, its height equal or slightly larger
than height of first dorsal, its base length equal or slightly less than base length
of first dorsal. Posterior margin of second dorsal moderately concave.

Anal fin much smaller than second dorsal, its height 44—46 percent of first
dorsal height, base length about ' to *5 of second dorsal base. Anal origin
posterior to second dorsal origin by distance equal to 11 to 27 percent of second
dorsal base length. Anal insertion anterior to second dorsal insertion by distance
about 25 to 36 percent of second dorsal base length. Anal posterior margin
nearly straight, not deeply concave or notched.

Caudal fin with ventral lobe poorly developed in adults. Postventral caudal
margin not notched. Subterminal caudal margin about 7 to “40 of terminal
margin. Caudal short, not tapelike, in adults about % (19.4-21.2 percent) of
total length. No lateral undulations in dorsal caudal margin. Terminal sector
of caudal between 3 and 4 times in dorsal caudal margin. A weak supracaudal
crest of enlarged, circular or oval-crowned denticles present along the anterior
half of the dorsal caudal margin.

Cranium with lateral and medial rostral cartilages fused at their tips to
form a tripod (fig. 2). Rostral node not yokelike, not penetrated by a rostral
fenestra. Rostrum very long, length of medial rostral cartilage only 1.3 in
nasobasal length. Distance between bases of lateral rostral cartilages about 2.7
times in length of medial rostral. Lateral rostrals with their bases far anterior
to anterior fontanelle but with their dorsal edges connected to fontanelle by
a low ridge.

Nasal capsule oval in shape, transversely elongated. Transverse width of
capsule about 1.4 times its length. Greatest width across nasal capsules 1.4
in nasobasal length. Nasal aperture and nasal fontanelle broadly continuous
with each other, the fontanelle not separated from the nasal aperture by a bridge
of cartilage. Greatest width of nasal aperture about 5 times distance between
nasal apertures. Ectethmoid chamber opening inside nasal capsule on its

268 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

posteroventral wall, with its aperture just above posterior edge of nasal fontanelle
and not separated from the latter by a horizontal ledge beneath it. No true
ectethmoid condyles, though a pair of low condyles are present on the postero-
ventral surfaces of the nasal capsules (but well lateral to the position of the
ectethmoid condyles of carcharhinoid genera that have them). Subethmoid
fossa absent. No separate ectethmoid foramen on the ventral surface of the
nasal capsule.

Anterior fontanelle rhomboidal in shape, its greatest width only 7 times
in nasobasal length. Cranial roof flattened between orbits.

No deep notch separating suborbital shelf from nasal capsule. Basal plate
expanded just posterior to nasal capsules into a broad suborbital ledge, the
least width across which is about 2.0 in nasobasal length. Arterial foramina
on ventral surface of basal plate include a pair of small internal carotid foramina
about half as far from each other as from each of the tiny stapedial foramina
distal to them. Basal plate slightly arched, without keels.

Supraorbital crest present, with an arcuate lateral edge. Dorsal edge of
crest extending well above level of cranial roof between orbits. Width across
narrowest part of crest about 3.6 in nasobasal length. Postorbital processes
narrow, exserted beyond supraorbital crest and sphenopterotic ridge, with tips
not bifurcated. Suborbital shelf very wide, greatest width across shelves 1.4
in nasobasal length.

Otic capsules short, not greatly enlarged and inflated. Length of otic
capsule about 4 in nasobasal length. Sphenopterotic ridge extending medial
to sides of otic capsule.

Occipital condyles very small, without dorsal and ventral covering plates for
base of vagus nerve.

Total vertebral counts 166 (2). MP centra 30.7, DP centra 35.6, and DC
centra 33.7 percent of total counts. DP/MP ratio 1.16, DC/MP ratio 1.10.
“A” ratio 129 to 141, “B” ratio 87 to 99. No stutter zone. Last few MP
centra not greatly enlarged.

Vertebral centra without wedgelike intermedialia or diagonal calcified lamel-
lae. Notochordal canal rather large through apices of calcified double cones.

Valvular intestine with a spiral valve having about 11 turns.

Development ovoviviparous (Kato, 1968; J. A. F. Garrick, personal com-
munication ).

Ground color brownish gray dorsally, lighter below. No color pattern.

Size small-medium, males mature at 932 to 955 mm. total length, females
at 1015 mm.

COMPARISON WITH OTHER GENERA. Gollum belongs to Compagno’s (1970)
group of ‘scyliorhiniform triakoids,’ in which it is a singular and isolated member.
Ctenacis, Proscyllium, and Eridacnis all differ from Gollum in having shorter
snouts, only 4.3 to 6.7 percent of total length and not bell-shaped in dorsoventral

VoL. XXXIX] COMPAGNO: NEW SHARK GENERA 269

view; no concavity or depression on the dorsal surface of the subocular ridge;
gill rakers present; internarial space only 0.5 to 1.2 times nostril width; large
papillae present in buccal cavity; fewer tooth rows, 46-86/49-88; interdorsal
ridge absent; pelvic bases much closer to first dorsal base than second dorsal
base; midpoint of first dorsal base much closer to pelvic bases than pectoral
bases; first dorsal free rear tip over or posterior to pelvic origins; no supracaudal
crest of denticles; rostral cartilages much shorter, medial rostral length 3 to 4
in nasobasal length; no lateral condyle on posteroventral wall of nasal capsule;
postorbital process not exserted from supraorbital crest; sphenopterotic ridge
not medial to sides of otic capsule; and size smaller, not exceeding 700 mm.
total length when adult.

In addition Ctenacis differs from Gollum in having the head longer than
the distance between pectoral and pelvic bases; second dorsal insertion over
the anal insertion; distance across narrowest part of supraorbital crest greater,
2.6 in nasobasal length; total vertebral count 136; vertebral centra with knoblike
diagonal calcified lamellae; and a color pattern of dark blotches and spots.
Eridacnis also differs from Gollum in having the head about equal to the distance
between pectoral and pelvic bases or longer than it; posterior nasal flaps not
fringed; gynandric heterodonty more or less developed, in males spikelike teeth
present in one or both jaws at the symphysis; head and trunk considerably
shorter than the tail; pelvic inner margins, free rear tips, and posterior margins
elongate-attenuate; anal origin under second dorsal origin or slightly anterior
to it; caudal very long, tapelike, with dorsal caudal margin 24 to 30 percent of
total length; and total vertebral counts only 113 to 144. Proscyllium also differs
from Gollum in having the shortest gill opening less than half as long as the
longest; anterior nasal flaps very large, broadly triangular, and nearly reaching
the upper symphysis; edges of posterior nasal flaps not fringed; head and trunk
only 39 to 41 percent of total length; dorsal edges of clasper groove fused
together between hypopyle and apopyle; exorhipidion very strong, with clawlike
clasper hooks on its distal edge; first dorsal origin well posterior to pectoral free
rear tip; anal origin slightly anterior to second dorsal origin; DP/MP ratio 1.6
to 1.8; knoblike diagonal calcified lamellae present in vertebral centra; develop-
ment oviparous; and a color pattern of dark spots and bars present.

Triakis and Mustelus differ from Gollum in lacking a bell-shaped snout
profile in dorsoventral view (fig. 4C); no concavity or depression in dorsal
surface of subocular ridge (fig. 4D); nictitating lower eyelid external or transi-
tional in young but usually transitional or internal in adults; anterior nasal
flaps somewhat elongated; posterior nasal flaps rudimentary or absent; tooth
roots low, with transverse groove and notch usually present; crowns of teeth
towards ends of tooth band not comblike, with cusps and cusplets reduced or
absent; longest distal radials of pectoral fin equal in length or somewhat longer
than longest proximal radials; claspers without exorhipidion; anal fin insertion

270 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

slightly posterior to second dorsal insertion; anal posterior margin moderately
concave to deeply notched; no supracaudal crest of denticles; rostrum shorter,
length of medial rostral cartilage 1.9 to 4.4 in nasobasal length; bases of lateral
rostral cartilages not connected to edge of anterior fontanelle by a ridge; least
distance across supraorbital crest 1.5 to 2.3 in nasobasal length; suborbital
shelf separated from nasal capsule by a deep notch; no suborbital ledge between
nasal capsule and suborbital shelf; postorbital processes bifurcated distally;
sphenopterotic ridge above side of otic capsule or slightly distal to it, not medially
situated; and vertebral centra wtih intermedialia and diagonal calcified lamellae.

In addition Triakis differs from Gollum in having a thick short snout (fig.
4D), bluntly rounded or obtusely triangular in dorsoventral view (fig. 4C)
and bluntly rounded in lateral view; an arcuate mouth; very long labial furrows
extending along margins of upper and lower jaws; fewer tooth rows, 44—-65/
34-56; and a well developed ventral caudal lobe in adults. Mustelus also differs
from Gollum in having a snout less acutely wedge-shaped in lateral view; labial
furrows usually longer, extending well onto the upper and lower jaws; teeth
forming a regular pavement, with primary cusps and cusplets reduced or absent
in most forms; and a true ectethmoid condyle present on the nasal capsule.

DERIVATION OF NAME. Gollum (treated as a masculine noun), named for
the antihero of J. R. R. Tolkien’s Lord of the Rings trilogy, to whom this shark
bears some resemblance in form and _ habits.

Species. Only one known, Gollum attenuatus (Garrick, 1954).

STUDY MATERIAL. Dominion Museum (DM) 3636, 955 mm. adult male,
from about 25 miles off Cape Brett, North Island, New Zealand; DM-4841,
1015 mm. adult male, off Karamea Bight, South Island, New Zealand.

ACKNOWLEDGMENTS

Stewart Springer (National Marine Fisheries Service Systematics Laboratory,
U. S. National Museum) discussed the generic classification of Ctenacis
fehlmanni with me on several occasions both in person and in correspondence.
In addition, he allowed me to examine the holotype of this species and dissect
a ‘door’ in its head to expose the neurocranium. J. A. F. Garrick (Victoria
University, Wellington, New Zealand) advised me on the classification of Gollum
attenuatus and of additional specimens of this species collected since his original
description. J. M. Moreland (Dominion Museum, Wellington, New Zealand)
sent me two specimens of G. attenuatus.

LITERATURE CITED

ALLis, EpwARD PHELPS, JR.
1923. The cranial anatomy of Chlamydoselachus anguineus. Acta Zoologica, vol. 4,
pts. 2-3, pp. 123-221, pls. 1-23.
Compacno, L. J. V.
1970. Systematics of the genus Hemitriakis (Selachii: Carcharhinidae), and related

VoL. XXXIX] COMPAGNO: NEW SHARK GENERA 271

genera. Proceedings of the California Academy of Sciences, Fourth Series,
vol. 38, no. 4, pp. 63-98, figs. 1-8.
Compacno, L. J. V., and STEWART SPRINGER
1971. Jago, a new genus of carcharhinid sharks, with a redescription of 7. omanensis.
Fishery Bulletin, vol. 69, no. 3, pp. 615-626, figs. 1-6.
Garrick, J. A. F.
1954. Studies on New Zealand Elasmobranchii. Part III. A new species of Triakis
(Selachii) from New Zealand. Transactions of the Royal Society of New
Zealand, vol. 82, part 3, pp. 695-702, figs. 1-2.
GEGENBAUR, CARL
1872. Untersuchungen zur Vergleichenden Anatomie der Wirbelthiere. Drittes Heft.
Das Kopfskelet der Selachier, ein Beitrag zur Erkenntnis der Genese des
Kopiskeletes der Wirbelthiere. Wilhelm Engelmann, Leipzig. Pp. i-x, 1-316,
pls. 1-22.
Hortmcren, NIts
1941. Studies on the head in fishes. Embryological, morphological and phylogenetical
researches. Part II: Comparative anatomy of the adult selachian skull, with
remarks on the dorsal fins in sharks. Acta Zoologica, vol. 22, part 1, pp.
1-100, figs. 1-74.
Kato, SuSUMU
1968. Triakis acutipinna (Galeoidea, Triakidae), a new species of shark from Ecuador.
Copeia, 1968, no. 2, pp. 319-325, figs. 1-2.
LEIGH-SHARPE, W. HAroLp
1921. The comparative morphology of the secondary sexual characters of elasmobranch
fishes. The claspers, clasper siphons, and clasper glands. Memoir II. Journal
of Morphology, vol. 35, pp. 359-380, figs. 1-15.
RipEwoop, W. G.
1921. On the calcification of the vertebral centra in sharks and rays. Philosophical
Transactions of the Royal Society of London, Ser. B., Zoology, vol. 210,
pp. 311-407, figs. 1-38.
SMmrrHee|i i. B.
1957. A new shark from South Africa. South African Journal of Science, vol. 53,
no. 10, pp. 261-264, figs. 1-2.
SPRINGER, STEWART
1968. Triakis fehlmanni, a new shark from the coast of Somalia. Proceedings of the
Biological Society of Washington, vol. 81, pp. 613-624, figs. 1-5.

7 76

Ce.
oneagy tae

Tee GW

7

OF THE

CALIFORNIA ACADEM

ae
OF SCIENCES

FOURTH SERIES

Vol. XXXIX, No. 15, pp. 273-284; 3 figs. July 9, 1973

CHEMICAL EXTRACTION TECHNIQUES TO
FREE FOSSIL SILICOFLAGELLATES FROM
MARINE SEDIMENTARY ROCKS

By
York T. Mandra

Research Associate, California Academy of Sciences and
Professor of Geology, California State University, San Francisco

A. L. Brigger

Research Associate, California Academy of Sciences
and

Highoohi Mandra

8 Bucareli Drive, San Francisco, California

Abstract. Techniques of extracting fossil silicoflagellates and other siliceous micro-
fossils from marine sedimentary rocks are described. The extraction is achieved by
dissolving and chemically disaggregating all the rock (or the cementing agents of
the rocks) except the siliceous fraction, and by removing the nonfossiliferous fraction
so that the fossils will not be hidden.

INTRODUCTION

This paper, while written primarily for micropaleontologists who work with
silicoflagellates, can also be used by those who study other siliceous micro-
fossils such as diatoms, radiolarians, ebriata, sponge spicules, and _ rocellids.

It is our intent that this paper be a useful working tool for micropaleontologists
who are not either chemists or geochemists. It is not intended to be a mono-
graphic study, recording, comparing, and evaluating other techniques. Hence

1 Silicoflagellates are defined here as marine, planktonic Mastigophora (Protozoa) with a pseudopodium,
a flagellum, and a skeleton of hollow siliceous rods. These organisms also contain color pigment organelles and

therefore are treated by some workers as plants (Algae) and by others as an animal-plant group (Protista).

[273]

274 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

no long list of prior papers dealing with this subject is cited. Papers omitted
from our much abridged list have not been judged less worthy by us. Their
omission results simply from our wish to present a short list. The reader who
wishes to compare other techniques is referred to the following papers and their
bibliographies: Mann (1922), Hustedt (1958), Hanna (1937), Burma (1965),
Gray (1965), and Schopf (1965). The first four workers have techniques that
are similar to each other, and similar to our techniques. However, there are
sufficient differences between our methodology and the earlier procedures to
justify making our data available to those who work with siliceous microfossils.
Gray and Schopf deal with pollen, but their papers include some techniques
and procedures that are transferable to the study of siliceous microfossils.

Rocks that contain fossil silicoflagellates vary from friable (e.g. pure di-
atomite) to well indurated (e.g. limestone). The techniques presented here
can be used for all marine sedimentary rocks except those which have essentially
the same chemical composition as silicoflagellates. Examples of rocks for which
these techniques will not work are chert and jasper. Siliceous microfossils in
these kinds of rocks must be studied by thin sections and polished surfaces.
The reader is referred to any standard petrographic text for these techniques.
They are not included here because it has been our experience that the study of
silicoflagellates and diatoms by thin sections and polished surfaces is at best
not satisfactory and, in most cases, not worth the effort.

OBJECTIVES

The purpose of this paper is to describe our techniques of extracting fossil
silicoflagellates and other siliceous microfossils from marine sedimentary rocks.
If the procedures given here are followed, all individual specimens of these micro-
organisms will be relatively clean (i.e., no large particle of debris will be cemented
to the specimens).

These results are achieved by: (1) dissolving and chemically disaggregating
all the rock (or all the cementing agent of the rock) except the siliceous fraction
of the rock and the siliceous microfossils in the rock; and (2) decanting away
the nonfossiliferous part of the rock so that silicoflagellates will not be hidden
by these materials. In general only acids and other chemicals that do not attack
siliceous microfossils are used, the one exception being the use of sodium hy-
droxide (NaOH), which can dissolve silicoflagellates. However we use it as a
dilute solution (0.25—1.0 N) and only for a relatively short time. No appreciable
damage to silicoflagellates or diatoms by such use of sodium hydroxide has
been observed.

DEEP-SEA CORE SAMPLES AND OUTCROP SAMPLES. There is such a wide vari-
ation in the lithology, mineralogy, and chemistry of marine sedimentary rocks
containing fossil silicoflagellates that no one simple chemical extraction (or

Vor. XXXIX] MANDRA, ET AL.: EXTRACTION TECHNIQUES 275

simple generalized method) will be applicable to all marine sedimentary rocks.
However, one generalization is possible: most deep-sea core samples are more eas-
ily prepared chemically, as compared with most older Cenozoic samples taken from
outcrops of well indurated rock. Therefore many steps of the procedure pre-
sented here may not be necessary for deep-sea core samples. In contrast, a
large number of important New Zealand Early Tertiary samples taken from
outcrops consisting of well indurated rock and cemented by complex compounds,
have taken as long as two weeks of continuous work per sample to prepare.

NUMBER OF SAMPLES TO BE PROCESSED AT ONE TIME. If the worker has ade-
quate experience and adequate equipment, and no problems develop, then six
samples can be processed as a batch—at one time. However, at the sulfuric
acid fuming? step only one sample at a time can be worked. Frequently we
find that three or four samples are all that the most experienced worker can
handle at one time. Again, this excludes the sulfuric acid fuming step, when
but one sample should be handled. Furthermore it is not uncommon that one
sample could be such a continous source of trouble that it should be processed
alone. Beginners, for reasons of safety and efficiency, should work with only
one sample at a time.

DIRECTIONS AND COMMENTS
Read each set of directions completely before you start to do each step.

Rubber gloves, plastic apron, face mask, and goggles should be used at all times
while handling acids or sodium hydroxide solutions or hydrogen peroxide so-
lutions. Use a fume hood when working with beakers containing acids.

if Place the rock sample which is to be prepared on a layer of newspapers.
With a clean ice pick break a piece from each large fragment of the rock
sample so that the material to be treated will be representative of the whole
sample. The remaining untreated portion of the rock sample should be
preserved for future study. Reduce all the pieces to about one-fourth inch
in size with the ice pick. The maximum amount of the sample to be treated
should be about 100 grams.

After each sampling has been made, carefully roll up and discard the
newspapers, and clean the ice pick. This procedure will prevent contam-
ination of one sample by another.

2, Label a clean 1000-ml. Pyrex beaker with sample number. Place about
100 grams of the one-fourth inch pieces into the beaker. If the samples are
accurately weighed, the abundance of silicoflagellates in the fossil plankton
can be quantitatively determined. (Tappan et al., 1971)

2In this paper we use the term “sulfuric acid fuming” in the sense of concentrated H,SO, which is
boiling and decomposing to H,O and SO, white fumes. We do not mean “fuming sulfuric acid” which
is H,S,O, (H,SO, with SO, in solution).

bo

CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Cover the sample in the 1000-ml. beaker with about 300 ml. of distilled
water. Large beakers are needed because some chemical reactions can be
quite vigorous.

If a worker is familiar enough with the microorganisms in his source of
tap water to recognize them as contaminants, if the mineral content of the
tap water is relatively low, and if the expense of distilled water is a factor
to be considered, then good tap water filtered at the tap could be used in
many of the steps described in this paper.

Place a heavy watch glass over the beaker and add concentrated, chem-
ically pure hydrochloric acid (HCl) through the lips in increments of 10 ml.
if effervescence is evident. If there is no effervescence, slowly add all 100
ml. at one time. However, care must be taken because if the sample has a
high calcium carbonate (CaCO;) content and if a large amount of hydro-
chloric acid is added at once, excessive effervescence might result in a
spill-over.

Calcium carbonate will neutralize the acid, hence more HCl may be
needed to insure an excess of acid after all the CaCOs has reacted.

If chemically pure acids are not available to the worker and if expense
is a factor in the operation, then technical quality acids could be used.
However, because the chemically pure acids have smaller amounts of im-
purities their use is recommended.

The purpose of the HCl treatment is to dissolve oxides and salts (other
than silicates) of metals whose chlorides are soluble. These soluble chlo-
rides can then be decanted away. This treatment will work for most metals
except silver and lead in the marine sedimentary rock sample.

Place the beaker on a hot plate and keep the solution at a gentle boil
until all reactions have ended. Suggested hot plate dimensions are: top,
three-eighths inch thick, good quality stainless steel, 20 X 20 inches; 10-
inch-high stainless steel legs; and 1'2-inch-high removable stainless steel
railing to prevent beakers from falling off the edge. Heat should be sup-
plied by two or three Meker burners. Rubber tubing should not be used
because of the high heat. Tygon or other heat resistant tubing should be
used, or, better yet, a direct connection with steel piping with valve control.

Add distilled water to the beaker until half-filled, set aside until no
microfossils are suspended in the solution. This should be determined by
examination of a drop of the solution under the microscope at 100 X.

Decant carefully so as not to lose any of the microfossils. Fill the
beaker half full with distilled water. Cover with a watch glass and again
bring the solution to a gentle boil. More soluble chlorides will now be
taken into solution as will be indicated by the color of the water. Boil gently
for at least an hour, cool, and examine a drop of the solution under a micro-

Vor. XXXIX] MANDRA, ET AL.: EXTRACTION TECHNIQUES 277

®

uae %

Ficure 1. Sample broken down with HCl and water, retained after washing on 400-
mesh screen.

scope at 100 X. If there are no microfossils in the solution, decant. If
microfossils are still in suspension, wait until they settle, then decant.

F Add distilled water to the beaker until half-filled. Remove the fine
sediment by vigorously swirling the solution in the beaker so that the fine
fraction (i.e. the decomposed and disaggregated material of the rock
sample) goes into suspension. Then immediately pour off the solution with
its suspended matter into another clean 2000-ml. Pyrex beaker marked with
the sample number and an ‘A’, leaving the coarser material in the original
beaker.

Repeat step 5 several times until the solution in the original beaker is
neutral to litmus paper.

6. To the original beaker containing the hydrochloric acid-treated sample
that is not yet decomposed and not yet disaggregated, add a solution of
sodium carbonate NazCO; (50 grams per liter of distilled water) until
the sample is covered with the solution.

Heat slowly on the hot plate and gradually add a 30 percent solution
of chemically pure hydrogen peroxide (H2O2). This must be done care-
fully, a few drops at a time, because the reaction could be quite vigorous
and some of the sample could be lost. Continue this careful addition of

CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

FicurE 2. Sample fumed with H2SO;:, washed and retained on 400-mesh screen.

hydrogen peroxide until 10 ml. have been added. Boil very slowly for
30 minutes. Then remove the beaker from the hot plate to cool and allow
the sediment to settle.

The boiling with hydrogen peroxide helps to disaggregate the particles
in the sample and also to oxidize the organic matter. The organic matter
in the sample probably is the residue of plankton, seaweed, etc., partially
decomposed and combined with other materials desiccated and oxidized
while at outcrop. If the sample has not been exposed to the atmosphere
(e.g. a deep-sea core sample), it will not be oxidized and desiccated; the
organic matter will usually go into solution easily and chemical extraction
of microfossils is simple. This is why we stated earlier that most deep-sea
cores are relatively easy to prepare.

If there are no microfossils in suspension, decant and discard the solution
which is in beaker ‘A’. Test for acidity with litmus paper. If acidic, wash
several times and decant until the solution is neutral. Examine a drop of
the solution with the microscope at 100 X before each decantation to be
sure that no microfossils will be lost.

Decant the sodium carbonate - hydrogen peroxide treated fine sediment
of step 6 into beaker ‘A’. Wash several times by swirling the original beaker
until the fine disaggregated and decomposed fraction of the sample is in
suspension. Continue washing and decanting until beaker ‘A’ is nearly filled.

Vor. XXXIX] MANDRA, ET AL.: EXTRACTION TECHNIQUES

ins)
~
Ko)

FicurE 3. Fumed sample treated with NaOH and H.O., washed and retained on 400-
mesh screen.

Then set beaker ‘A’ aside for the suspension to settle. Decant when the
solution above the precipitate in beaker ‘A’ is clear.

Place the original beaker which contains the undisaggregated material
back on the hot plate and add about 400 ml. of distilled water. Bring to
boil and keep at a low boil for about three-quarters of an hour.

9. In most cases the sodium carbonate and hydrogen peroxide boiling at
this stage should have decomposed or disaggregated most of the remaining
cementing agents which have held the rock sample together. The boiling
water in step 8 should have completely disaggregated the sample.

However, there are samples that will not react, or will barely react, to
this treatment. In such a case, after decanting the sodium carbonate solution
into beaker ‘A’ cover the remaining undecomposed and undisaggregated sam-
ple in the original beaker with a solution of sodium hydroxide (20 grams
NaOH to 1000 ml. of distilled water) and carefully add 1 ml. of 30 percent
chemically pure hydrogen peroxide (H2O2). Bring the solution to a boil
and continue the boiling for five minutes. Remove from the hot plate, cool
and decant the solution and fine sediment into beaker ‘A’. Add 400 ml.

10.

dell.

CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

of distilled water to the remaining residue in the original beaker and boil
for 30 minutes at which time the sample should be completely disaggregated.
Remove from the hot plate, cool, swirl the solution, and decant the sus-
pended matter and solution into beaker ‘A’. If at this stage undecomposed
material still exists in the original beaker, dry it overnight in an oven at
100° C. Label it ‘Partially Treated’ and store it in a container that also
clearly records its locality data.

All the fine material that has been separated from the larger pieces
of the marine sedimentary rock sample is now in beaker ‘A’. Fill this
beaker with distilled water and allow the solution to settle. Examine a drop
of the solution under the microscope at 100 X. If there are no microfossils
in suspension, decant and discard the solution.

If in the acid treatment all of the metal ions have not been removed,
they will now exist as reprecipitated carbonates or hydrates. To remove
them, add 100 ml. of concentrated chemically pure hydrochloric acid (HCl)
and 400 ml. of distilled water, cover with a watch glass and boil for one
hour or more. Remove beaker ‘A’ from the hot plate, allow it to cool and
allow the sediment to settle. Examine the suspended matter in the solution.
If the microfossils have settled, decant and discard the solution. Continue
to wash the sediment by repeated decantations until the solution is neutral,
making sure that no microfossils are poured away.

The purpose of this step is to reduce the bulk of the sample. Use a
clean 2-inch-deep, 12-inch-diameter, 400-mesh all stainless steel screen, and
a clean 10-liter bucket to retain the screened material. Before screening
the material, wet the screen on both sides with water; if this is not done
water will not pass through this fine mesh.

Add water to beaker ‘A’ until about half full, swirl the solution in the
beaker until the fine sediments are in suspension. Allow the heavier par-
ticles to settle: a minute should be sufficient time. Then pour onto the
screen a little of the suspended material while agitating the screen by gently
striking it on the side with the palm of the hand. To aid the screening,
use a plastic wash bottle so held that water will flow onto the screen. The
plastic wash bottle should have plastic tubing in order to protect the ex-
pensive screen against accidental contact with the tubing. Continue agitating
and washing the material on the screen until the water passing through
carries no more sediment. Do not use your fingers or any tool to force
material through the screen as this will damage both screen and sample.
Tilt the screen slightly and wash the retained material to the edge of the
screen. With the wash bottle, carefully wash the retained material on
the screen into the beaker to be used for the sulfuric acid fuming step. The
beaker size should be either 400 or 250-ml. depending upon the amount of

Vor. XXXIX] MANDRA, ET AL.: EXTRACTION TECHNIQUES 281

12:

13:

residue remaining on the 400-mesh screen.* Continue this screening process
until all the sample that is being treated has been washed on the 400-mesh
screen and transferred to either the 250 or 400-ml. beaker.

Because the screen openings of the 400-mesh screen are 38 microns in
size, some small silicoflagellates and other small microfossils will pass
through these screen openings. These microscopic fossils should be retained
for study. However, it has been our experience that the —400 mesh fraction
frequently does not contain many silicoflagellates. If there are no micro-
fossils in suspension, decant the solution from the 10-liter bucket, dry its
sediment overnight in an oven at 100° C., label, and store for examination
and study.

The +400 fraction which does not go through the screen will have un-
decomposed fragments of the sample containing fossils that are smaller than
38 microns. However, after completing all the steps these remaining speci-
mens, smaller than 38 microns in size, will be freed and available for study.

Clean the screen after use with a surfactant solution. If possible use an
ultrasonic cleaning device.

In this dangerous step the sample is boiled in sulfuric acid in order to
dissolve organic cementing agents.

After the microfossils have settled to the bottom of the 250 or 400-ml.
beaker, decant the water. If the layer of the sediment is more than ™%-inch
thick, use a 400-ml. beaker.

Extreme caution should be used while the sample is subjected to the
sulfuric acid treatment and a face mask and eye goggles should be worn.
Place a heavy watch glass over the Pyrex beaker and with the aid of a
pipette to which a rubber bulb is attached, slowly add small increments of
chemically pure concentrated sulfuric acid through the lip of the beaker.
These small amounts of acid should be run down the side of the beaker
rather than added directly to the contents of the beaker. Agitate the beaker
occasionally as the acid is being added in order to avoid any vigorous action.
Continue the addition of sulfuric acid slowly until at least 150 ml. has been
added, and the sample is covered with at least one inch of sulfuric acid.
Rotate the beaker in an arc with the aid of beaker forceps to mix the acid
with the contents of the beaker.

Place the beaker on a ring stand over a Nichrome wire gauze and adjust
the Meker burner so that it is about two inches below the wire gauze. This
system is necessary because the previously used hot plate will not produce
temperatures sufficiently high for this step.

The beaker lip should be kept pointed away from the worker.

3 The material that is retained on the 400-mesh screen is called the +400 fraction, and is read as ‘plus’

400. Similarly, that which passes through the 400-mesh is called the —400 fraction, and is read as ‘minus’ 400.

14.

15.

16.

CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

If the contents start to bump, hold the beaker with the beaker forceps
and swirl the solution with its sediment. This should minimize the bumping.
If the solution becomes viscous, remove from heat and slowly and carefully
add more sulfuric acid. Then continue heating. If the solution should start
to foam excessively, slide the watch glass partially off the top of the beaker
so that the solution is exposed to the air and remove from heat. As soon
as the foaming has subsided, heat again. This tendency to foam generally
lasts only a short time. Never add water to prevent foaming, because a
violent reaction would take place. After white fumes appear, continue
heating for about ten minutes. Then put the solution aside for about two
hours until the beaker has cooled to room temperature.

With the aid of a pipette to which a rubber bulb is attached, slowly and
carefully add 10 ml. of concentrated chemically pure nitric acid (HNOs)
to the fumed sample through the lip of the beaker which is covered with a
watch glass, and again bring the solution to fumes over the Meker burner.
The combination of the two acids will oxidize the sample and change the
color of the sediment to white. Set this solution aside to cool to room tem-
perature. Through the lip of the beaker which is covered with a watch
glass carefully add small amounts of distilled water down the side of the
beaker, a few drops at a time. Swirl the solution with beaker forceps after
each addition of water until there is no reaction.

Fill the beaker three-fourths full with water and allow to settle until
the solution above the sediment is clear. At this point do not examine the
solution under a microscope because of the high acidity.

Decant carefully and discard the solution making sure that none of the
sediment is lost. Fill the beaker three-fourths full with distilled water.
Swirl the solution in the beaker as water is added to insure that the sample
is being washed. Allow the sediment to settle and examine it under a micro-
scope, as in earlier washings, for the presence of microfossils. Continue the
washings, examinations, and decantations until the solution gives a neutral
reaction to litmus paper.

Add 100 ml. of approximately 1.0 N solution of sodium hydroxide* (40
grams of sodium hydroxide per liter of distilled water) and two ml. of a
30 percent solution of hydrogen peroxide to the solution. Place the beaker
on a ring stand over gauze and adjust the flame of the Meker burner to
give a low heat.

Continue to use extreme care. Gently rotate the beaker holding it with
beaker forceps in such a way that the heat of the flame contacts the outer

4 NaOH will dissolve siliceous microfossils. Therefore a test was made in order to determine within what

limits sodium hydroxide can be used. A solution containing 20 grams of sodium hydroxide per liter of water
was placed in a beaker containing a sample of fossil silicoflagellates and diatoms and heated to dryness.
Water was then added and the microfossils examined. There was no apparent damage to the fossils.

Vot. XXXIX] MANDRA, ET AL.: EXTRACTION TECHNIQUES 283

We

18.

edges of the beaker. Care should be taken to prevent the flame from coming
in contact with the interface between the solution and air, for even a Pyrex
beaker might then crack. As soon as the solution becomes warm the H2O2
will start decomposing and bubbles of oxygen will become attached to
particles of sediment in the beaker, eliminating to a great extent the hazard
of bumping. Without the addition of hydrogen peroxide, bumping could be
severe enough to cause loss of most of the contents of the beaker. Allow
the solution to boil for two minutes. Remove from heat and allow the sedi-
ment to settle so that the solution above the sediment does not contain any
microfossils in suspension.

The purpose of this sodium hydroxide treatment is to decompose the
siliceous fraction of the sample that is cementing the sediment together and
to decompose those siliceous particles that are adhering to the microfossils.

Decant the solution and add 20 ml. of HCl. Cover the beaker with a
watch glass and bring the acidified solution to a boil on the hot plate and
continue at a gentle boil for about 30 minutes. Remove from the hot plate
and cool. Decant after the sediment and microfossils have settled. This last
decantation may be used to determine qualitatively what metallic ions are
left in the sample.

Add distilled water to the beaker, allow microfossils to settle, then decant,
using the prior described precautions of not decanting any of the sample.
Test for acidity. If acidic, wash again by decantation until the solution is
neutral to litmus paper. The sample should now be completely clean.

Nest three stainless steel screens 150, 250, and 400-mesh. This will
yield four fractions: +150, —150 +250, —250 +400, and —-400-mesh. Such
a division of the treated sample makes it possible to study specimens more
effectively as well as to make better slides, because smaller specimens will
not be obscured by larger ones. Pour the sample onto the top of the 150-
mesh screen of the nested screens, which in turn are placed on top of a
2000-ml. Pyrex beaker. The lip of the beaker will serve as an air vent.
Wash each screen well with a wash bottle until little or no sediment passes
through the screen. When no more sediment will pass through the 150-mesh
screen, tilt it and carefully wash the sample with the wash bottle into a
beaker marked with the sample locality description and ‘+150-mesh’.

As the screening proceeds, check to see that the volume of water is not
increased in the 400 and 250-mesh screens to a point of overfill. If this
occurs, place the top two screens onto another clean, labeled 2000-ml. beaker
and agitate the 400-mesh screen by striking it gently with the palm of the
hand until it no longer contains water. Then place the 250-mesh screen on
top of the 400-mesh screen and continue the process until the sample has
been thoroughly washed and all the sediment transferred to properly marked
beakers with distilled water.

284 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Prepare the bottles in which the four samples will be kept by washing the
bottles with distilled water, and attaching labels giving the locality data
and the appropriate mesh number data (e.g. +150, -150 +250, —250 +400,
and —400-mesh).

Allow the fraction of the sample which has passed through the 400-mesh
screen to settle for at least two hours. Examine for microfossils in suspen-
sion. If fossils are in a drop of the solution, wait until they have settled.
Then decant and carefully transfer this fraction to its bottle. Preserve each
sample with the addition of four drops of 37 percent solution of formalde-
hyde in bottles of approximately 30 cc. capacity. Fill bottles about three-
fourths full with distilled water. Use leak-proof caps.

ACKNOWLEDGMENTS

We wish to thank Dr. Peter F. Linde, Professor of Chemistry, California
State University, San Francisco, and Dr. Milka Zhivadinovich, Professor of
Chemistry, California State University, Hayward for their critical reviews of
our manuscript.

REFERENCES CITED

Burma, B. H.
1965. Radiolarians, i7 Handbook of Paleontological Techniques. Edited by Bernhard
Kummel and David Raup, pp. 7-14. Freeman Co., San Francisco.
Gray, JANE
1965. Palynological Techniques, 7x Handbook of Paleontological Techniques. Edited
by Bernhard Kummel and David Raup, pp. 471-481. Freeman Co., San
Francisco.
Hanna, G D.
1937. Invertebrates, 7x Methods in Paleontology. By Camp, C. L., and G D. Hanna,
pp. 117-121. University of California Press, Berkeley.
HusteptT, FRIEDRICH
1958. Praparation und Untersuchungsmethoden fossiler Diatomeen, 7x Handbuch der
Mikroskopie in der Technik, vol. 2, pt. 3, pp. 427-450.
Mann, ALBERT
1922. Suggestions for collecting and preparing diatoms. Proceedings of United States
National Museum, 60, pp. 1-8.
ScHOPF, JAMES
1965. A method for obtaining small acid-resistant fossils from ordinary solution residues,
in Handbook of Paleontological Techniques. Edited by Bernhard Kummel
David Raup, pp. 301-304. Freeman Co., San Francisco.
Tappan, HELEN, WILLIAM C. CORNELL, AND E. REED WICANDER
1971. Quantification of fossil phytoplankton fluctuations. Abstract of Research, Fif-
teenth Annual Report on Research, Period Ending August 31, 1970, American
Chemical Society, Petroleum Research Fund, pp. 195-196.

ariné Biological Laboratory
LIBRARY
AUG 2 11973

Weéds Hala; Mass.

PROCEEDINGS

OF THE

CALIFORNIA ACADEMY OF SCIENCES

FOURTH SERIES

Vol. XX XIX, No. 16, pp. 285-310; 10 figs.; 1 table. August 9, 1973

TWO NEW SPECIES OF THE SCORPIONFISH
GENUS RHINOPIAS, WITH COMMENTS ON
RELATED GENERA AND SPECIES

By
William N. Eschmeyer

California Academy of Sciences, Golden Gate Park, San Francisco,
California, U.S.A. 94118

Yoshitsugu Hirosaki
Enoshima Aquarium, Fujisawa City, Japan 251
and
Tokiharu Abe

Tokai Regional Fisheries Research Laboratory,
Ministry of Agriculture and Forestry, Katchidoki 5-5-1, Kyobashi P.O.,
Tokyo 104, and Zoological Institute, Faculty of Science,
University of Tokyo, Hongo, Tokyo 113, Japan

Apstracr. A new scorpionfish, Rhinopias argoliba, is described from Japan. A new
species name, Rhinopias aphanes, is given for a specimen identified as R. frondosa
(Giinther) by Whitley (1964), and this single known specimen from New Caledonia
is described and figured. Additional specimens of Rhinopias xenops (Gilbert) and
Rhinopias frondosa are described. Rhinopias godfreyi Whitley is referred to the genus
Pteroidichthys. The genus Pteroidichthys Bleeker is thought to be closely related to
Rhinopias Gill. The monotypic genus Pogonoscorpius Regan, known only from the
holotype of P. sechellensis Regan, possibly is a synonym of the genus Rhinopias.
Remarks are given on the monotypic genera Hipposcorpaena and Pteropelor, both
described by Fowler (1938) from juvenile specimens. Comments on shedding of ‘skin’
in these and other scorpaenid fishes are given.

INTRODUCTION

Specimens of the genus Rhinopias are very rare and are among the most
striking of fishes in coloration and form. Species are known from the Indian

[285]

286 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Ocean and the central and western Pacific Ocean. The two new species de-
scribed here bring to four the number of species in the genus. The few speci-
mens known previously have received considerable attention (Palmer, 1963;
Smith, 1966; and others). We include with the descriptions of the new species
information on the other two species, including new records, and offer comments
on related genera and species.

The Japanese specimen on which one new species description is based al-
ready has had an interesting history. It was collected by Mr. Hajime Masuda
and maintained alive in the Enoshima Aquarium, Fujisawa, Japan. This speci-
men was depicted in color on the cover of Fish Magazine (1971) and it was the
subject of articles on shedding of skin by one of us (Hirosaki, 197la-c). The
second new species is based on a specimen which was first displayed in the
Nouméa Aquarium in New Caledonia and was figured by Whitley (1964) as
Rhinopias frondosa (Giinther).

Shedding of ‘skin’ in scorpaenid fishes has received some attention, most
recently by Hirosaki (1971a-c) and Wickler and Nowak (1969). Information
is provided on additional genera of scorpaenid fishes observed to shed.

ACKNOWLEDGMENTS

We wish to express our sincere thanks to Mr. Hajime Masuda, Izu Ocean
Park, It6 City, Japan, for his generosity and thoughtfulness. He provided the
first and only known specimen of Rhinopias argoliba, allowing it first to be
displayed alive in the Enoshima Aquarium, and second to be studied by us.
We gratefully acknowledge the valuable services rendered by Dr. Chiichi Araga,
Seto Marine Biological Laboratory (SMBL), Kyoto University, and Dr. Osamu
Okamura, Kochi University (KU); they provided information and sent on
loan Japanese specimens and color slides. Similarly, we are grateful to Mr. Paul
J. Struhsaker, National Marine Fisheries Service, Honolulu, Hawaii, for provid-
ing an unreported specimen of Rkinopias xenops from Hawaii and two speci-
mens of R. frondosa from the Caroline Islands. We are grateful to Dr. R. Catala,
Nouméa Aquarium, New Caledonia, for providing a color slide, photographs,
and additional information on the holotype of R. aphanes.

Funds from a United States National Science Foundation grant (NSF 15811)
permitted the first author to visit museums. The following persons assisted
the present study during visits: John R. Paxton, the Australian Museum, Sydney
(AMS); M.L. Bauchot and Jean Claude Hureau, Museum National d’Histoire
Naturelle, Paris (MNHN); Alwyne C. Wheeler, British Museum of Natural
History, London (BMNH); Jgrgen Nielson, Zoologisk Museum, Copenhagen
(ZMC); M. Boeseman, Rijksmuseum Van Natuurlijke, Leiden (RMNH); Adolf
Kotthaus, Biologische Anstalt Helgoland, Hamburg (BAH); and the staff of
the U. S. National Museum of Natural History, Washington, D. C. (USNM).

Vor. XXXIX] ESCHMEYVER ET AL.: NEW SCORPIONFISHES 287

M. L. Bauchot, Jorgen Nielson, and John R. Paxton loaned specimens. Alwyne
Wheeler provided additional information for some specimens. Richard Krejsa,
California State Polytechnic College, analyzed shed ‘skin’ from scorpionfishes
for us. Pearl Sonoda, Tomio Iwamoto, Maury Giles, Kathy Smith, Kathryn
Boyer, and other staff members of the California Academy of Sciences (CAS)
aided the study.

We wish to thank Lillian J. Dempster, CAS, for her help with literature, for
her comments on the manuscript, and for assisting in the selection of new spe-
cies names. K. V. Rama Rao provided comments on the manuscript.

METHODS

Methods of measuring and counting follow Eschmeyer (1969); exceptions
to measurements commonly used to describe teleosts include the following:
pectoral-fin length is measured from the base of the first ray to the end of the
longest ray with the fin pointing back; measurements originating from the ante-
rior end of the fish are taken from the most anterior point of the left pre-
maxillary, and measurements are made on the left side of specimens. The last
soft ray of the dorsal fin and anal fin is a double ray and the expression XII +
9% is used to signify 12 spines and 9 soft rays (last double). Terminology of
head spines follows Matsubara (1943) and Eschmeyer (1969). Abbreviations
for depositories of specimens are given in the Acknowledgments section except for
the following: EA—Enoshima Aquarium, Fujisawa.

Genus Rhinopias Gill
Rhinopias GILL, 1905, p. 225 (type-species Scorpaena frondosa GUNTHER, 1891, by original
designation; monotypic).
Peloropsis GILBERT, 1905, p. 630 (type-species Peloropsis xenops GILBERT, 1905, by original
designation; monotypic).

Remarks. The genus Rhinopias has about six months priority over the
genus Peloropsis. The monotypic genus Pogonoscorpius Regan (1908), with
the type-species P. sechellensis, may also be a synonym of Rhinopias. Pogono-
scorpius sechellensis is still known only from the holotype (see our treatment
of Pogonoscorpius). Rhinopias godfreyi Whitley (1954) is referred to the genus
Pteroidichthys Bleeker.

Diacnosis. Dorsal fin XII + 9%; anal fin II] + 5%; pectoral rays 15-18,
some rays branched in larger specimens; scales on body small and cycloid, over
60 vertical scale rows; head unscaled; vomerine teeth present; palatine teeth
absent; vertebrae 24; body compressed; second suborbital bone becoming wider
posteriorly, attached to preopercle; third suborbital bone absent; no slit behind
fourth gill arch.

SPINATION. Head spination is similar for all species. Most spines poorly
developed. Nasal spines small, usually absent in one species. Preocular, supra-

288 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

ocular, and postocular spines present; sometimes preocular and supraocular
spines reduced; supraocular spine obscured by tentacle; postocular spine as
broad shelf above rear of orbit. Parietal and nuchal spines well developed,
joined. Sphenotic spine (as group of small spines) absent in some species.
Pterotic spine well developed. Upper posttemporal spine small; lower post-
temporal spine well developed. Supracleithral and cleithral spines rounded.
Opercular spines blunt at tips. Tympanic, coronal, and interorbital spines absent.
Supplemental preopercular spine present or absent. First three preopercular
spines moderate, fourth small, fifth absent or barely perceptible. Suborbital
ridge with four low spines or lumps, first on lateral face of preorbital bone,
followed by one under front of eye, and two just before preopercular bone;
sometimes these spines may be absent or poorly developed. Preorbital bone with
2 poorly defined spines over maxillary (second and third preorbital lobes of
Matsubara, 1943).

Key TO THE SPECIES OF THE GENUS Rhino pias

1. Pectoral rays 18 [17 and 19 should be expected]; no black spot or ocellus on soft

@orsal “fin: 22 ee ee 2
1. Pectoral rays 16 or 15 [17 should be expected]; black spot or ocellus present on soft

Gorsalee tiny. eee ee ee meee ee 3
2. Skin flaps absent on lower jaw; few skin flaps on body; third dorsal spine about 3

times in head length and about two-thirds of snout Rhinopias argoliba (figs. 1-2A)

2. Skin flaps present on lower jaw; head, body, and fins with numerous skin flaps; third
dorsal spine about two times in length of head or less than 2 times if third spine is

especially elongate, third spine longer than snout Rhinopias xenops (figs. 2B, 3-4)
3. Body, head, and fins with round and oblong pale (dark rimmed) spots or blotches;
sometimes almost entirely pallid in preservative — Rhinopias frondosa (figs. 5-7)

3. Body, head, and fins with dark reticulations on a paler background —._________
paloek EDS Sets ee Repose,

SPECIES ACCOUNTS

Rhinopias argoliba Eschmeyer, Hirosaki, and Abe, new species.

(Figures 1-2A.)

No scientific name used, HrrosakI, 1971a, pp. 4-5, 2 figs. (shedding); 1971b, pp. 26-27,
front cover, 5 figs. (shedding; cover in color showing live coloration); 1971c, p. 170,
fig. (shedding).

Inimicus didactylum (not of Pallus), AxELRop and Burcess, 1972, figs. 466, 470 (misiden-
tification; poor color reproduction of specimen when alive in the Enoshima Aquarium).

Hototyrr. Enoshima Aquarium no. 1 (129 mm. S.L., 167 mm. T.L.),
Japan, Sagami Bay, caught off Izu Ocean Park, Ito City, at a depth of 50
meters, 15 January 1971 [transferred alive to Enoshima Aquarium on 30 Jan-
uary 1971, where it died on 23 May 1971].

Description. (Based on the holotype and only known specimen.) A large-
headed, compressed scorpionfish. Dorsal fin with 12 spines and 9% soft rays;

VoL. XX XIX] ESCHMEYER ET AL.: NEW SCORPIONFISHES 289

Ficure 1. Rhinopias argoliba, holotype, EA no. 1, 129 mm. S.L., Japan; note white
coloration below eye and above pectoral fin (from the specimen when alive).

third dorsal spine not elongate, fourth spine the longest, eleventh dorsal spine
about ™ the length of the twelfth spine. Anal fin with 3 spines and 5% soft
rays; first spine slightly more than one-half length of second, third spine long-
est. Pectoral rays 18, only one or two branched rays in each fin (branching of
pectoral rays variable with size in other species). Gill rakers (including rudi-
ments) total 23, 7 on upper arch, 16 on lower arch including one at angle, all
rakers as short spiny knobs. Scales on body small and cycloid; about 80 verti-
cal scale rows (count made a few scale rows above lateral line from above first
lateral-line scale to end of hypural plate). Head without scales.

Spination similar to that of other species of Rhinopias (see account under
genus). Nasal spines small. Supplemental preopercular spine present, small,
lying above first preopercular spine. Sphenotic spines absent in available speci-
men. Suborbital ridge with 4 spines; first a sharp one on preorbital bone, fol-
lowed by one under the eye and two lumps before the preopercular bone.

290 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Ficure 2. A. Rhinopias argoliba, holotype, EA no. 1, 129 mm. S.L., Japan. B. Rhinopias
xenops, USNM 209415, 107 mm. S.L., Hawaii.

Measurements for the holotype in table 1.

Tentacles on body few. No supraocular tentacle in available specimen; no
skin flaps on lower jaw as in other species; small skin flaps on nostrils, on eye,
on lower margin of preopercle, on some lateral-line scales, and a few very tiny
ones on the pectoral fin. (The development of tentacles and skin flaps is some-
what variable in scorpaenid fishes. See the ‘Comparisons and Remarks’ section
below. )

VoL. XXXIX] ESCHMEYVER ET AL.: NEW SCORPIONFISHES 291

Coloration in preservative almost entirely pallid (fig. 2); no reticulations
present, no dark spot on soft dorsal fin. White stripe before eye slightly visible;
some faint dark areas below eye (melanophores contracted). Color in life bright
red, including all of body, head, and fins, with the exception of the white stripe
before the eye, white patches near base of lower pectoral rays, a few scattered
white spots on body and snout, one conspicuous spot about size of pupil just
above lateral line over pectoral fin, and another spot on dorsal part of caudal
peduncle. About 15—20 small (0.2—0.5 mm. in diameter) dark blue specks on
head, most concentrated below eyes just above suborbital ridge. Spinous dorsal
fin tipped with white. The white stripe (fig. 1) before the eye expanded into
a circular spot at distal end (resembling an exclamation point |!]), ‘spot’ sep-
arated from stripe on left side, partially joined to stripe on right side (see
Hirosaki, 1971c, fig. on p. 170, or Axelrod and Burgess, 1972, figs. 466, 470).

COMPARISONS AND REMARKS. This species is easily distinguished from
R. frondosa and R. aphanes by having a higher pectoral fin ray count (18 versus
16 or 15). The development of tentacles and skin flaps on the head and body
is variable but presence on the lower jaw is a stable feature in species of other
genera. Of the four species of R/inopias only R. argoliba lacks tentacles on
the lower jaw. Rhinopias argoliba appears to be closely related to R. xenops
with which it shares a pectoral fin ray count of 18. The holotype of R. xenops
has the third dorsal spine notably elongate, but the additional specimens from
Hawaii and two Japanese specimens do not, although the third spine is slightly
elongate in two. In some scorpionfishes which have one or more dorsal spines
elongate, this elongation is variable and does not occur until the fish reaches
the subadult stage or even the adult stage; we do not know if this difference will
be useful for distinguishing large specimens. The dorsal spines are proportion-
ally shorter in the specimens of R. argoliba than in R. xenops. The second anal
spine is shorter than the third in R. argoliba, but about equal in length to the
third in R. xenops. Most fin spines are proportionately shorter in R. argoliba
than in R. xenops. The pectoral fin in R. argoliba is shorter than in R. xenops.
The holotype of R. argoliba is as large or larger than the known specimens of
R. xenops but has fewer branched pectoral rays (2 or 3 versus 5 to 7). Branch-
ing of pectoral rays in scorpaenids begins in juveniles and increases with increase
in size to a maximum number of branched rays. The dentition on the premax-
illaries is stronger in R. xenops than in R. argoliba. Differences in coloration
of R. argoliba and R. xenops may be useful in distinguishing these species. The
entire specimen of R. argoliba was bright red in life, including the head, body,
and fins, with the exception of milky white spots on the body and the white
stripe before the eye as given in the description above. The coloration in R.
xeno ps is quite different (see account of R. xenops).

We do not know what affect, if any, the 114-day stay in the Enoshima Aquar-

292 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

ium had on the growth of the holotype of R. argoliba as might be reflected in
unnatural lengths of some body parts. The only damage to the specimen seems
to be that a callus formed at the symphysis of the lower jaw, caused by friction
against glass.

DisTRIBUTION. Known only from the holotype collected from Japan, off Its,
Sagami Bay, at a depth of 50 meters.

Name. The scientific name ‘argoliba’ (treated as an adjective) is from
Greek, argos (white) plus lébos (tear or drop), alluding to the milky-white
teardrop below the eye. The Japanese common name Namida-kasago is pro-
posed and is based on “namida,” or tear, plus “kasago,” the Japanese name
for Sebastiscus marmoratus.

Rhinopias xenops (Gilbert).

(Figures 2B, 3-4.)

Peloropsis xenops GILBERT, 1905, pp. 630-631, text fig. 245 (original description; type locality
Hawaii, Auau Channel, between Maui and Lanai Islands, depth 32 to 43 fathoms,
Albatross station 3872, 12 April 1902; holotype USNM 51604); JorpAN and SEALE, 1906,
p. 379 (listed; Hawaii) ; JorDAN and EvERMANN, 1926, p. 10 (listed; Hawaii) ; Fow er,
1928, p. 287 (compiled description) ; 1938, p. 290 (listed; Hawaii); Tinker, 1944, pp.
262-264, fig. (compiled; figure from Gilbert, 1905); Gostinre and Brock, 1960, pp. 284,
286, 341 (in key; compiled); PALMER, 1963, pp. 701-704, pl. XX (in part; wrongly
included Japanese specimens [Kamohara, 1942, et seq.]; compared with R. frondosa) ;
GOSLINE, 1965, p. 825 (depth distribution; compiled).

Rhinopias xenops, WHITLEY, 1954, p. 61 (placed in genus Rhinopias; distinguished from
other species) ; SMITH, 1966, pp. 74, 77-79 (in key, distinguished from R. frondosa).

Peloropsis frondosus, KAMOHARA, 1959, pp. 5-6 (in part; R. xenops in synonymy; thought
R. xenops may be variant of R. frondosus rather than a distinct species).

MatTertaL. USNM no. 51604 (1, 110 mm. S.L., holotype), Hawaii, Auau
Channel, between Maui and Lanai Islands, depth 32 to 43 fathoms, Albatross
station 3872, 12 April 1902. USNM no. 209415 (1, 107 mm. S.L.), Hawaii,
Haleiwa, 21°39.6’ to 21°42’N., 158°07.3’ to 158°05’W., 41-ft. shrimp trawl,
in 95-110 meters, Townsend Cromwell cruise 36, station 19, 3 May 1968.
SMBL no. 7201 (1, 114 mm. S.L.), Japan, off Shirahama Town, Nishimuro
County, Wakayama Prefecture, rocky bottom, 31 October 1971. SMBL no.
7202 (1, 129 mm. S.L.), Japan, off Nambu Town, Hidaka County, Wakayama
Prefecture, bottom gill net on rocky substrate, 31 January 1969.

DISTINGUISHING FEATURES. Preserved specimens mostly pallid; pectoral
fin rays 18; no ocellus or dark spot on soft dorsal fin; vertical scale rows about
70; skin flaps present on lower jaw.

Remarks. The specimens reported here agree with the holotype in most
features. The third dorsal spine is elongate in the holotype (fig. 3) but it is
not especially elongate in the other specimens (figs. 2B, 4). The specimens agree

in counts. All have 18 pectoral rays, rays 2—7 branched in the holotype (also

Vou. XXXIX] ESCHMEVER ET AL.: NEW SCORPIONFISHES 293

Z2
PEE |
CLO

ZZ
GAIT LL
ERS

Ficure 3. Rhinopias xenops, holotype, USNM no. 51604, 110 mm. S.L., Hawaii.

the 9th on left side in the holotype), rays 1—7 or 2 through 6 or 7 branched
in the other specimens. Gill rakers (including rudiments) total 21 to 23, 6(7)
on upper arch, 15—17 on lower arch. Vertical scale rows number about 70-75,
and lateral line scales 22 or 23. Head spination as for the genus. Nasal spines
small. Supplemental preopercular spine present, small. Sphenotic spines absent
or small.

An extensive color description was given by Gilbert (1905, p. 631) as follows:

“Head, body, and fins bright vermilion, upper parts of head and body darkened with
olive tint, and with small scattered purplish spots, which are also found on upper half
of pectoral fin; head, lower parts of body, and fins mottled with yellowish-white;
flaps and tentacles narrowly edged with bright lemon-yellow; a large blackish blotch
below eye, one on opercle and one at base of pectoral; a conspicuous broad, yellowish
white bar on each side of compressed part of rostrum; three groups of brownish spots
along base of dorsal fins; conspicuous white spots on back of tail and at base of
eighth and ninth dorsal spines; a larger blotch below the latter just above lateral line.”

No information on fresh coloration is available for the additional specimen from
Hawaii. A Kodachrome slide of a Japanese specimen (SMBL 7202) revealed
the following: body and head mostly orange-red; small purplish specks on
upper part of head and body; large patch of white on body below dorsal spines

294 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH Ser.

Ficure 4. Rhinopias xenops. A. SMBL no. 7201, 114 mm. S.L., Japan. B. SMBL no.
7202, 129 mm. S.L., Japan.

7-10, with a smaller white spot just above lateral line below this large white
patch; other white patches on head, pectoral fin, and scattered on body; small,
tapering pale stripe leading forward and down from eye. The coloration of the
Japanese specimen is similar to that given by Gilbert for the holotype with a
few exceptions. The Japanese specimen had more orange and less red pigment
on the head and body and lacked most of the lemon-yellow pigmentation as
found by Gilbert. Some differences may result from occurrence at different

VoL. XX XIX] ESCHMEYER ET AL.: NEW SCORPIONFISHES 295

depths; the Japanese specimen was on display in an aquarium when photo-
graphed while Gilbert’s description was taken from the holotype recovered from
32-43 fathoms. Dusky patches were similar as evidenced by the preserved
condition. The major features of the distribution of white patches were very
similar.

Coloration in preservative (figs. 2B, 3-4) for all specimens is basically
similar, the most conspicuous feature being the dusky pigment at the base of
the soft dorsal fin.

Measurements are given in table 1.

DIsTRIBUTION. Rhinopias xenops is now known from Hawaii and Japan.
Previous records of this species from Japan were based on specimens of R.
frondosa. Depths of capture for the Hawaiian specimens were between 32 and
43 fathoms and between 52 and 60 fathoms. Depths of capture for the Japanese
specimens are not available, but they were captured offshore. Habitat appears
to be rocky or coralline areas.

Name. ‘Nise-boro-kasago’ is proposed as the Japanese common name, from
‘nise’ (pseudo) + ‘boro’ (ragged or tattered) + ‘kasago’ (Sebastiscus marmor-
atus or similar fish). Boro-kasago is the Japanese name for Rhinopias frondosa.

Rhinopias frondosa (Giinther).
(Figures 5-7.)

Scorpaena frondosa GUNTHER, 1891, pp. 482-483, pl. XXXIX (original description; type
locality Mauritius; holotype BMNH 1891.4.30.3).

Peloropsis xenops (not of Gilbert), Kamowara, 1942, pp. 27-28, fig. 1 (description; differences
between his specimens and the holotype of R. xenops; two specimens from Shirahama,
Wakayama Prefecture, Japan) ; 1950, p. 219, fig. 165 (compiled; figure from Kamohara,
1942).

Peloropsis frondosa, DERANIYAGALA, 1952, p. 109, pl. 32 (specimen from Ceylon; figure
poor; lengths and body proportions incorrect [see SMITH, 1966, p. 77]).

Peloropsis frondosus, Munro, 1955, p. 250, pl. 48, fig. 726 (compiled; figure copied from
Deraniyagala, 1952); Kamonara, 1959, pp. 5-6, fig. 2 (description; one specimen from
Susaki Fish Market; figure from Kamohara, 1942); 1964, pp. 73-74, fig. 47 (listed;
Kochi Prefecture, Japan; figure from Kamohara, 1942) ; FouRMANoIR and NHU-NHUNG,
1965, p. 93, fig. 59 (two specimens from Viet Nam; line drawing) ; FourRMANoIR and
GuEkzE, 1966, pp. 56-57, fig. III d (Reunion Island, 90 m.)

Rhinopias frondosa, SmirH, 1957, p. 62, pl. 4, fig. C (compiled; copied from Giinther) ;
PALMER, 1963, pp. 701-704, pl. XX (summary of earlier records of Rhinopias; treated
R. xenops and R. frondosa as separate species; radiographs of R. xenops and R. frondosa) ;
KotrHaus, 1966, p. 122 (figure of a specimen from the northern Indian Ocean) ; SMITH,
1966, pp. 74-79, fig. B on pl. 14 (synonymy; description; new record for South African
waters; summary of earlier knowledge on R. xenops and R. frondosa and differences
between them; retouched photo of a 45 mm. S.L. specimen).

MatTERIAL. BMNH no. 1891.4.30.3 (1, 146 mm. S.L., holotype), Mauritius,

collected by Robillard. MNHN 1967-550 (1, 79.5 mm. S.L.), Reunion Island.
BAH uncat. (1, 145 mm. S.L.), off Somalia, 6°06’ to 6°03’N., 49°05’ to 49°03’E.,

296 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Ficure 5. Rhinopias frondosa, holotype, BMNH no. 1891.4.30.3, 146 mm. S.L., Mauritius
(from Giinther, 1891, pl. 39).

in 55-65 meters, Meteor station 123. KU no. 8465 (1, 152 mm. S.L.), Japan,
Kochi Prefecture, Susaki Fish Market, 12 March 1959. SMBL no. 7203 (1,
127 mm. S.L.), Japan, Nambu Town, Hidaka County, Wakayama Prefecture,
gill-net on rocky bottom, 17 January 1959. SMBL no. 7204 (1, 146 mm. S.L.),
Japan, off Iwashiro, Hidaka County, Wakayama Prefecture, lobster gill-net
on rocky bottom, 18 April 1959. USNM no. 209416 (1, 103 mm. S.L.) and
CAS no. 15320 (1,115 mm. S. L.), Caroline Islands, Condor Reef, in 30 fathoms,
12.5-m. shrimp trawl, Townsend Cromwell Cruise 57, 24 May 1972.

DISTINGUISHING FEATURES. Pectoral rays 16 (rarely 15, but 17 rays would
not be unusual); dark spot on membrane of soft dorsal fin between rays 7-9,
sometimes more restricted; body and fins of preserved specimens covered with
large brown circles or with pale spots or oblong blotches on light brown back-
ground, vertical scale rows about 70-75; skin flaps present on lower jaw.

Remarks. The known specimens up until now of this species have been
reviewed in two articles (Palmer, 1963; Smith, 1966). Both authors reached
the conclusion that R. frondosa is distinct from R. xenops. The color patterns
of all the known preserved specimens of R. frondosa are similar except for one
from Nouméa reported on by Whitley (1964, p. 9, pl. IT). The Nouméa speci-
men is felt by us to represent a separate species and is treated as a new species,
R. aphanes, in a separate section of this paper.

VoL. XXXIX] ESCHMEVER ET AL.: NEW SCORPIONFISHES 297

FicurE 6. Rhinopias frondosa. A. KU no. 8465, 152 mm. S.L., Japan. B. MNHN no.
1967-550, 79.5 mm. S.L., Reunion Island.

Measurements and counts are presented in table 1. The holotype, speci-
mens from Japan, Reunion Island, and the Caroline Islands are figured (figs.
5-7).

Color slides provided by Dr. Araga of 2 Japanese aquarium specimens (both
about 230 mm. T.L.) revealed the following: specimen 1 with body, head, and
fins reddish pink (approaching maroon); all of specimen covered with mostly

298 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Tasie 1. Counts and measurements for specimens of the genus Rhinopias. (Measurements
are in mm.; percent standard length in parentheses; see Materials Examined for catalog num-
bers and locality information; holotype marked with an asterisk.)

: R. xenops se R.

R. argoliba aphanes frondosa
Standard length 129* 107 110* 114 129 178* 79.5
Dorsal rays XII-+9% XII-+9% XII-+9% XII-+-9% XII-+9% XII-+9% XII-+9%
Anal rays TWI-+5% WI-+5% I-+5% I11-+5% Ii-+5% TWi-+5% Ii+5%
Pectoral rays 18-18 18-18 18+18 18-418 18-208 (ios Simonetta
Gill rakers 7416 6415 6416 6417 6-7--16 “S6-oIe ero ons
Vertebrae 24 24 24 24 24 24 24
Vertical scale rows 80 70 68 76 73 70 —

(approximate)

Head length 59.5(46) 51.7(48) 51.2(46) 53.9(47) 60.1(47) 85.5(48) 36.8(46)
Body depth 55.2(43) 45.7(43) 48.7(44) 47.4(42) 53.3(41) 82.1(46) 35.0(44)
Orbit diameter 10.3(08) 9.5(09) 10.0(09) 10.0(09) 11.1(09) 13.5(07) 7.0(09)
Snout length 26.5(21) 21.1(20) 20.6(19) 22.2(19) 25.6(20) 40.0(22) 14.8(19)
Interorbital width 5.8(04) 4.4(04) 5.3(05) 4.5(04) 6.0(05) 10.5(06) 4.2(05)
Jaw length 29.5(23) 26.4(25) 26.4(24) 28.4(25) 31.2(24) 42.0(24) 16.9(21)
Predorsal-fin length 47.7(37) 40.7(38) 42.2(38) 45.6(40) 48.8(38) 74.1(42) 29.3(37)
Caudal fin length 32.6(25) — — 33.6(29) 38.6(30) 52.6(30) 30.5(38)
Pectoral fin length 37.5(29) 33.3(31) 39.9(36) 42.2(37) 45.6(35) 67.7(38) 30.3(38)
Pelvic fin length 29.2(23) 29.3(27) 33.6(30) 31.3(27) 33.2(26) 49.3(28) 23.9(30)
Length 1st anal spine 9.6(07) 11.6(11) 13.2(12) 10.6(09) 13.2(10) 16.1(09) 8.7(11)

Length 2nd anal spine 17.2'(13)) 225021). 22/6(21)| 23:0(20)) 2417) 2851s) eet ets))
Length 3rd anal spine 20:0(116)' 21-420)" 222721), 217/19) 21S) SZ 14D) Sent)
Length 3rd dorsalspine 20.3(16) 25.2(24) 40.0(36) 34.7(30) 35.3(27) 41.7(23) 22.6(28)

Length 4th dorsalspine 21.2(16) 25.8(24) 31.2(27) 32.8(29) 30.9(24) — 24.9(31)
Length 11th dorsalspine 11.7(09) — — 12.1(11) 11.7(09) 8.5(05) 6.3'(08)
Length 12th dorsal spine 12.2(10) -- — 16.3(14) 16.8(13) 21.9(12) 13.6(17)

circular white spots or blotches, but with central area of blotches or spots usually
colored same as body; pigment around edges of these pale areas slightly darker
than surrounding areas; some small pale blotches between larger circular areas.
Specimen 2 with more pink on fins and tentacles, and with head and body more
brownish red. Transparent, oblong areas between upper pectoral rays at about
midheight of fin visible in specimen two. Kamohara (1959) reported that the
body (our KU 8465, 152 mm. S.L.) was uniform reddish; head bluish, lower
parts of body and fins mottled with white, and black blotch on membrane be-
tween seventh and eighth dorsal rays. Smith (1966, p. 77, pl. 14B) reported
the coloration of his 45 mm. S.L. specimen as follows:

“Colour in life. . milky yellow, with orange tinge over the opercle. The body is
covered with thin brown lines forming irregular loops of varying size and shape. The
iridal flaps are almost black, giving a radial effect. There are a few prominent dark

VoL. XXXIX] ESCHMEVER ET AL.: NEW SCORPIONFISHES 299
TABLE 1 (Continued)
R. frondosa (continued)
Standard length 103 115 127 145 146* 146 152
Dorsal rays XII+9% XII-+9% XII+9% XII-+9% XII-+9'% XII-+9'% XII+91%4
Anal rays T-+5% WWI-+5% WI-+5% WI+5% I1+5% WI+5% IlI+5%
Pectoral rays 16+16 16+16 16+16 16+16 16+16 16+16 16+16
Gill rakers 6+15 6+15 7+15 —- 6+15 6-7+15 6+14
Vertebrae 24 24 24 --= 24 24 24
Vertical scale rows 70-75 U2 -- -= — 75 75
(approximate)

Head length 46.3(45) 57.1(50) 59.0(46) 65.3(45) 66.6(46) 68.0(46) 67.3(44)
Body depth 46.8(45) 59.6(52) 61.6(48) 66.3(46) 66.8(46) 69.8(48) 70.7(46)
Orbit diameter 8.2(08) 11.2(10) 10.3(08) 11.4(08) 12.4(08) 13.0(09) 12.0(08)
Snout length 20.5(20) 23.8(21) 26.6(21) 30.2(21) 29.9(20) 28.9(20) 29.1(19)
Interorbital width 6.3(06) 7.7(07) 8.3(06) 8.0(06) 8.6(06) 9.7(07) 8.3(06)
Jaw length 20.5(20) 26.0(23) 25.8(20) 30.1(21) 30.8(21) 30.7(21) 32.1(21)
Predorsal-fin length 37.6(36) 46.2(40) 47.8(38) 50.6(35) 58.0(40) 56.5(39) 56.4(37)
Caudal fin length 35.7(35) 43.7(38) 41.9(33) 48.0(33) 49.0(34) 47.4(32) 48.5(32)
Pectoral fin length 37.3(36) 48.4(42) 47.4(38) 49.5(34) 49.5(34) 53.9(37) 55.1(36)
Pelvic fin length 26.2(25) 32.3(28) 33.4(26) 39.2(27) 42.1(29) 40.3(28) 37.8(25)
Length 1st anal spine 8.4(08) 9.3(08) 8.8(07) — 10.5(07) 11.2(08) 10.0(07)
Length 2nd anal spine IBLOKALS) UWS SGEDY aS (CI) --- 19.0(13) 17.8(12) 17.9(12)
Length 3rd anal spine 16.3(16) 18.5(16) 17.1(13) — 21.7(15) 20.9(14) 21.7(14)
Length 3rd dorsal spine 27.4(27) 39.7(34) 38.0(30) — AG NGD) EVO Sees)
Length 4th dorsal spine 27.5(27) 38.6(34) 33.0(26) —_— 53.0(36) 47.7(33) 36.2(24)
Length 11th dorsal spine 5.3(05) 5.8(05) 6.2(05) — 9.0(06) 9.7(07) 7.8(05)
Length 12th dorsal spine 13.6(13) 18.2(16) 17.1(14) — 19.0(13) 24.4(17) 20.4(14)

1 Broken at tip.

marks on the fins, one rectangular, low, between the 5—6th dorsal spines, one, smaller,
behind the base of the ninth dorsal spine, one across the upper part of the second
dorsal ray, a small one on the upper and another at the lower part of the caudal.
The largest is low down between the seventh and eighth dorsal rays, and the smallest
(but distinct) between the apices of the first and second (upper) rays of the pectoral

fin.”

Deraniyagala (1952, pl. 32) illustrates a 126-mm. T.L. specimen as mostly
orange-red. The general body coloration appears to change from mostly yel-
low in small specimens to mostly red or crimson in large specimens. Our speci-
men from Reunion Island (fig. 6B) has the dark pigment most concentrated
in the areas mentioned by Smith as prominent dark marks. The retention in
preservative of the dark, mostly circular lines (outlining the pale areas) is vari-
able (figs. 5-7) and seems to depend on the amount of black or brown pigment

300 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Ficure 7. Rhinopias frondosa, CAS 15320, 115 mm. S.L., Caroline Islands.

present initially and perhaps on the method of preservation. Kotthaus (1966,
pl. 22) provides an excellent figure of a 145-mm. S.L. specimen.

Spination as for the genus. Nasal spines usually absent. Supplemental pre-
opercular spine absent or poorly developed. Sphenotic spines present.

The reader is referred to the papers cited above for a detailed description of
R. frondosa.

DiIstRIBUTION. Rhinopias frondosa is known from Mauritius, Reunion
Island, Natal, Somalia, the Arabian Sea, Ceylon, Vietnam, Japan, and the
Caroline Islands. Few depths of capture are available: dredged in 13 meters
in Ceylon (Deraniyagala, 1952); in 55-65 meters off Somalia, in 90 meters at
Reunion, and in 55 meters at the Caroline Islands. Habitat is rocky or coralline
areas.

Name. The Japanese name ‘Boro-kasago’ has been used for this species;
from ‘boro’ (ragged or tattered) + ‘kasago’ (Sebastiscus marmoratus or similar
species ).

Rhinopias aphanes Eschmeyer, new species.

(Figure 8.)

Rhinopias frondosa (not of Giinther), WuitLry, 1964, p. 9, pl. II (3-sentence account re-
porting a specimen from New Caledonia [this specimen now holotype of R. aphanes]).
Hototyre. AMS IB7079 (178 mm. S.L., 235 mm. T.L.), New Caledonia,

outside barrier reef facing Nouméa, at a depth of 30 meters, collected by Yves

Merlet and Madame R. Catala-Stucki [transferred alive to the Nouméa Aquar-

ium where it lived for less than a week].

VoL. XXXIX] ESCHMEYVER ET AL.: NEW SCORPIONFISHES 301

Ficure 8. Rhinopias aphanes, holotype, AMS no. IB7079, 178 mm. S.L., New Caledonia.

Description. (Based on the holotype and only known specimen.) A large-
headed, strongly compressed scorpionfish. Dorsal fin with 12 spines and 9%2
soft rays; no dorsal spine notably longer than the others, third and fourth spines
the longest, eleventh dorsal spine about % the length of the twelfth spine. Anal
fin with 3 spines and 5’ soft rays; third spine slightly longer than second, third
extends past second when fin is depressed. Pectoral rays 16, rays 1 or 2 through
6 branched. Gill rakers, including rudiments, total 20, 6 on upper arch, and 14
below, including one at angle; all rakers as short spiny knobs. Scales on body
small and cycloid; about 70 vertical scale rows (counting a few scale rows above
lateral line from above first lateral line scale to end of hypural plate). Head
without scales.

Spination similar to that in other species of Rhinopias; see account under
the genus. Preorbital bone with two slight lumps over maxillary, obscured by
tentacles. Suborbital ridge with 4 spines, first as tiny lump on lateral face of
preorbital bone, followed by one below anterior part of orbit and two posterior
ones (poorly defined) just before preopercle bone. Nasal spines small. Pre-
ocular spine well defined; supraocular spine obscured by tentacle; postocular
spine very large and broad. Sphenotic spine as a lump. Supplemental pre-
opercular spine small, obscured by tentacle. First and second preopercular
spines longest; third and fourth smaller, fifth almost imperceptible.

Measurements for the holotype in table 1.

Tentacles and skin appendages numerous; most slender; black or striped
with black; most branched.

Preserved coloration as in figure 8. Transparent ovoid and round areas

302 CALIFORNIA ACADEMY OF SCIENCES [Proc, 4TH SER.

present between the pectoral rays, most prominent as a row near mid-height of
pectoral fin. A slide and color photographs taken by Madame Catala-Stucki
when the specimen was alive show that the holotype was a beautiful yellow and
black. The pallid areas in figure 8 were lemon yellow, with a few traces of
white (particularly in one patch on the suborbital stay), and the dark areas
in figure 8 were black to dark brown. The skin flaps and tentacles were black
bordered by yellow. The transparent areas of the pectoral fin were apparent.

COMPARISONS AND REMARKS. Although this specimen was originally identi-
fied as R. frondosa (Whitley, 1964; Smith, 1966), its color pattern seems dis-
tinctive and it is thought to represent a separate species. All of the reported
specimens of R. frondosa seem to have a fairly consistent color pattern as shown
in our figures 5—7, and the figures reported in the synonymy. The holotype of
R. aphanes is a little larger in size than the holotype and largest known specimen
of R. frondosa, but it seems unlikely that the color pattern would change this
much with growth; there is no indication that the roundish pale areas of R.
frondosa would become elongate as in R. aphanes (see especially the head). The
differences in coloration are apparent by comparing figures 5—7 with figure 8.
Other differences between R. frondosa and R. aphanes are as follows: Rhinopias
aphanes appears to have a smaller orbit, longer snout, a longer jaw, a longer
predorsal-fin length, a shorter caudal fin, and some shorter fin spines (table 1) ;
some of these differences may not remain when additional material is found.

Rhinopias aphanes is easily separated from R. xenops and R. argoliba on
the basis of pectoral rays (16 instead of 18) and coloration.

DISTRIBUTION. This species is known only from the holotype collected near
Nouméa, New Caledonia.

NamME. The species name is from the Greek word aphanes (aph’-a-nés), a
noun, meaning that which is unapparent or inconspicuous, alluding to the cam-
ouflage provided by its remarkable coloration and skin flaps.

The common name ‘Merlet’s scorpionfish’ is suggested. Doctor Yves Merlet,
an avid diver and underwater naturalist, captured the holotype. Over a 12-
year period he made available numerous valuable specimens for display in the
Nouméa Aquarium and for study by systematists. He died in a diving accident
in 1969.

RELATED GENERA AND SPECIES

Other genera and species which appear to be related to, or members of,
Rhino pias are discussed below. The subfamily name Pteroidichthyinae Fowler
is available. Fowler (1938, p. 51, in key) provided the following meager de-
scription for the subfamily: ‘Pterodichthyinae, new subfamily. Dorsal spines
Mie ranaleSpInes: Zecca. Pteropelor.” The subfamily name is based on the genus
Pteroidichthys Bleeker. Fowler’s incorrect spelling of the subfamily (no i before

VoL. XXXIX] ESCHMEYVER ET AL.: NEW SCORPIONFISHES 303

the d) was emended by Whitley (1954). In the subfamily description in the
key Fowler listed the dorsal spines as 11 and anal spines as 2, but in his de-
scription of Pteropelor, the only genus he specifically included in the subfamily,
he listed dorsal spines as 12 and anal spines as 3. Problems surrounding the
dorsal and anal spine counts are discussed below. Whitley (1954, p. 60) is
evidently the only subsequent worker to use the subfamily name, and Whitley
recognized Pteropelor as a synonym of Rhinopias and referred Rhinopias to the
subfamily Pteroidichthyinae. It seems likely that the nominal genera Rhino pias,
Pteropelor, and Pteroidichthys, as well as the genera Pogonoscorpius Regan and
Hipposcorpaena Fowler, are closely related. They appear to be derivatives of
the subfamily Scorpaeninae, differing mainly in having a more compressed body;
but whether they should be retained as a separate subfamily must await a more
detailed study.

The monotypic genus Pogonoscorpius Regan is here referred to the subfamily
Pteroidichthyinae; in fact it may be a synonym of the genus Rhinopias. The
monotypic genus Hipposcorpaena Fowler, based on a juvenile, also seems closely
related to Rhinopias. The nominal genera Pteroidichthys and Pteropelor are
each known from a single species based on 1 or 2 specimens. Rhinopias godfreyi
Whitley, also known from a single specimen, seems referable to Pteroidichthys.
The status of each of these nominal genera and species is discussed below, and
some corrections for information given in the original descriptions are made based
on examination of the type specimens by the first author.

1. Pogonoscorpius.

Pogonoscorpius REGAN, 1908, p. 236 (type-species Pogonoscorpius sechellensis by original
designation; monotypic).

Pogonoscorpius sechellensis REGAN, 1908, p. 236, fig. 3 on pl. 28 (original description; type
locality Seychelles; holotype BMNH 1908.3.23.172).

Pogonoscorpius seychellensis, SMITH, 1957, pp. 51 and 59, fig. D on pl. 4 (compiled; figure
from Regan, 1908; misspelled species).

The genus Pogonoscorpius and the species P. sechellensis are known only
from the holotype of the species. It is clear that Regan intended the species
name to be spelled sechellensis rather than seychellensis, since the same spelling
was used both for the description and figure, and Regan also used this same
spelling for Champson sechellensis, Synchiropus sechellensis, and Scaeops sechel-
lensis, three other new species described by him in the same work. Smith’s spell-
ing (seychellensis) is an unjustified emendation.

The holotype agrees with species of the genus Rhinopias in meristic features,
dentition, spination, and body shape. It differs from them in coloration, dis-
tribution of skin flaps (particularly the presence of a median chin barbel), and
a less elevated orbit. The drawing accompanying the original description (Regan,
1908, fig. 3 on pl. 28) is poor. We have reproduced a photograph of the holotype

304 CALIFORNIA ACADEMY OF SCIENCES [PRoc. 4TH SER.

Ficure 9. Pogonoscorpius sechellensis, holotype, BMNH no. 1908.3.23.172, 51.4 mm.
S.L., Seychelles.

(fig. 9) which better illustrates the body shape. Additional study when speci-
mens become available may show that this species is best placed in the genus
Rhino pias.

The following counts for the holotype were recorded: dorsal rays XII + 9%;
anal rays III + 5%; pectoral rays 18+18, all rays appear to be unbranched;
vertical scale rows estimated at about 60 as given by Regan, many missing; gill
rakers difficult to count, 7 on upper arch, 9 (including one at angle) plus addi-
tional rudiments on the lower arch. Spination about as in species of the genus
Rhinopias; supplemental preopercular spine present only on left side, most
spines small or low. Some measurements (in mm., percent standard length in
parentheses) were recorded as follows: standard length 51.4; total length about
66; head length 23.7 (46); body depth 19.8 (38); orbit diameter 4.5 (09);
snout length 10.1 (20); interorbital width 2.2 (04); jaw length 12.1 (24); pre-
dorsal-fin length 20.3 (40) ; caudal fin length 14.4 (28); pectoral fin length 15.9
(31); pelvic fin length 12.3 (24).

Coloration as given by Regan, “yellowish; fins tinged with reddish.”

This species appears to be a valid one and is known only from the holotype
collected in the Seychelles in 37 fathoms.

2. Hipposcorpaena.

Hipposcorpaena Fow er, 1938, pp. 71-72 (type-species Hipposcorpaena filamentosa, by
original designation; monotypic).

Hipposcorpaena filamentosa Fow ier, 1938, pp. 72-73, fig. 31 (original description; type
locality Philippine Islands, Gulf of Davao, 7°04’48”N., 125°39'38”E., in 28 fathoms,
Albatross station D.5253, 18 May 1908; holotype USNM 98819).

The genus Hipposcorpaena was based on a single specimen of about 29 mm
S.L., USNM 98819. One additional Philippine specimen is available (USNM
168183, Albatross station 5174) and is of about the same size as the holotype.

VoL. XXXIX] ESCHMEYER ET AL.: NEW SCORPIONFISHES 305

The holotype is now in poor condition. Fowler (1938, p. 73) gives a dorsal fin
count of XI + 10, but this should have been XII + 9%; he lists anal rays as 9,
but they total 8’ and seem to be III + 5%. Pectoral rays appear to number
14 as given by Fowler. The rays near the lower end of the pectoral fin are the
longest, and the fifth from the bottom extends past the posterior base of the
anal fin (not as in Fowler’s figure 31); the lower principal caudal rays are also
the longest. The vertical scale rows were given by Fowler as 30; although many
scales are missing, the scale rows probably number about 35—40. Hipposcorpaena
filamentosa appears to be a valid species known only from the Philippines.

3. Pteroidichthys.

Pteroidichthys BLEEKER, 1856, pp. 33-34 (type-species Pteroidichthys amboinensis Bleeker
by original designation; monotypic ) ; 1876, p. 5 (generic diagnosis).

Pteroidichthys amboinensis BLEEKER, 1856, pp. 34-35 (original description; type locality
Amboina and Manado, Celebes; two syntypes, RMNH 5873); GUNTHER, 1860, p. 127

(compiled) ; BLEEKER, 1876, pp. 5, 9-10, 12-13, 57-58, fig. 5 on pl. 1 (description com-
piled from Bleeker, 1856; figure good); 1879, fig. 1 on pl. 414 (figure from Bleeker,
i876) ; Herre, 1952, p. 436 (compiled from Bleeker) ; pe BrAurort im Weber and de

Beaufort, 1962, p. 54 (re-examined types; figure redrawn from Bleeker, 1876).

The two syntypes, RMNH 5873, were briefly examined. They are in fairly
poor condition and now measure about 67 and 80 mm. in total length and 51 and
57 mm. in standard length. The counts as given by Bleeker and by de Beaufort
are dorsal rays XI + 11 and anal rays II + 7. The twelfth element in the dorsal
fin does not appear to be segmented, although this is difficult to determine and
the ray is long and flexible; counting by the methods used in this paper the dorsal
rays number XII + 912, with the twelfth element very long and at the leading
edge of the soft dorsal fin; anal rays I] + 6%. For further information see
‘Remarks’ below.

4. Pteropelor.

Pteropelor FOWLER, 1938, p. 77 (type-species Pteropelor noronhai Fowler by original des-
ignation; monotypic).

Pteropelor noronhai Fow er, 1938, pp. 78-79, fig. 34 (original description; type locality
vicinity of Hong Kong, China Sea, 21°33’N., 116°13’E., in 100 fathoms, Albatross
station D.5310, 4 November 1908; holotype USNM 98892, paratype USNM 99009).
The genus and species were based on two juvenile specimens of 32.7 (USNM

98892) and 35.5 (USNM 99009) mm. in standard length. Fowler (1938, in

key, p. 51) states that the dorsal spines number 11 and the anal spines 2, but

in the description (pp. 78-79) he reports 12 dorsal spines and 3 anal spines. As
with Pteroidichthys amboinensis and Rhinopias godfreyi, there is difficulty in
determining if the twelfth dorsal fin element and the third anal fin element are
spines or segmented rays. In Pteropelor noronhai these fin rays appear to be
unsegmented. The dorsal spine count would be XII + 9%. The paratype has

306 CALIFORNIA ACADEMY OF SCIENCES [PRroc. 4TH SER.

a total of 8% anal rays (? III + 5%) and the holotype a total of 9% (? III+
6’2). Fowler states that there are only 5 or 6 lateral line scales anteriorly, but
examination of the types shows that the lateral line is probably complete, but
many scales have been rubbed off; at least nine tubed scales are present in one
specimen. Fowler gives the vertical scale rows as 25 + 2; we estimate that they
number considerably more than this, but most scales have been rubbed off.
Further information is provided in the ‘Remarks’ below.

5. Rhinopias godfreyi.

Rhinopias godfreyi WHITLEY, 1954, pp. 60-61, pl. 3, fig. 2 (original description; type locality

Exmouth Gulf, northwestern Australia; holotype AMS IB2977).

This species is known only from the holotype, a specimen 46.1 mm. in stan-
dard length. The counts for the specimen are pectoral rays 15, anal rays II +
6%, and the dorsal count is either XI + 10’ or XII + 9%; the twelfth dorsal fin
element appears to be unsegmented; the third anal fin element is segmented
distally. The vertical scale rows number about 40; the lateral line seems to
have been complete, although some scales are rubbed off. (Whitley incorrectly
lists the pectoral rays as 14 and anal rays as III + 6; he gave the dorsal ray
count as XI + 10.)

Remarks. Pogonoscorpius and Hipposcorpaena are retained for now as
separate monotypic genera which appear to be closely related to the genus
Rhino pias. On the other hand, Pteroidichthys amboinensis, Pteropelor noronhai,
and Rhinopias godfreyi are very closely related species, and Pteropelor is con-
sidered a synonym of Péeroidichthys. (Whitley (1954) went even further and
considered Pteroidichthys a synonym of Rhinopias.) It is possible that more
thorough study of the available specimens and additional specimens as they
are captured will show that Pteroidichthys amboinensis Bleeker 1856, Pteropelor
noronhai Fowler 1938, and Rhinopias godfreyi Whitley 1954 are all based on
different sized specimens of the same species, or at least that some of them are
synonyms of one another. These specimens are characterized by having dorsal
fin rays XII + 9'2 (the twelfth element which is at the leading edge of the
soft dorsal fin may be segmented in large specimens), anal rays normally II +
6% (possibly with the third element unsegmented in small specimens); and
pectoral rays 14 or 15, rays unbranched (?). They are further characterized
by having a very compressed body, very long fins (particularly the anal fin),
and long supraocular tentacles which are branched in large specimens. They
differ from species of the genera Rhinopias, Pogonoscorpius, and Hipposcorpaena
most notably in the condition of the third anal fin element (two rather than three
anal spines) and also in such features as a less deep body, shorter head, and
lower pectoral ray count. If these three nominal species are all the same species
then the localities from which it has been collected are Hong Kong, Celebes,

VoL. XXXIX] ESCHMEVER ET AL.: NEW SCORPIONFISHES 307

Ficure 10. Pteroidichthys amboinensis, ZMC uncat., 36.4 mm. S.L., Vietnam.

Amboina, and northwestern Australia. It is an offshore species, probably living
in coral in depths from perhaps 30 to 100 fathoms.

One additional specimen (fig. 10) was found which is referable to this com-
plex. The specimen measures 36.4 mm. in standard length and is from the col-
lection of the Zoological Museum of Copenhagen with the following data: col-
lected at Nhatrang Bay, Vietnam, 16 November 1959, by J. Knudsen. This
specimen has counts of dorsal rays XII + 9%, anal rays II + 6%, and pectoral
rays 16+ 15 (all simple). In coloration it is most like the type of Pteropelor
noronhai.

SHEDDING OF ‘SKIN’

Sloughing of ‘skin’ in the scorpaenid Taenianotus triacanthus has been dis-
cussed recently by Wickler and Nowak (1969), although they missed an earlier
account of shedding in Taenianotus by Faulkner (1961). The first mention of
shedding was by Gilchrist (1920) for Agriopus (subfamily Congiopodinae).
Hirosaki (1957) discussed shedding in scorpaenids, including the genera Pterois,
Parapterois, and Erosa, and other fishes, and (1971a—c) in Rhinopias argoliba.
Faulkner (1961) also reported sloughing of skin in Pterois sphex. From museum
specimens we can report that this shedding also occurs in stonefishes (Synanceia,
subfamily Synanceiinae), the genus /nimicus, and evidently all species of Pterois
(subfamily Pteroinae). The holotype of Rhinopias aphanes also has pieces of
‘skin’ hanging from it as do some specimens of R. xenops and R. frondosa.

Previous authors who have commented on the shed layer have described it
as skin, and have correctly pointed out that it is not mucous. Samples have been
analysed for us by Richard Krejsa, and he informs us that the shed layer is
best termed a cuticle, an epidermal product; some cells are lost with the shed
layer but the basic matrix is cuticular. Further histological studies are being
made by Dr. Krejsa.

Hirosaki (1971a—c) found that in R. argoliba shedding occurred in the
Enoshima Aquarium 9 times in 114 days at regular intervals (January 30, Feb-

308 CALIFORNIA ACADEMY OF SCIENCES [PRroc. 4TH SER.

ruary 13 and 28, March 12 and 30, April 11 and 24, May 8 and 18). The cuticle
from the entire body was shed each time. It appears that shedding was a natural
event not caused by any sort of shock. The function of the shedding probably
involves the removal of algae, external parasites, and other objects which might
accumulate on the skin of these basically sedentary fishes.

The genera /nimicus, Minous, and Choridactylus are peculiar scorpaenids
which have one to three lower pectoral rays thickened and free. Each free ray
is tipped with a ‘friction pad’ or ‘cap.’ The caps are easily dislodged in preserved
specimens. These thickened caps, covering the distal one-fourth to one-tenth
of each ray, are made of the same cuticular substance, although much thicker,
as the shed layer of the scorpaenids listed above; almost certainly these caps are
used as an aid for ‘walking’ on the bottom with the free pectoral rays. (These
‘caps’ will be discussed more fully in a subsequent paper on the subfamily
Minoinae. )

LITERATURE CITED

AXELROD, HERBERT R., and WARREN E. BuRGESS
1972. Pacific marine fishes, Book 1. T. F. H. Publications, Neptune City, New Jersey,
280 pp., 489 color figs.
BLEEKER, PIETER
1856. Beschrijvingen van nieuwe en weinig bekende vischsoorten van Amboina, . . .
Acta Societatis Scientiarum Indo-Neerlandicae, vol. 1, pp. 1-76.
1876. Mémoire sur les espéces insulindiennes de la famille des Scorpénoides. Verhande-
lingen der Koninklijke Akademie van Wetenschappen, vol. 16, pp. 1-100,
pls. 1-5.
1879. Atlas ichthyologique des Indes Orientales Néerlandaises. . .Amsterdam, vol. 9,
pls. 355-360, 363-420.
DERANTYAGALA, P. E. P.
1952. A colored atlas of some vertebrates from Ceylon. Vol. 1. Ceylon National Mu-
seums Publication, 149 pp., 34 pls., 60 text figs.
ESCHMEYER, WILLIAM N.
1969. A systematic review of the scorpionfishes of the Atlantic Ocean (Pisces: Scor-
paenidae). Occasional Papers of the California Academy of Sciences, no. 79,
130 pp., 13 figs.
FAULKNER, S. DOUGLAS
1961. Fishes that shed skin, sway like ocean plants. Taenianotus triacanthus. Aquarium
Journal, April, 1961, pp. 169-170.
FourMANoIR, P., and P. GUEZE
1967. Poissons nouveaux ou peu connus provenant de la Réunion et de Madagascar.
III. Sept espéces intéressantes trouvees récemment dans les eaux Malgaches
et Réunionnaises. Cahiers ORSTOM, Océanography, vol. 5, no. 1, pp. 55-58.
FourMANoIR, P., and Do-THI NHU-YUNG
1965. Liste complémentaire des poissons marins de Nha-trang. Cahiers ORSTOM,
Oceanography, no. spécial, Juillet, 1965, 114 pp.
FOWLER, HENRY W.
1928. The fishes of Oceania. Memoirs of the Bernice P. Bishop Museum, vol. X, pp.
1-540, pls. I-XLIX.

Vor. XXXIX] ESCHMEYER ET AL.: NEW SCORPIONFISHES 309

1938. Descriptions of new fishes obtained by the United States Bureau of Fisheries
Steamer “Albatross”, chiefly in Philippine Seas and adjacent waters. Pro-
ceedings of the United States National Museum, vol. 85, no. 3032, pp. 31-135,
text figs. 6-61.
GILBERT, CHARLES HENRY
1905. The deep-sea fishes of the Hawaiian Islands. Pp. 577-713, pls. 66-111, figs. 230-
276, in David Starr Jordan and Barton Warren Evermann, The aquatic re-
sources of the Hawaiian Islands. Bulletin of the United States Fish Com-
mission, vol. 23, for 1903, part 2.
Gitcurist, J. D. F.
1920. Ecdysis in a teleostean fish, Agriopus. The Quarterly Journal of Micrescopical
Science, vol. 64, pp. 575-587.
GILL, THEODORE
1905. Note on the genera of Synanceine and Pelorine fishes. Proceedings of the United
States National Museum, vol. 28, no. 1394, pp. 221-225, 1 fig.
GosLINE, WILLIAM A.
1965. Vertical zonation of inshore fishes in the upper water layers of the Hawaiian
Islands. Ecology, vol. 46, no. 6, pp. 823-831.
GosLINE, WILLIAM A., and VERNON E. Brock
1960. Handbook of Hawaiian fishes. University of Hawaii Press, 372 pp., 277 text figs.
GUNTHER, ALBERT
1860. Catalogue of the acanthopterygian fishes in the collection of the British Museum,
vol. 2, xxi-+ 548 pp.
1891. Description of a remarkable fish from Mauritius, belonging to the genus Scorpaena.
Proceedings of the Zoological Society of London, year 1891, pp. 482-483, pl. 39.
Herre, ALBERT W. C. T.
1952. A review of the scorpaenoid fishes of the Philippines and adjacent seas. The
Philippine Journal of Science, vol. 80, no. 4, pp. 381-482.
Hirosaki, YOSHITSUGU
1957. Observations on ecdysis in Japanese fishes (Preliminary Report). Journal of
the Faculty of Science of Hokkaido University, ser. VI, Zoology, vol. 13, nos.
1-4, pp. 178-179.
1971a. Uo no dappi (Ecdysis in fishes). Asahi-Lalousse, Shukan Sekaiddbutsu Hyakka,
no. 4, pp. 4-5. (In Japanese).
1971b. Dappi o tsuzukeru figawarina kaisuigyo (A strange marine fish which continues
shedding). Fish Magazine, Tokyo, vol. 7, no. 5, pp. 26-27, plus front cover.
(In Japanese).
1971c. Kaichiiddbutsu no fushigi (Wonders of marine animals). Jz Sea Fantasy—the
world under the ocean surface. Tokyo, pp. 153-174. (In Japanese).
Jorpan, Davin S., and Barton W. EVERMANN
1926. A check list of the fishes of Hawaii. Journal of the Pan-Pacific Research Insti-
tution, vol. 1, no. 1, pp. 3-15.
Jorpan, Davi S., and ALVIN SEALE
1906. The fishes of Samoa. . . Bulletin of the Bureau of Fisheries, vol. 25 (for 1905),
pp. 173-455, 111 text figs., pls. 38-53.
KAmoHarA, TOSHIJI
1942. Rare fishes of the Provinces of Kishu and Tosa [= Wakayama and Kochi
Prefectures]. Dobutsugaku Zassi [Zoological Magazine], Tokyo, vol. 54, no.
1, pp. 25-28. (In Japanese).

310 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

1950. Description of the fishes from the Provinces of Tosa and Kishi, Japan. Kochi,
368 pp., 220 text figs. (In Japanese, with English title).
1959. New records of fishes from Kochi Prefecture, Japan. Reports of the USA
Marine Biological Station, vol. 6, no. 2, pp. 1-8, 3 text figs.
1964. Revised catalogue of fishes of Kochi Prefecture, Japan. Reports of the USA
Marine Biological Station, vol. 11, no. 1, pp. 1-99, 63 text figs.
KotrrHaus, ADOLF
1966. Fischforschung im Indischen Ozean ‘“Meteor’-Expedition 1964/65. Umschau in
Wissenschaft und Technik, jarhgang heft 1966, pp. 118-123, 8 figs.
MAstuBarA, KtyoMatTsu
1943. Studies on the scorpaenoid fishes of Japan. The Transactions of the Sigenkagaku
Kenkyusyo, no. 1, 486 pp., 4 pls.
Munro, IAn F. R.
1955. The fishes of New Guinea. Department of Agriculture, Stock and Fisheries,
Port Morsby, XX XVII -+ 651 pp., 6 color pls., 78 pls., 23 text figs.
PALMER, G.
1963. The scorpaenid fishes of the genera Rhinopias Gill and Peloropsis Gilbert. Annals
and Magazine of Natural History, ser. 13, vol. 6, pp. 701-704, pl. 20
REGAN, C. TATE
1908. Report on the marine fishes collected by Mr. J. Stanley Gardiner in the Indian
Ocean. The transactions of the Linnean Society of London, vol. 12, part 3, no.
14, pp. 217-255, pls. 23-32.
SmirTH, J. L. B.
1957. The fishes of the family Scorpaenidae in the western Indian Ocean, part 1. The
subfamily Scorpaeninae. Ichthyological Bulletin, Rhodes University, Grahams-
town, no. 4, pp. 49-72, pls. 1-4.
1966. Certain rare fishes from South Africa with other notes. Occasional Papers,
Department of Ichthyology, Rhodes University, Grahamstown, no. 7, pp. 65—
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TINKER, SPENCER W.
1944. Hawaiian fishes. Tongg publishing Co., Honolulu, 404 pp., text figs. 8 pls.
Weser, Max, and L. F. DE BEAUFORT
1962. The fishes of the Indo-Australian Archipelago, XI. Scleroparei, Hyostomides. . .
Xenopterygii. E. J. Brill, Leiden, 481 pp., 100 text figs.
WHITLEY, GIvBeErT P.
1954. More new fish names and records. The Australian Zoologist, vol. 12, part 1,
pp. 57-62, pl. 3.
1964. Camouflaged fish from New Caledonia. The Australian Naturalist, vol. 12, part 4,
p. 9, pl. II.
WICKLER, WOLFGANG, and CHRISTEL NOWAK
1969. Hautung und andere Verhaltensweisen von Taentanotus triacanthus, einem
verwandten der Skorpionfische. Natur und Museum, vol. 99, no. 10, pp. 441—
456, 9 figs.

OF THE

CALIFORNIA ACADEMY OF SCIENCES

FOURTH SERIES

Vol. XXXIX, No. 17, pp. 311-336; 6 figs. August 9, 1973

THE BRACHYURAN CRABS OF EASTER ISLAND'

By

John S. Garth

Allan Hancock Foundation
University of Southern California

INTRODUCTION

Easter Island or Rapa Nui is a volcanic island lying in the eastern South
Pacific at Latitude 27° 10’ S. and Longitude 109° 26’ W. (Cook Bay or Hanga-
Roa). Roughly triangular in shape, it measures approximately 11 miles along
its major dimension, from southwest to east, and approximately 8 miles along
each of its two shorter legs. Each corner is dominated by a volcanic crater or
shield, of which La Pérouse Mountain (Rana Hana Kana), near North Cape,
is highest (1767 feet). The principal anchorages are at Cook Bay (Hanga-Roa)
on the west side and La Pérouse Bay on the northeast side. Neither of these is
protected in all weather and a ship must be prepared to up-anchor on an hour’s
notice, a circumstance that accounts in part for the infrequency with which the
island has been visited by scientific parties. Of these only one is known to have
made significant collections of marine animals: the Albatross Eastern Pacific
Expedition of 1904-1905. According to Alexander Agassiz (1906, p. 56), expe-
dition leader, “Considerable collecting was done at Easter Island .... We col-
lected a number of shore fishes, and made a small collection of the littoral fauna.
The fishes have a decided Pacific look, and the few species of sea urchins . . .
have a wide distribution in the Pacific.”” The Brachyura of this expedition were
reported on by Rathbun (1907), who listed the following six species from Easter
Island:

1 Allan Hancock Foundation Contribution No. 345.

311

312 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

XANTHIDAE: Pseudoliomera remota (Rathbun)
Trapezia ferruginea Latreille.

GrapsIpAaE: Leptograpsus variegatus (Fabricius)
Pachygrapsus transversus (Gibbes)
Ptychognathus easteranus Rathbun
Plagusia dentipes (De Haan).

The Heyerdahl Expedition of 1955-1956 collected a single brachyuran on
Easter Island. Not hitherto reported, it is included herewith:

GRAPSIDAE: Geograpsus crinipes (Dana).

The Medical Expedition to Easter Island (METEI) visited the island in
1964-1965. Collections of brachyuran crabs were made from late December to
early February at the following localities: on the north side of the island at
Anakena Bay; on the west side of the island at Hanga-Roa, Hanga-piko, and
Mataveri (inland from Hanga-piko); on the south side of the island at Vaihu.
Most of the collecting was intertidal, but that at Anakena was in water of 20-
25 foot depth. Specimens were collected by rotenone poisoning at Vaihu, and
the stomach of a large starfish from an unknown locality yielded crabs of the
genus Trapezia, usually associated with corals. Except where noted, as in the
case of children and islanders, who provided native names for crabs, all col-
lecting was done by Ian E. Efford and Jack A. Mathias.

Fourteen species of brachyuran crabs are included in the METEI collection,
a number double that obtained by earlier expeditions. Among the 14 are all
previously collected species except Pseudoliomera remota, obtained by the
Albatross Expedition, and Geograpsus crinipes, obtained by the Heyerdahl Expe-
dition. A second specimen of Ptychognathus easteranus of the opposite sex
from the unique holotype was collected. The number of crab families known
from Easter Island was increased from two to four with the addition of the
Dromiidae and Portunidae; the number of genera was increased from seven to
twelve with the addition of Dromidia, Portunus, Carpilius, Chlorodiella, and
Cyclograpsus; and among the seven species new to the island is a second species
of Plagusia new to science.

The Expedicién Isla de Pascua (EIP) of the Instituto Central de Biologia,
Universidad de Concepcion, Chile, visited the island in August, 1972. Collec-
tions of brachyuran crabs were made intertidally at Anakena Bay, Hanga-Roa,
and Hotu-Iti by H. I. Moyano, at Tahai by T. Cekalovic, and by diving in 8-
10 meters by V. A. Gallardo. Specimens were sent to the author for identifi-
cation in November, 1972, by M. A. Retamal, who has kindly permitted in-
corporation of the new records in the present manuscript.

Included in the EIP collection are twelve species of brachyuran crabs. Among
them are five species also collected by the METEI and one species collected

Vol. XXXIX] GARTH: BRACHYURAN CRABS 313

by the Heyerdahl Expedition. The remaining six species include five widely
ranging Indo-west Pacific species not previously known from Easter Island
and one that appears to be new to science. The number of genera known from
Easter Island is increased from twelve to sixteen with the addition of Thalamita,
Etisus, Liomera (= Carpilodes), and Lophozozymus. Description of the new
species will appear subsequently in a Chilean journal, at Professor Retamal’s
request.

The astute carcinologist will note that two large groups of Brachyura, the
Oxystomata and the Oxyrhyncha, are lacking from Easter Island collections
made to date, the list being composed entirely of Dromiacea and Brachyrhyncha.
It may be confidently predicted that when present terrestrial and intertidal col-
lecting is extended to include the subtidal, representatives of these groups will
be added; for while a few oxystomatous and oxyrhynchous crabs occur in
weeds and among corals in the tropical Pacific, more are to be found by shallow
dredging.

HisTor1tcaL NOTE

Through the research of Dr. L. B. Holthuis of the Royal Netherlands Mu-
seum of Natural History, Leiden, I am able to mention what is probably the
oldest observation of crabs on Easter Island. It is found in the account of Don
Juan Hervé, first pilot or senior navigating officer of the San Lorenzo, one of
two vessels comprising the expedition to the islands under Don Felipe Gonzalez
in 1770 (Hervé, 1908, p. 122), as reprinted in the Works of the Hakluyt Society,
transcribed and edited by B. G. Corney. In the description of the small, rocky
islets at the southwest tip of the island, now known as Needle Rock or Motu-
kaokao and Flat Rock or Motu-nui, is the statement: “We passed on to the
outer one, where we succeeded in landing, and on which we found two large
masses of seaweed, many black flints, some sea urchins and small crabs, eggs of
seagulls and their fledglings . . .” It is tempting to speculate, in the light of
METEI findings, what these small crabs might have been.

ACKNOWLEDGMENTS

When preparing the account of the brachyuran decapod Crustacea of Chile
a decade and a half ago (Garth, 1957) it was realized that this work was in-
complete in that coverage of the Chilean Isla de Pascua, as Easter Island is
known to the Spanish-speaking world, was lacking. Since no new specimen
material was then available, as in the case of Juan Fernandez Island, it was
thought better to await the arrival of fresh collections than to redescribe the
older ones. Such an opportunity presented itself with the extensive collections
made by Ian E. Efford and Jack A. Mathias on the Medical Expedition to Easter

314 CALIFORNIA ACADEMY OF SCIENCES [Proc. 47TH SER.

Island, and to them the author is indebted for the privilege of extending his
knowledge of the Chilean fauna to this island outpost of the Pacific.

Inclusion in the present report of the single specimen of brachyuran crab col-
lected at Easter Island by the Heyerdahl Expedition of 1955-1956 was made
possible through the courtesy of Mr. Nils Knaben of the Oslo Museum, with
Dr. L. B. Holthuis of the Leiden Museum facilitating arrangements.

Inclusion in the present report of the Easter Island collections of the Insti-
tuto Central de Biologia, Universidad de Concepcion, Chile, was made possible
through the kindness of Prof. M. A. Retamal.

The writer is indebted to Mme. Daniéle Guinot of the Muséum National
d’Histoire Naturelle, Paris, for identifying the Chlorodiella species from Easter
Island, and to Dr. T. Sakai of Kamakura, Japan, for confirming the identity
of Plagusia dentipes from Easter Island with Japanese specimens. He wishes to
thank Dr. C. B. Goodhart of the Museum of Zoology of Cambridge University,
Cambridge (see Garth, 1971), for permission to examine specimens of Pseudo-
liomera remota and P. lata from the Indian Ocean, and Dr. L. B. Holthuis of
the Rijksmuseum van Natuurlijke Historie, Leiden, and Mr. J. Forest of the
Muséum National d’Histoire Naturelle, Paris, for permission to examine the
type-specimens of Plagusia species at their respective institutions.

Consultation of type-specimens in European museums was made possible by
a grant from the National Science Foundation (GB-3849). Illustration of the
new species of Plagusia was provided by a grant from the Research and Publi-
cation Fund (National Defense Education Act) of the University of Southern
California.

The sponge covering of Dromidia was identified by Dr. Gerald J. Bakus of
the University of Southern California. The coral hosts of Trapezia species were
identified by Dr. John W. Wells of Cornell University and the star fish from the
stomach of which Trapezia species were recovered was identified by Mr. James
Clark of Harvard University. Dr. F. A. Chace, Jr. of the U. S. National Mu-
seum reviewed the manuscript and made valuable suggestions.

ACCOUNT OF SPECIES

RESTRICTION OF SYNONYMIES. The synonymies that follow are restricted
to the original description, the first use of the name in its present combination,
a good illustration if not included in the above two, the first record from Easter
Island, and, in order to provide documentation for the zoogeographical dis-
cussion to follow, records from adjacent Pacific islands from which the species
might have migrated to Easter Island. In cases of involved synonymies, ref-
erence is made to one of the regional monographs, such as Alcock (1900), Ed-
mondson (1922, 1954, 1959, 1962), Forest & Guinot (1961), Ihle (1913),
Rathbun (1906, 1918, 1930), or Sakai (1936, 1939).

Vol. XXXIX] GARTH: BRACHYURAN CRABS 315

LIST OF SPECIES

Alba- Heyer-

tross dahl METEI

1904— 1955- 1964— EIP
1905 1956 1965 1972

DROMIACEA

DROMIIDAE
Dromidia unidentata (Ruppell). xX

BRACHYGNATHA-BRACHYRHYNCHA

PORTUNIDAE
Portunus pubescens (Dana) xX
Thalamita xX
XANTHIDAE
Car pilius convexus (¥Forskal) xX
Liomera rugata (Milne Edwards) x
Actaea parvula (Krauss) xX
Pseudoliomera remota (Rathbun) xX
Lophozozymus dodone (Herbst) x
Etisus electra (Herbst) xX
Chlorodiella cytherea (Dana) Xx xX
Trapezia areolata Dana x xX
Trapezia cymodoce (Herbst) x xX
Trapezia danai Ward (= maculata Dana) xX
Trapezia ferruginea Latreille xX xX
Species incertae sedis.” x xX
GRAPSIDAE
Geograpsus crinipes (Dana) X xX
Leptograpsus variegatus (Fabricius) xX xX x
Pachygrapsus transversus (Gibbes) xX xX
Ptychognathus easteranus Rathbun xX xX
Cyclograpsus longipes Stimpson x
Plagusia dentipes De Haan x X
Plagusia integripes Garth, new species xX
6 1 14 i?

2 One small xanthid obtained by the METEI is unreported pending further study.

316 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Section BRACHYURA
Subsection DROMIACEA
Superfamily DRomiIDEA
Family DRroMrIDAE
Dromidia unidentata unidentata (Rippell).

Dromia unidentata RUPPELL, 1830, p. 16, pl. 4, fig. 2; pl. 6, fig. 9 (Red Sea) ; AtcocK, 1900,

p. 139, literature; Cuiiton, 1911, p. 554 (Kermadec Islands).

Dromidia unidentata, KOSSMANN, 1880, p. 67; ILE, 1913, p. 31 (New Guinea), synonymy;

SAKAI, 1936, p. 13, pl. 6, fig. 2; text-figs. 2a, b (Japan).

MATERIAL EXAMINED. Anakena, Easter Island, METEI Station F 85, 20-
25 feet, 15 January 1965, 1 male, Ian E. Efford and Jack A. Mathias. The
crab was covered by a sponge identified as Hymeniacidon species by Dr. Gerald
J. Bakus.

MEASUREMENTS. Male specimen: length of carapace 13.4 mm., width
11.3 mm.

DISTRIBUTION. From Red Sea and East Africa to New Guinea and Japan;
Kermadec Islands and, as subspecies D. u. Aawaiiensis Edmondson (1922, p.
34), Hawaii.

Remarks. The Easter Island specimen allies itself with the widely distrib-
uted Indo-west Pacific subspecies, rather than with the Hawaiian subspecies
described by Edmondson (1922, p. 34). The only other South Pacific record
is from Meyer Island in the Kermadec Islands (Chilton, 1911), which lie north-
east of New Zealand in Latitude 30° S., a few degrees more southerly than
Easter Island. Other islands lying south of the Tropic of Capricorn and, to-
gether with Rapa Nui (Easter) and Sala y Gomez, completing an arc to San
Felix and San Ambrosio off the coast of Chile are Rapa, Morotiri (Bass), Pit-
cairn, Henderson, and Ducie.

Subsection BRACHYGNATHA
Superfamily BRACHYRHYNCHA
Family PoRTUNIDAE

Portunus pubescens (Dana).

Lupa pubescens DANA, 1852b, p. 274; 1855, pl. 16, fig. 9 (Sandwich Islands).

Portunus pubescens, RATHBUN, 1906, p. 870, pl. 14, fig. 1; Epamonpson, 1923, p. 22 (Palmyra) ;
STEPHENSON and CAMPBELL, 1959, p. 99, figs. 2C, 3C, pl. 1, fig. 3; pls. 4C, 5C, synonymy.

Neptunus pubescens, SAKAI, 1934, p. 303 (Japan).

Neptunus tomentosus HASWELL, 1882, p. 547 (Australia).

MATERIAL EXAMINED. Hanga-Roa, near camp, January 1965, 1 female, Ian
E. Efford and Jack A. Mathias.

Vol. XXXIX] GARTH: BRACHYURAN CRABS 317

MEASUREMENTS. Female specimen: length of carapace 28.0 mm., width
including lateral spines 44.6 mm., without spines 39.0 mm.

DisTRIBUTION. Australia, Japan, Hawaii, and Line Islands (Palmyra).
The early record from Port Jackson, New South Wales, is based on Stephenson
and Campbell’s synonymy of Neptunus tomentosus.

Remarks. The occurrence at Easter Island of Portunus pubescens, until
recently known only from the north Central Pacific and Japan, becomes more
plausible with the establishment of its identity with the Australian P. tomentosus
and its recognition as a southern hemisphere species, Easter Island being in the
Latitude of Brisbane. It is probable that P. pubescens occurs widely through-
out the South Pacific but has not been reported because of insufficient collecting.
Traces of reddish brown color remain on the metagastric and epibranchial regions
after three and a half years in alcohol.

Family XANTHIDAE

Carpilius convexus (Forskal).

Cancer convexus ForSKAL, 1775, p. 88 (Red Sea).

Carpilius convexus RUPPELL, 1830, p. 13, pl. 3, fig. 2; pl. 6, fig. 6 (Tahiti); Dana, 1852b,
p. 159; 1855, pl. 7, fig. 5 (Sandwich Islands); RAtusBun, 1907, p. 37 (Tuamotu) ;
Boone, 1934, p. 89, pls. 43-45, synonymy; BARNARD, 1950, p. 205 (Durban) ; EpmMonpson,
1962, p. 223, fig. lb; GarTH, 1965, p. 16, figs. 8 (male pleopods), 13 (Clipperton Island).
MATERIAL EXAMINED. Anakena, Station M 5, 1 meter, 23 December 1964,

2 females, collected by islanders. Mataveri, Station M 41, 21 January 1965,

1 female, Ian E. Efford and Jack A. Mathias.

MESUREMENTS. Specimens, all females, measured 62.8 X 85.4, 70.7 X 95.1,
and 73 X 96 mm. in length and width of carapace, respectively.

DisTRIBUTION. From the Red Sea and South Africa to Hawaii, Tahiti, and
Tuamotu. Clipperton Island.

Remarks. The color of the Easter Island specimens resembled nothing that
this writer has encountered previously with this or any other brachyuran spe-
cies. It was as if each specimen had been dipped, like an Easter egg, in a series
of dyes: orange, pink, and blue, while being held each time by a different ap-
pendage. The result was a piebald appearance of utmost irregularity that could
not be called a pattern. Suspecting some artifact in the preserving process, the
writer queried Tan E. Efford, but was assured that this was their natural color-
ing before preservation. The native name for this spectacular crab is Pikea
Tutuau.

Widely distributed throughout the Indo-west Pacific, Carpilius convexus
has now been reported twice from the eastern Pacific, once at Clipperton Island
and now at Easter Island, in the same Longitude, 109° W. Some 37° of Lat-

318 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

itude, or 2,200 nautical miles, separate the two localities. The genus occurs also
in the western Atlantic, but is absent from the American west coast.

Liomera rugata (Milne Edwards).

Carpilodes rugatus, A. Mi1nr Epwarps, 1865, p. 230, pl. 12, figs. 3, 3a, 3b; ORTMANN, 1893,

p. 468 (Tahiti) ; ODHNER, 1925, p. 20, pl. 1, fig. 16 (Gilbert Islands) ; EpDMonpson, 1962,

p. 249, fig. 9a, synonymy.

Carpilodes monticulosus, RATHBUN, 1906, p. 884 (Hawaii); 1907, p. 37 (Tuamotu). [Not

C. monticulosus A. Milne Edwards. |
Liomera rugatus, BARNARD, 1950, p. 237.

MATERIAL EXAMINED. Hotu-iti, 8 August 1972, one female, H. I. Moyano,
1 ie

MEASUREMENTS. Female specimen, length of carapace 12.2 mm., width
21.4 mm.

DISTRIBUTION. From Red Sea and Mauritius to Tahiti, Tuamotu, and
Hawaii.

Remarks. This common Indo-west Pacific species has been confused with
Liomera monticulosa (A. Milne Edwards) by such authorities as Borradaile
(1902) and Rathbun (1907), according to Odhner (1925), whose figure is the
basis for our identification.

Actaea parvula (Krauss).

Menippe parvulus Krauss, 1843, p. 34, pl. 2, figs. 2, a-c (Natal).

Actaea parvula, LENz, 1910, p. 549 (Madagascar) ; ODHNER, 1925, p. 51, pl. 3, fig. 13 (Galle,
Ceylon; Fiji; Marquesas); BarNnarp, 1950, p. 234, text-fig. 43, g, h (Delgoa Bay),
synonymy.

MATERIAL EXAMINED. Anakena Bay, 9 August 1972, 2 males, one ovigerous
female, H. I. Moyano, University of Concepcion, Chile.

MEASUREMENTS. Male specimens, 8.9 X 11.7 and 10.5 X 14.1 mm.; ovigerous
female specimen 11.5 X 15.5 mm. in length and breadth of carapace, respectively.

CoLoR IN ALCOHOL. Carapace yellow, with red-orange blotches on gastric and
hepatic regions (three blotches in all). Chelipeds and legs yellow, fingers white
with brown pigment visible beneath the surface. Hairs golden.

Remarks. Easter Island specimens were compared with a 9.3 X 12.5 mm.
female collected by the writer at Galle, Ceylon, in 1964, which had in turn been
compared with specimens in the British Museum (Natural History), also from
Galle, Ceylon, identified by T. Odhner. The salient, tridentate tooth on the
immovable finger of the chela is characteristic of the species.

Pseudoliomera remota (Rathbun).

Actaea remota RatHBUN, 1907, p. 43, pl. 1, fig. 9; pl. 7, fig. 1 (Easter Island); 1911, p. 217
(Salomon, Coetivy) ; ODHNER, 1925, p. 63 (Hilo, Hawaii) ; SaKat, 1939, p. 490, pl. 93,
fig. 4 (Tosa Bay) ; Gutnot, 1962, p. 237, figs. 8, 9 (Maldive Islands).

Vol. XXXIX] GARTH: BRACHYURAN CRABS 319

Actaea nana KLUNZINGER, 1913, p. 86 (Koseir, Red Sea).
Pseudoliomera remota, GUINOT, 1967, p. 561; 1969, p. 228.

PREVIOUS COLLECTION. Easter Island, shore, 20 December 1904, 1 male,
Albatross (Rathbun, 1907).

MATERIAL EXAMINED. None from among METEI collections. Through
the kindness of Dr. C. B. Goodhart of the University Museum of Zoology,
Cambridge, it was possible to examine the young female from Salomon collected
by the Sealark.

MEASUREMENTS. Young female: length of carapace 4.0 mm., width 4.6 mm.
The holotype male (USNM no. 32849) measures 6.0 mm. in length and 8.7 mm.
in width of carapace (Rathbun, 1907).

DISTRIBUTION. Red Sea and western Indian Ocean to Japan, Hawaii, and
Easter Island. The west Australian record (Guinot, 1962, fig. 9) is an error of
provenience resulting from a confusion of names between the Eastern Islands
of the Houtman Abrolhos group in the eastern Indian Ocean and Easter Island
(ile de Paques) in the eastern South Pacific. A Christmas Island also exists
in both eastern Indian and central Pacific oceans further to confound the
zoogeographer.

Remarks. As noted from the Indian Ocean specimen examined, Pseudo-
liomera remota is a patterned species with the inner protogastric areoles broadly
continuous with the front and the outer protogastric areoles discontinuous with
the inner orbital areoles. The fingers resemble those of Pseudoliomera lata
(Borradaile) from the Maldive Islands (see Garth, 1971), also seen at Cam-
bridge University (cf. Guinot, 1962, figs. 6 and 8). The synonymy of Actaea
nana Klunzinger (cf. Odhner, 1925) permits the inclusion of the Red Sea in
the already extended longitudinal range of the species.

Lophozozymus dodone (Herbst).

Cancer dodone Hersst, 1801, p. 37, pl. 52, fig. 5 (East Indies).
Atergatis elegans HELLER, 1862, p. 519; 1865, p. 7, pl. 1, fig. 3 (Tahiti).
Lophozozymus dodone, ALcock, 1898, p. 108 (Andaman); Batss, 1938, p. 39 (Fiji, Tahiti).

Forest and Guinot, 1961, p. 54, text-figs. 39a, b (Tahiti), synonymy.

MATERIAL EXAMINED. Anakena Bay, shore, 9 August 1972, 1 male, H. I.
Moyano, University of Concepcién Expedition.

MEASUREMENTS. Male specimen: length of carapace 6.1 mm., width 9.2 mm.

DISTRIBUTION. From South Africa to Hawaii and Tahiti.

Remarks. The Easter Island specimen was compared with a specimen from
Hawaii of like size and sex. The slight discrepancies noted are believed to lie
within the normal range of variation for this widely ranging Indo-west Pacific
species.

320 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Etisus electra (Herbst).

Cancer electra HErsst, 1801, p. 34, pl. 51, fig. 6 (East Indies).

Etisus rugosus JACQUINOT, 1852, pl. 4, fig. 2; 1853, p. 33 (Tuamotu).

Etisodes sculptilis A. MILNE Epwarps, 1873, p. 236, pl. 9, fig. 2 (New Caledonia).
Etisodes electra, Nos, 1907, p. 390 (Tuamotu) ; RATHBUN, 1907, p. 42 (Tuamotu, Gilbert

Islands) ; Forest and Gurnot, 1961, p. 89, figs. 82a, b (Tahiti), synonymy.

Etisus electra, BARNARD, 1950, p. 245, figs. 45a, b (South Africa); Gurnot, 1964, p. 59.
Etisus (Etisodes) electra, HoLTHUutIs, 1953, p. 21 (Marianas, Gilbert Islands).

MATERIAL EXAMINED. Anakena Bay, sand, 9 August 1972, 1 male, 1 young,
H. I. Moyano, University of Concepcion, Chile.

MEASUREMENTS. Male specimen, length of carapace 5.4 mm., width 6.8 mm.

DISTRIBUTION. From the Red Sea and Mozambique (Delgoa Bay) to Tahiti
and Tuamotu (Manga Reva).

Remarks. The Easter Island specimens compare with the figure of Etisodes
sculptilis A. Milne Edwards, a synonym of Etisus electra (Herbst), according
to Guinot (1964). The far-flung distribution of this small xanthid makes it
a likely colonizer of island outposts like Easter.

Chlorodiella cytherea (Dana).
Chlorodius cytherea DANA, 1852a, p. 79; 1852b, p. 213; 1855, pl. 12, figs. 2a-c (Tuamotu and

Hawaii).

Chlorodiella cytherea, HottHuts, 1953, p. 14 (Marianas and Gilbert islands); Forest and

Gu1not, 1961, p. 95, figs. 90-92, 98a, b (Tahiti), synonymy.

MATERIAL EXAMINED. Hanga-piko, tide pool, Station M 10, 31 December
1964, 1 ovigerous female, Ian E. Efford and Jack A. Mathias. Anakena Bay,
on the beach, 9 August 1972, 1 young male, H. I. Moyano, EIP.

MEASUREMENTS. Female specimen: length of carapace 6.1 mm., width 9.6
mm.

DISTRIBUTION. From the Red Sea and Madagascar to Hawaii, Tahiti, and
Tuamotu.

REMARKS. With the recognition by Holthuis (1953) of Chlorodiella cytherea
among Pacific coral atoll collections and its reestablishment by Forest and
Guinot (1961) upon firm morphological characters, it becomes possible to dis-
tinguish four common C/lorodiella species in the Indo-west Pacific, the others
being C. laevissima (Dana), with which the present species was formerly con-
fused, C. nigra (Forskal), and C. barbata (Borradaile). The Easter Island
specimen was seen and its identification confirmed by Mme. D. Guinot of the
Paris Museum.

Trapezia areolata Dana.

Trapezia areolata DANA, 1852a, p. 83; 1852b, p. 259; 1855, pl. 15, fig. 8a (Tahiti) ; Forest and
Guinot, 1961, p. 135, fig. 133 (Hikueru).

Trapezia ferruginea areolata, ORTMANN, 1897, p. 206, synonymy.

Trapezia reticulata STIMPSON, 1858, p. 37.

Vol. XXXIX] GARTH: BRACHYURAN CRABS 321

MATERIAL EXAMINED. Hanga-piko, 2 February 1965, 2 males, 2 ovigerous
females, collected by Norma (METEI). No station or date, 2 males, from the
stomach of a large starfish collected by Ian E. Efford and Jack A. Mathias and
identified as Leiaster leachii by James Clark of Harvard University. SCUBA
diving, coral reef, 8-10 meters, 8-9 August 1972, 1 male, V. A. Gallardo, EIP.

MEASUREMENTS. Males measure 9.3 X 10.7, 9.5 X 10.8, 9.6 X 10.9, and
12.1 X 13.7 mm., ovigerous females 9.8 X 12.2 and 12.1 X 15.0 mm. in length
and width, respectively.

DISTRIBUTION. From Ceylon and the Nicobar Islands to Tahiti and Tuamotu
(Hikueru = Melville Island).

Remarks. The large starfish from whose stomach the two males were col-
lected, along with four specimens of Trapezia ferruginea Latreille, was presum-
ably feeding on the corals with which the Trapezia species are invariably asso-
ciated throughout their extensive range.

Trapezia cymodoce (Herbst).

Cancer cymodoce Hersst, 1801, p. 22, pl. 51, fig. 5 (East Indies).

orf

Trapesia cymodoce, GERSTAECKER, 1857, p. 125; ORTMANN, 1897, p. 203, synonymy; EpMonD-
son, 1923, p. 20 (Palmyra).

Grapsillus cymodoce, RATHBUN, 1906, p. 685, pl. 11, fig. 6 (Hawaii).

Trapesia hirtipes JacQuiNoT and Lucas, 1853, p. 44, pl. 4, fig. 14 (Marquesas).

MATERIAL EXAMINED. Hanga-piko, 2 February 1965, 1 ovigerous female
without chelipeds, collected by Norma (METEI).

MEASUREMENTS. Ovigerous female: length of carapace 8.5 mm., width
10.8 mm.

DISTRIBUTION. From the Red Sea and Dar-es-Salaam to Tahiti and the
Marquesas. Hawaii and the Line Islands (Palmyra).

Remarks. Although no mention is made of an invertebrate host, the Tra-
pezia species are customarily collected by cracking large heads of Pocillopora
coral, of which three species, P. damicornis, P. danae, and P. diomedae, occur at
Easter Island, according to Dr. John W. Wells of Cornell University. Trapeszia
cymodoce is most readily distinguished from T. ferruginea by the fine woolly
hair on the outer side of the palm of the cheliped. In the absence of chelipeds,
reliance may be placed in the lateral teeth of the carapace, which are pointed even
in grown specimens, according to Ortmann (1897).

Trapezia danai Ward.

Trapezia maculata Dana, 1852b, p. 256 (part); 1855, pl. 15, fig. 4d (mot 4a-c); Stmrpson,
1860, p. 219 (Socorro Island). [Not Grapsilius maculatus MacLeay.]

Trapesia danae Warp, 1939, p. 13, figs. 17, 18. [Name substituted for T. maculata Dana, pre-
occupied by 7. maculata (MacLeay).]

Trapesia aff. danai, SERENE, 1969, p. 136, figs. 14A, 14B, 15, 16, 21, 22, 24.

322 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

MATERIAL EXAMINED. Easter Island, 8-10 meters, SCUBA diving, 9-10
August 1972, one female, V. A. Gallardo, EIP.

MEASUREMENTS. Female specimen, length of carapace 8.5 mm., width
10.8 mm.

DISTRIBUTION. Western Pacific at Samoa, Tahiti, and Hawaii; eastern
Pacific doubtfully at Socorro Island; specimens sent to Stimpson by John Xantus
probably were from Hawaii also.

ReMaRKS. Since Dana’s material probably contained more than one species
(Serene, 1969), it is with some hesitation that the name now applied to one of
them is used here, as it is by no means certain that the Easter Island specimen,
a female, is identical with either the Hawaiian or Tahitian specimens figured
by Dana (1855). For the present, it can only be said that Easter Island supports
a finely spotted species of Tvapezia resembling both T. danai Ward and T. wardi
Seréne, yet differing in some particulars from each of them. A series of speci-
mens, including males, will be needed to resolve the question.

Trapezia ferruginea Latreille.

Trapezia ferruginea LATREILLE, 1825, p. 695 (Red Sea) ; ORTMANN, 1897, p. 205, synonymy.
Trapezia cymodoce ferruginea, RATHBUN, 1907, p. 58 (Easter Island) ; 1930, p. 557, pl. 228,
figs. 1, 2 (Clarion Island); Boonr, 1927, p. 240, text-fig. 88 (Galapagos Islands) ;

HErRTLEIN and Emerson, 1957, p. 5 (Clipperton Island).

Trapezia miniata JAcQuINOT and Lucas, 1853, p. 43, pl. 4, fig. 10 (Marquesas).

PREVIOUS COLLECTION. Easter Island, shore, 20 December 1904, 1 male, 1
ovigerous female, A/batross (Rathbun, 1907).

MATERIAL EXAMINED. Easter Island, without precise locality or date, 3
males, 1 ovigerous female, from the stomach of a large starfish collected by
Tan E. Efford and Jack A. Mathias and identified as Leiaster leachii by James
Clark:

MEASUREMENTS. Males measure from 7.5 X 8.4 to 12.3 X 14.4 mm., oviger-
ous female 8.7 X 11.4 mm. in length and width, respectively.

DIstTRIBUTION. From the Red Sea and Zanzibar to Hawaii and the Marquesas
in the western Pacific; Clarion, Clipperton, Easter, and Galapagos islands in
the eastern Pacific; from the Gulf of California to Colombia on the American
mainland.

REMARKS. Tvrapezia ferruginea is one of two Indo-Pacific species of Tra-
pezia known to occur on the west coast of tropical America, 7. digitalis Latreille
being the other (Rathbun, 1930). Both are inhabitants of the Pocillopora coral
colony but, whereas 7. ferruginea is bright red, T. digitalis is a somber brown.
The latter species was not found among METEI collections, nor has it been
reported from Easter Island. Rathbun, (1907), who considered T. ferruginea
a subspecies of 7. cymodoce (Herbst), noted that the chelipeds of Easter Island
specimens were covered with fine spots.

Vol. XXXIX] GARTH: BRACHYURAN CRABS 323

Family GRAPSIDAE
Geograpsus crinipes (Dana).

Grapsus crinipes DANA, 1851, p. 249; 1852b, p. 341; 1855, pl. 21, figs. 6a—d (Sandwich Islands).
Geograpsus crinipes, ORTMANN, 1894, p. 706 (Samoa); De Man, 1895, p. 83, pl. 28, figs.
17a-c; EpMoNpDSON, 1923, p. 10 (Fanning, Palmyra) ; 1959, p. 162, fig. 4a; SAKAI, 1939,

p. 652, pl. 107, fig. 2 (Japan), synonymy.

MATERIAL EXAMINED. Poike, eastern peninsula of Easter Island, in a hole at
an altitude of 250 m., 24 February 1956, 1 male, Thor Heyerdahl, collector
(Oslo Museum). Tahai, 4 August 1972, T. Cekalovic, EIP.

MEASUREMENTS. Male specimens: 38.7 X 45.1 and 47 X 55 mm., in length
and breadth of carapace, respectively.

DisTRIBUTION. Throughout the Indo-west Pacific from the Red Sea and
Zanzibar to Japan, Samoa, Hawaii, and the Line Islands (Fanning, Palmyra).

Remarks. The male specimen collected by the Heyerdahl Expedition lived
for 19 days, or until 14 March 1956. It is not known whether the hole in which
it was found contained water, or if so, whether this was fresh or brackish. The
altitude at which the Easter Island specimen was collected serves to underscore
the terrestriality of the species. In the Marshall Islands, which are low islands,
Geograpsus crinipes occurs among litter on the forest floor and is seldom seen
by day except when accidentally disturbed.

The Oslo Museum specimen was identified by Dr. L. B. Holthuis of the
Leiden Museum, who kindly forwarded the essential information for inclusion.

Leptograpsus variegatus (Fabricius).

Cancer variegatus FABRIctus, 1793, p. 450.
Leptograpsus variegatus, MILNE Epwarps, 1853, p. 172; RATHBUN, 1907, p. 29 (Easter Island) ;

1918, p. 234, pl. 36; Gartu, 1957, p. 94 (synonymy).

Leptograpsus ansoni MILNE Epwarps, 1853, p. 171 (Juan Fernandez Island).

PREVIOUS COLLECTIONS. Easter Island, shore, 16, 20 December 1904, 3
males, 5 females. La Pérouse Bay, 17 December 1904, 7 males, 5 females, Al-
batross (Rathbun, 1907). The single male reported by Rathbun (1907) with
the date of 21 December 1899, is not presently on record at the U. S. National
Museum and may have been part of a collection not involving the Albatross,
which on the date in question was at sea between Fiji and Funafuti in the Ellice
Islands on an earlier cruise that did not stop at Easter Island (F. A. Chace,
Jr., personal communication).

MATERIAL EXAMINED. Near Hanga-Roa, Station F 42, 25-26 December 1964,
2 males, 1 female, collected by children. Vaihu, Station F 94, 25 January 1965,
rotenone collection in 6 to 10 feet of water, 1 male, soft shell, Ian E. Efford and
Jack A. Mathias. Hanga-Roa, 5 February 1965, 1 female, without chelipeds,
Ian E. Efford and Jack A. Mathias. Hanga-Roa Tai, 5 August 1972, 1 male,
H. I. Moyano.

324 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

MEASUREMENTS. Males measure from 21.8 X 24.5 to 43.5 X 50.5 mm.,
ovigerous female 30.1 * 34.2 mm., non-ovigerous female 44.2 X 49.8 mm. in
length and width of carapace, respectively.

DISTRIBUTION. Australia and New Zealand in the western Pacific; Easter
Island and Juan Fernandez Island in the eastern Pacific; Peru and Chile on the
west coast of South America.

Remarks. In sub-tropical southern hemisphere areas from which Grapsus
grapsus (Linnaeus) is absent, its dominance of the upper intertidal of rocky
shores is assumed by Leptograpsus variegatus. It is therefore probable that the
crabs observed in 1770 by Don Juan Hervé (1908, p. 122) and mentioned in the
introduction of this paper were of this species, although the fact that he re-
ferred to them as small crabs might indicate Pachygrapsus transversus (Gibbes)
instead. Unlike Grapsus grapsus, which keeps to the supra-tidal level, Lepto-
grapsus variegatus was reported by Lund University Chile Expedition collectors
both in rocky crevices around high water level and in quiet water between
boulders in the lower part of the littoral (Garth, 1957). Its presence in from
6 to 10 feet of water at Easter Island would support the latter observation.

Pachygrapsus transversus (Gibbes).

Grapsus transversus GIBBES, 1850, p. 181.
Pachygrapsus transversus, GIBBES, 1850, p. 182; RATHBUN, 1902, p. 279 (Galapagos) ; 1907,
p. 29 (Easter Island) ; 1918, p. 244, pl. 61, figs. 2, 3, synonymy.

PREVIOUS COLLECTION. Easter Island, shore, 16 December 1904, 1 female,
Albatross (Rathbun, 1907).

MATERIAL EXAMINED. Hanga-Roa, 23 December 1964, 1 male, Ian E. Efford
and Jack A. Mathias. Hanga-piko, tide pool, Station M 10, 31 December 1964,
2 males, Ian E. Efford and Jack A. Mathias. Vaihu, tide pool, 3 January 1965,
3 males, 2 females, Ian FE. Efford and Jack A. Mathias.

MEASUREMENTS. Males measure from 5.2 X 6.3 to 11.8 X 14.5 mm., females
5.2 X6.6 and 6.0 X 7.7 mm. in length and width of carapace, respectively.

DISTRIBUTION. West coast of America from Gulf of California to Peru;
Galapagos Islands; Easter Island. East coast of America from Florida to Ur-
uguay; Bermuda. West coast of Africa from Mediterranean to northern Angola;
Cape Verde Islands.

REMARKS. Since the Indo-west Pacific species of Pachygrapsus, P. planifrons
De Man and P. minutus A. Milne Edwards, were encountered by the Scripps
International Geophysical Year (IGY) Expedition at Clipperton Island (Garth,
1965), it was with interest that the writer verified from freshly collected METEI
material the Easter Island record (Rathbun, 1907) of the amphi-American
species, P. transversus (Gibbes). Thus, in the eastern North Pacific, as at
Clipperton and Clarion islands, it is the western Pacific Pachygrapsus species

Vol. XXXTX] GARTH: BRACHYURAN CRABS 325

that have gained foothold; in the eastern South Pacific, as at Galapagos and
Easter islands, it is a New World Pachygrapsus species that has established itself.

Ptychognathus easteranus Rathbun.
Ptychognathus easterana RATHBUN, 1907, p. 31, pl. 2, fig. 4; pl. 7, figs. 4, 4a (Easter Island).

PREVIOUS COLLECTION. Easter Island, shore, 20 December 1904, 1 male,
Albatross (Rathbun, 1907).

MATERIAL EXAMINED. Hanga-piko, tide pool, Station M 10, 31 December
1964, 1 young female, Ian E. Efford and Jack A. Mathias.

MEASUREMENTS. Young female: length of carapace 4.2 mm., width 5.0 mm.
The male holotype (USNM No. 32845) measured 10.6 mm. in length and 12.7
mm. in width of carapace (Rathbun, 1907).

DISTRIBUTION. Known only from Easter Island, where it was first collected
by the U. S. Fish Commission Steamer Albatross in 1904. The genus ranges
throughout the Indo-west Pacific.

Remarks. The single young female collected by the METEI differs from
the male holotype in being more straight-sided, thereby increasing its resem-
blance to Ptychognathus polleni De Man (1895, p. 94, pl. 28, fig. 20a), and
would not have been recognized as the same species as the male except for the
external maxilliped, which is identical with the figure of Rathbun (1907).

Cyclograpsus longipes Stimpson.

Cyclograpsus longipes Stimpson, 1858, p. 105 (Bonin Islands); De Man, 1896, p. 355, pl.
32, figs. 43a-c (Atjeh) ; RATHBUN, 1907, p. 36 (Tahiti, Tuamotu) ; SaKat, 1939, p. 690,
synonymy; CAMPBELL and GRIFFIN, 1966, p. 139, key.

MATERIAL EXAMINED. Shore above Vaihu, 25 January 1965, 2 males, 1
female, Ian E. Efford and Jack A. Mathias.

MEASUREMENTS. Male specimens: 4.6 X 5.7 and 5.4 X 7.0 mm.; female
specimen: 7.0 X 8.8 mm.

DistRIBUTION. Sumatra (Atjeh = Achin), Bonin Islands, Fiji Islands,
Tahiti, and Tuamotu.

Remarks. According to Campbell and Griffin (1966), Cyclograpsus longipes
is distinguished from other Cyclograpsus species with entire anterolateral margins
by having the lateral margins of the carapace straight and markedly divergent
posteriorly, the epigastric lobes prominent, and the suborbital ridge interrupted
two or three times. The Easter Island female differs from the two males in
lacking the felted patch on the propodus of the second pair of walking legs and
in having the suborbital ridge entire. In this respect it agrees with a larger
series of eight males and three females from the Marshall Islands in the collec-
tions of the Allan Hancock Foundation.

326 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Plagusia dentipes De Haan.
Plagusia dentipes DE HAAN, 1835, p. 58, pl. 8, fig. 1 (Japan); RATHBUN, 1907, p. 36 (Easter

Island) ; SAKAI, 1939, p. 79, fig. 3, synonymy.

PREVIOUS COLLECTION. Easter Island, shore, 21 December 1904, 2 males,
1 female, Albatross (Rathbun, 1907).

MATERIAL EXAMINED. Easter Island, near camp, January 1965, 1 male, 1
female, Ilan E. Efford and Jack A. Mathias.

MEASUREMENTS. Male specimen: length of carapace 38.5 mm., width
41.6 mm.; female specimen: length 34.4 mm., width 36.7 mm.

DISTRIBUTION. Japan and Formosa. Easter Island. Any continuity be-
tween the two regions must be assumed, as records from intermediate localities
are lacking.

Remarks. The Easter Island specimens, while large, are not as large as
a male from Simoda, length of carapace 44 mm., width 59.5 mm. (Sakai, 1939).
Easter Island specimens were seen by Dr. Sakai, who confirmed their identity
with Japanese specimens. The writer also had the privilege of examining two
male and one female syntypes of von Siebold’s collecting in the collections of the
Leiden Museum.

Plagusia integripes Garth, new species.
(Figures 1-6.)

Type. Female holotype (AHF No. 6511), from shallow water off Hanga-
Roa, Easter Island, 2 February 1965, Ian E. Efford and Jack A. Mathias, col-
lectors (METEI).

MEASUREMENTS. Female holotype: length of carapace 30.2 mm., width of
carapace 31.1 mm., of front 6.2 mm., between inner orbital spines 12.2 mm.,
between exorbital spines 19.7 mm., length of chela 15.0 mm., of dactyl 8.4 mm.

Diacnosis. Carapace tuberculate; three teeth behind external orbital angle.
Coxae of walking legs entire: meri smooth anteriorly, a single spine subter-
minally. Exognath of outer maxilliped without a flagellum.

DESCRIPTION. Carapace roughened everywhere by low scabrous tubercles
arranged in transverse rows and becoming more prominent on the slopes of the
major elevations; intervening depressions free of tubercles but filled with a
tomentum of short, stout, hooked hairs directed forward and inward. Larger
tubercles distributed as follows: two pairs on the front, forming a square; one
behind the orbit, one outer branchial, and one post lateral, each with one or more
subordinate tubercles. Front with about eight tubercles on each side arranged
in a forward-curving arc, the anterior three independent, the posterior five co-
alesced; a blunt tooth on outer slope of inner orbital lobe. Anterolateral border
with four strong teeth, including the exorbital tooth; teeth acute, directed for-
ward, upward, and slightly inward, separated by broad U-shaped sinuses, and
decreasing in size from first to last; last tooth considerably smaller than the

Vol. XXXTX] GARTH: BRACHYURAN CRABS 327

oy ye? egiee ae

eaeeronmeane ee pi

10mm

10mm

Ficure 1. Plagusia integripes Garth, new species, female holotype (AHF No. 6511), dorsal
view. FicurE 2. Same, outer view of left cheliped. Ficure 3. Same, frontal view of orbital
region. FicuRE 4. Same, left outer maxilliped. Ficure 5. Same, abdomen, restored. FIGURE
6. Same, coxa of right third leg. Drawings by Carl Petterson.

preceding tooth. Lower orbital margin continuous with epistome and similarly
denticulate; epistome not readily divisible into lobes. Merus of outer maxilliped
with a groove parallel to obliquely truncated anteroexternal margin; exognath
lacking a flagellum.

Chelipeds of the female (the male is not known) moderately robust, equal,

328 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

tuberculate, and hairy; tubercles forming a weak crest backed by feathered
setae along inner margins of merus, carpus, and manus, irregularly arranged
on upper surface of carpus and manus, and forming rows along ridges leading
to the dactyls on outer surface of palm; dactyls basally tuberculate, movable
finger grooved, tips hollowed and meeting with a slight gape; hooked hairs in
depressions of carpus and manus.

Merus of walking legs with anterior crest entire, terminating in an acute
spine or tubercle, fringed with plumose setae, and paralleled by a superior row
of low tubercles; carpus and propodus similarly crested and fringed but lacking
subterminal spine; dactylus short, curved, a double row of bristles internally,
fringing hairs externally. Coxal lobes entire.

Abdomen of unique female holotype crushed, but clearly showing the 7-seg-
mented arrangement as restored in figure 5.

RemMArKs. The new species from Easter Island allies itself with Plagusia
depressa (Fabricius), P. glabra Dana, and P. speciosa Dana in having the meri
of the walking legs smooth anteriorly and culminating in a single spine. Like
them also it lacks the flagellum of the exognath of the third maxilliped found
in Plagusia capensis De Haan and P. dentipes De Haan but in its place carries
a tuft of setae. It may be distinguished from Plagusia depressa in all its forms
by the denticulate lower margin of the orbit and inner margins of the merus,
carpus, and propodus of the cheliped, the corresponding surfaces in that species
being entire; by the flatter, more squarish carapace with clusters of sharp tu-
bercles separated by patches of hooked setae, rather than low squamiform tu-
bercles fringed with soft hair; and by the more robust chelae, at least in the
female sex, the male of the species being unknown; and from P. d. depressa by
the entire coxa of the walking legs (fig. 6), that of the nominate form being
bidentate (cf. Rathbun, 1918, text-fig. 154a).

The 2 holotype from Easter Island has been compared with each of the
following:

TYPE MATERIAL.

Plagusia chabrus (Linnaeus), 2 neotype from Cape of Good Hope, H. B. van Horstok;
Plagusia capensis De Haan, @ lectotype from Cape of Good Hope, H. B. van Horstok;
the above one and the same specimen in the Leiden Museum. (cf. Griffin, 1968)

Plagusia gaimardi Milne Edwards, 6 type from Tongatabu, Quoy and Gaimard;

Plagusia tomentosa Milne Edwards, 34 and 19 syntypes, Cape of Good Hope, attributed
to Latreille; 19 syntype, Cape of Good Hope, Delalande; both in the Paris Museum.
[The above three probably synonyms of Plagusia chabrus. |

Plagusia dentipes De Haan, 2¢ and 1Q syntypes, Japan, P. F. von Siebold, in the Leiden
Museum.

OTHER MATERIAL.

Plagusia capensis De Haan, 4 and @ from Long Reef, Callaroy, New South Wales;
Plagusia dentipes De Haan, é and @ from Easter Island;
Plagusia depressa (Fabricius), ¢ and 2 from Cubagua Island, Venezuela;

Vol. XXXIX] GARTH: BRACHYURAN CRABS 329

Plagusia depressa immaculata Lamarck, ¢ and @ from Cocos Island, Costa Rica;

Plagusia depressa tuberculata Lamarck, @ from IJIheya Shima, Ryukyu Islands;

Plagusia speciosa Dana, 2 from Apra Harbor, Guam; 2 from Clipperton Island. The above
are in the collections of the Allan Hancock Foundation.

Plagusia glabra Dana, 6 and 9 from Cogee, near Sydney, New South Wales. The above
courtesy of The Australian Museum, Sydney.

ZOOGEOGRAPHY

The study of the Brachyura of the METEI clearly establishes Easter Island
as the most easterly outpost of the Indo-west Pacific marine fauna in the South
Pacific. As such it is comparable to Clipperton Island, an atoll at the same
Longitude in the North Pacific, a relationship developed through the study of
the Brachyura of the Scripps IGY Expedition (Garth, 1965).

Easter Island species with ranges extending westward to the western margins
of the Indian Ocean are the following:

Dromidia unidentata (Ruppell) Red Sea, East Africa
Car pilius convexus (Forskal) Red Sea, South Africa
Liomera rugata (Milne Edwards) Red Sea, Mauritius
Actaea parvula ( Krauss) Red Sea, Natal
Lophozozymus dodone (Herbst) East Africa, Mauritius
Etisus electra (Herbst) Red Sea, Mozambique
Chlorodiella cytherea (Dana) Red Sea, Madagascar
Trapezia cymodoce (Herbst) Red Sea, Dar-es-Salaam
Trapezia ferruginea Latreille Red Sea, Zanzibar
Geograpsus crinipes (Dana) Red Sea, Madagascar.

Species with range extending westward to the Central Indian Ocean are
the following:

Pseudoliomera remota Rathbun Coetivy, Salomon
Trapezia areolata Dana Ceylon, Nicobar Islands
Trapezia danai Ward.

Species with ranges extending westward to the western margins of the Pacific
Ocean are the following:

Portunus pubescens (Dana) Australia, Japan
Thalamita species

Leptograpsus variegatus (Fabricius) Australia, New Zealand
Cyclograpsus longipes Stimpson Sumatra, Bonin Islands
Plagusia dentipes De Haan Japan, Formosa.

Species apparently endemic to Easter Island but representing Indo-west
Pacific or Pan-tropical genera are the following:

Ptychognathus easteranus Rathbun Plagusia integripes Garth, new species

330 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Species with ranges extending eastward to the shores of the Americas are
the following:

Trapezia ferruginea Latreille Galapagos Islands, Bay of Panama
Leptograpsus variegatus (Fabricius) Juan Fernandez Island, Chile
Pachygrapsus transversus (Gibbes) Galapagos Islands, Peru

It will be noted that the largest single segment are those that have negoti-
ated the distance from the Red Sea to Easter Island, representing 210° of Longi-
tude, or three-fifths of the circumference of the globe; also that the second
largest segment are those that have negotiated the distance from Australia to
Easter Island, representing 95° of Longitude, or one quarter of the circumference
of the globe. On the other hand, only three species have successfully negotiated
the distance from Chile to Easter Island, representing but 40° of Longitude,
or one-ninth the circumference of the globe, and of these only one species, Pachy-
grapsus transversus (Gibbes), appears to have made the crossing from east to
west, since it alone has incontestible New World affinities. That the migration
has proceeded predominantly in an easterly direction is apparent by the in-
creasing impoverishment of Pacific islands in Asiatic species from west to east,
without a corresponding enrichment in American species. In the case of the
Pachygrapsus species, which are susceptible to transport by ships’ hulls and
condensors, chance has determined the arrival and survival of P. transversus
at Galapagos and Easter Island, as also of the western Pacific P. planifrons
De Man and P. minutus A. Milne Edwards at Clarion and Clipperton in the
northern hemisphere (Garth, 1965), again underscoring the randomness of in-
sular dispersal.

The nearest islands to the west from which brachyuran species may have
reached Easter Island are Tahiti and Tuamotu. Fortunately, the fauna of
these islands is comparatively well known through the recent monograph of
Forest and Guinot (1961).

Species common to islands immediately to the west of Easter Island are the
following:

Car pilius convexus (Forskal) Tahiti, Tuamotu

Thalamita

Liomera rugata (Milne Edwards) Tahiti, Tuamotu (Fakarava)
Actaea parvula (Krauss) Marquesas

Lophozozymus dodone (Herbst) Tahiti

Etisus electra (Herbst) Tahiti, Tuamotu (Manga Reva)
Chlorodiella cytherea (Dana) Tahiti, Tuamotu

Trapezia areolata Dana Tuamotu (Hikueru = Melville)
Trapezia cymdoce (Herbst) Tahiti, Marquesas

Trapezia ferruginea Latreille Tahiti, Marquesas

Cyclograpsus longipes Stimpson Tahiti, Tuamotu.

Vol. XXXIX] GARTH: BRACHYURAN CRABS 331

There remain to be considered three species known only from localities
remote from Easter Island, rather than from adjacent islands. These are Dro-
midia unidentata (Ruppell), known in the Pacific from Japan, Hawaii, and
the Kermadec Islands; Portunus pubescens (Dana), known from Japan, Hawaii,
the Line Islands, and Australia; and Plagusia dentipes De Haan, known from
Japan and Formosa. The Marquesas Islands, mentioned above, suggest a pos-
sible migration route, since they lie northwest of Easter Island along an axis
which, if projected, leads through the Line Islands (Christmas, Fanning, Wash-
ington, and Palmyra) to Johnston and Midway islands, west of Hawaii. Such
a direct dispersal route seems highly improbable, however, since it cuts across
major ocean currents. The writer is inclined to favor, at least for the first two
species mentioned, a southern hemisphere dispersal route eastward from Aus-
tralia and would suggest a series of island stepping-stones for current-borne
larval stages lying south of the Tropic of Capricorn: Lord Howe, Norfolk,
Kermadec, Rapa, Morotiri (Bass), Pitcairn, Henderson, and Ducie. This island
arc is completed to South America by Sala y Gomez, San Felix, and San Am-
brosio. Juan Fernandez Islands, lying south as well as east of Easter Island,
have but one brachyuran species found also at Easter Island, according to Balss
(1924): Leptograpsus variegatus (Fabricius). The southern hemisphere island
arc is much more probably involved in its trans-Pacific distribution than the
hypothetical southern continent to which Balss (1924), following Arldt (1907,
p. 114), took recourse. Evidence is constantly mounting to show that planktonic
larvae of benthonic species, propelled by powerful currents, span greater oceanic
distances than were once believed possible (Garth, 1966; Briggs, 1967; Scheltema,
1968). Certainly, of all Pacific islands, with the possible exception of Clipperton
in the north Pacific, the sweepstakes route to Easter Island puts this hypothesis
to its severest test.

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iy | )

PROCEEDINGS —f “7/2. Biological Labor,

CALIFORNIA ACADEMY OF SCIENCES / ©

FOURTH SERIES Wit

Vol. XXXIX, No. 18, pp. 337-382; 13 figs.; 5 tables. October 24, 1973

TWO NEW STONEFISHES (PISCES,
SCORPAENIDAE) FROM THE INDO-WEST
PACIFIC, WITH A SYNOPSIS OF THE
SUBFAMILY SYNANCEIINAE

By
William N. Eschmeyer

California Academy of Sciences, Golden Gate Park, San Francisco, California 94118

and

Kaza V. Rama Rao
Zoological Survey of India, 10 Leith Castle South Street, Madyas-28, India

ABSTRACT. Two new stonefishes are described and placed tentatively in the genus
Synanceia. A discussion of the limits of the subfamily Synanceiinae is given, and
six genera (Synanceia, Erosa, Dampierosa, Trachicephalus, Leptosynanceia, and
Pseudosynanceia) are recognized. Related genera are discussed as are nomenclatural
problems. Brief descriptions and illustrations are provided for the species referable
to the scorpionfish subfamily Synanceiinae.

INTRODUCTION

The common name ‘stonefishes’ is applied here to all members of the scor-
pionfish subfamily Synanceiinae, although the name is more commonly used
only for the fishes of the genus Synanceia. Species of the genus Synanceia are
the most venomous of fishes, capable of causing death in man (see Halstead,
1970). Despite the interest in stonefishes, the species are poorly known, and
it is of interest that two new species have been found. The new species are
referred tentatively to the genus Synanceia, and their large venom glands sug-
gest that they may be just as venomous as the better-known and larger-sized
species Synanceia horrida and S. verrucosa. Generic allocation of the new spe-
cies necessitated an examination of other genera and species believed to be
related to the genus Synanceia. The genera referable to the subfamily Synancei-
inae contain few species and most of the species (nine or ten) are rarely en-

[337]

338 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

countered, so that it was thought desirable to provide a brief synopsis of the
subfamily. Fishes of the subfamily Synanceiinae are restricted to the warm
waters of the Indo-West Pacific faunal region. Some species are found on reefs,
but other members of the subfamily occur on muddy or sandy bottoms in
marine and brackish waters.

ACKNOWLEDGMENTS

Specimens of the two new species were obtained from five sources: The
United States National Museum of Natural History (USNM), Stanford Uni-
versity (SU) (collection now housed at the California Academy of Sciences
(CAS) ), the Zoological Survey of India (ZSI), the American Museum of Natu-
ral History (AMNH), and a collection made by Victor G. Springer in the Red
Sea as a part of a joint project of the United States National Museum of Natu-
ral History and the Hebrew University in Jerusalem (HUJ), Israel. We are
indebted to Springer and the late Dr. H. Steinitz for the loan of this Red Sea
material. William Smith-Vaniz first brought to Eschmeyer’s attention one of
the new species in 1967. A. G. K. Menon (ZSI) provided information on a
specimen in his care. Depositories of specimens other than those listed above
include the British Museum of Natural History (BMNH), Rijksmuseum van
Natuurlijke Historie (RMNH), Australian Museum, Sydney (AMS), Zoologi-
cal Institute, University of Tokyo (ZIUT), and the Academy of Natural Sci-
ences of Philadelphia (ANSP). Assistance by the staffs of these institutions
has been most generous, and we especially thank A. Wheeler, M. Boeseman, J.
Paxton, T. Abe, J. Atz, D. Rosen, and J. Bohlke for their assistance. Lillian J.
Dempster has offered valuable help in the preparation of this and other papers
on scorpionfishes. Her assistance with literature, selection of new scientific
names, and editorial comments have been substantial. Support for this study
came from the National Science Foundation grant GB-15811. A visiting scien-
tist fellowship provided by the California Academy of Sciences allowed Rama
Rao to participate in this study. The drawings were made by Katherine Smith.
M. Giles, W. Freihofer, P. Sonoda, T. Iwamoto, J. Gordon, F. Steiner, B. Wese-
mann, C. Pape, and K. Boyer assisted in the project.

METHODS

Methods follow those used by Eschmeyer (1969). Most scorpionfishes have
the last two elements in the dorsal and anal fins united or close together at
their bases and usually they are supported by a single pterygiophore. In the
subfamily Synanceiinae, however, species of some genera have the last element
single, well separated from the previous ray, and supported by its own pterygio-
phore. To distinguish between these two conditions the addition of ‘'2” or
the statement ‘last double” implies that the last soft ray is a double one and

Vor. XXXIX] ESCHMEYVER & RAMA RAO: NEW STONEFISHES 339

the absence of “2” or the statement “last single” indicates that the last ele-
ment is separate soft ray. In these fishes there is no depression or pit where
the premaxillaries join, so measurements originating from the anterior end are
taken from the symphysis of the premaxillaries.

Subfamily SYNANCEIINAE

The limits of this subfamily are in doubt. Bleeker (1874) included Pelor
|= Inimicus|, Synanceia, Leptosynanceia, and Polycaulus |= Trachicephalus |
in a family Synanceiidae (his “Synanceioidei’”’). J. L. B. Smith (1958) included
the genera Minous, Inimicus, Choridactylus, Synanceia, and Synanceichthys in
the family Synanceiidae (his ‘“‘Synanciidae”) for western Indian Ocean species.
De Beaufort (#7 Weber and de Beaufort, 1962) included Synanceia, Lepto-
synanceia, Polycaulus, Inimicus, and Minous in the family Synanceiidae (his
“Synanceidae”) for Indo-Australian species. Matsubara (1943) treated Minous
as a separate subfamily Minoinae, /nimicus as a separate subfamily Pelorinae,
and studied only Synanceia verrucosa as a representative of the subfamily
Synanceiinae. The genus Erosa is thought by us to be related to Synanceia;
Matsubara (1943) treated the genus Erosa as a separate subfamily Erosinae.
Non-Japanese genera were not studied by Matsubara.

At the present time we suggest the following allocation of these genera and
their synonyms (see also generic synonymies in text for additional synonyms):
Synanceiinae: Synanceia (including Synanceichthys), Leptosynanceia, Trachicepha-

lus [Polycaulus a synonym], Pseudosynanceia, Erosa, and Dampierosa.
Inimicinae (synonym Pelorinae): IJnimicus [Pelor a synonym] (including Choris-
mopelor), Choridactylus (including Choridactyloides) .

Minoinae: Minous (including Corythobatus, Decterias, Lysodermus, and Para-

minous ).

Species of the subfamily Minoinae are characterized externally by the pres-
ence of a free lower pectoral ray in each fin, while the species in the subfamily
Inimicinae have two (/nimicus) or three (Choridactylus) free pectoral rays.
(The monotypic genus Cheroscorpaena (Mees, 1964) has three free pectoral fin
rays; subfamilial placement of this genus is uncertain, but it probably belongs
near the distinctive genus A pistus.) Species of the subfamilies Minoinae and
Inimicinae will be treated in subsequent articles.

One feature which might prove to be important in the classification of these
fishes at the subfamilial level and below is the presence of what appear to be
skin glands or organs. These apparently pored structures tend to be scattered
over the body in fishes of the subfamily Synancelinae. They are present in rows
(usually above the anal fin or below the dorsal fin) in some genera, and in
Synanceia, Erosa, and Dampierosa they are reflected as warts. In the subfamily
Inimicinae these glands are concentrated in a widely spaced row above the

340 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

lateral line and in a patch behind the head. In the subfamily Minoinae, and
presumably in other scorpaenoid fishes, they are absent. Frequently the glands
contain a brown, hardened, wax-like substance in preserved specimens. A tiny
buried scale accompanying each gland can be seen in alizarin-stained speci-
mens. We do not know the function of these glands and have not studied them
in detail.

The following trends are noted for the subfamily Synanceiinae:

1. Dorsal spines increase from 12 to 17 (occasionally 11 spines in those species
which normally have 12 spines).

2. Dorsal soft rays decrease from 9 (10) to 4, but increase to 12-14 in one

genus.

Fin spines change from firm and strong to flexible.

Pelvic rays are reduced from I+ 5 to I1+3.

Pectoral fin rays decrease from about 18 or 19 to 11.

Eyes move to the dorsal surface of head; eyes become smaller.

Mouth shifts to a superior position.

Swimbladder is lost.

Vertebrae increase from 24 to about 30.

oon an nn SP W

The definition of genera within the subfamily Synanceiinae is difficult, par-
ticularly because of the (probably ‘rapid’) evolution of certain features listed
above as trends. For example, Synanceia is sometimes restricted to S. horrida,
and Synanceichthys to S. verrucosa (as verrucosus) (Smith, 1958, p. 173).
These 2 species and a presumed (by de Beaufort) hybrid (S. platyrhynchus)
were treated in the genus Synanceia by de Beaufort (72 Weber and de Beaufort,
1962); Whitley (1930) placed ‘platyrhynchus’ in its own subgenus Nofua. The
two new species described here further complicate this. One of the new species
has 11 pectoral rays, and the other normally 14 rays, while ‘platyrhynchus’ has
17, ‘horrida, 15-17, and ‘verrucosa,’ 18-19. Furthermore, one of the new species
has the pelvic rays reduced to I + 4, as do some specimens referable to ‘horrida.’
Given certain trends which occur in the subfamily, we do not feel that these
differences are so important and we have expanded the genus Synanceia. We treat
the genera Evosa, Dampierosa, Trachicephalus, Pseudosynanceia, and Lepto-
synanceia as monotypic, although more than one nominal species exists for some
of these genera. Subsequent workers may wish to unite Erosa and Dampierosa
in one genus.

SUBFAMILY DIAGNOsIS. Scales absent; no free pectoral rays; skin glands
present (appearing as ‘warts’ in some genera); dorsal spines 11—17, dorsal soft
rays 4-14 (last single or double); anal spines normally 2—4 (difficult to dis-
tinguish spines from soft rays in some species), anal soft rays 4-14, total anal
spines and rays 7-16; pelvic rays I+ 5, 1+ 4, or 1+ 3; pectoral rays 11-19;

VoL. XXXIX] ESCHMEYER & RAMA RAO: NEW STONEFISHES 341

second suborbital bone (= third infraorbital) broad, not T-shaped, attached to
preopercle; third and fourth suborbital (fourth and fifth infraorbital) bones
absent; vertebrae 23 to 30.

Key TO THE GENERA AND SPECIES OF THE SUBFAMILY SYNANCEIINAE

1. Mouth terminal, only slightly oblique; eyes placed laterally on head, directed out-

STAT, C1 Smee eee ee pt na ee Se ee ee 2
Mouth vertical or superior; eyes on dorsal surface of head, directed outwards and
Tip waALrdsmorcompletely* upwards 2. 6 eee eS 3
2. Dorsal spines 14; dorsal soft rays 544 to 64, usually 614; anal spines 3; anal soft
rays 514 to 64, usually 514; pectoral fin rays 14-16, usually 15 _._. Erosa erosa
Dorsal spines 12 (possibly 13); dorsal soft rays 914 (possibly 844); anal spines 2;
pectoral fin rays 12 (11 or 13 should be expected) ___-__.-_-_ Dampierosa daruma
See DOLsalmspimes: 0; Ob TOLe, 2 el ee 4
Worsalespines: Wl—14s . Sess he a ee ee ee ee ee ee ee 5
Amehewmicninewith spine and! 3 soft rays) 22 Pseudosynanceia melanostigma
Pelwicmtinewithe laspine and) 4 soft rays == 5 es Leptosynanceia asteroblepa
5. Anal fin with 3 spines and 414% (5) to 6% (7) soft rays, total anal fin elements
(10) 2A Sp. Se ee ee ee ee i ee ee 6

Anal fin with 2 spines and 12-14 soft rays, total anal fin elements 14-16 _.-_
En Selene Se ee ee ee _ Trachicephalus uranoscopus

6. (3 choices) Pectoral fin rays 11 (10 or 12 should also be expected) — Synanceia alula
Pectoral fin rays 14, rarely 15 (13 should be expected; one type specimen with 15 on

neem GMO TMNT et) ee oe Be as ee eee Synanceia nana

Pectoral fin rays 15 or more (if 15 or 16, eyes will have a crest above them) —_ 7

7. Pectoral fin rays 18-19; no bony crest above eye _______ Synanceia verrucosa

Pectoraletin rays 15-17; bony crest present above eye —= = == 8

8. Ridge connecting eyes across interorbit continuous, with no break in middle —___
a a Rr ED SV NONGCIURNONILOW

Eyes nearly connected by a bony ridge across interorbit but with a concavity at
Gentemlianspeciesot sumcertain) statusi]| = ==" Synanceia platyrhynchus

Genus Synanceia Bloch and Schneider

Synanceia Biocu and SCHNEIDER, 1801, p. xxxvii (brief description; listed species; type-
species Scorpaena horrida Linnaeus by subsequent designation of Jordan, 1919, p. 58).

Synanceja BLocu and SCHNEIDER, 1801, p. 194 (brief description; species descriptions; mis-
print for Synanceia [corrected in corrigenda on p. 573, see ‘Nomenclatural remarks’]).

Synanchia SwAINson, 1839, pp. 61, 180-181 (unjustified emendation of Synanceia [see ‘No-
menclatural remarks’]; three subgenera).

Bufichthys Swatnson, 1839, p. 268 [but not pp. 180-181] misprint for Synanchia [see ‘No-
menclatural remarks’]).

Synancidium MULier, 1843, p. 302 (type-species Scorpaena horrida Linnaeus by subsequent
designation of Swain [see ‘Nomenclatural remarks’]; Miiller proposed the genus for
“Synanceia mit Vomerzahnen”).

Synancydium, AcAssiz, 1845, p. 63 (unjustified emendation of Synancidium Miiller).

Synanceichthys BLEEKER, 1863, p. 234 (type-species Synanceia verrucosa = Synanceta brachio
by original designation).

Emmydrichthys JorpAN and RuTTER in Jordan, 1896, pp. 221-222 (type-species Emmydrich-
thys vulcanus Jordan and Rutter by monotypy).

342 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Nofua WuitLry, 1930, p. 24 (as a subgenus of Synanceja [sic]; type-species Synanceia
platyrhynchus Bleeker by original designation).
Deleastes SEALE, 1906, pp. 80-81 (type-species Deleastes daector by original designation).

NOMENCLATURAL REMARKS. Much confusion exists over the spelling of this
genus and its various synonyms, as well as the subfamily spelling. The genus is
spelled Synanceja by many recent authors (e.g., Smith, Whitley, de Beaufort),
but the correct spelling is clearly Synanceia. In the original description Bloch
and Schneider, on page xxxviil, spell the name as Synanceia, on page 194 they
spell it Synanceja, and in plate 45 Synanceia; in their “corrigenda” on page 573
they state ‘“‘|page| 194. genus 50. scribe: Synanceia.” Subsequent workers who
have commented on this problem missed the corrigenda (e.g., Briggs, 1961, p.
164).

Gill (1905, p. 221, et seqq.) said of the problem of the genus Synanceia and
some other genera, “Complication has resulted by reason of the intrusion of the
incompetent Swainson into the field.’ Swainson (1839) did add considerable
confusion, but this uncertainty easily can be resolved. As discussed by Gill,
Swainson attempted to reclassify the ““Synanceines” and named three subgenera,
but in three places (1839: pp. 61, 180-181, and 268) Swainson variously inter-
changed names and diagnoses. Gill summarized the equivalent categories as
follows:

p. 61 pp. 180-181 p. 268
Erosa = Bufichthys = Synanchia Cuvier (species erosa)
Synanchia == Synanchia = Bufichthys (species horrida and grossa)

Trichophasia = Trachicephalus = Trachicephalus (species elongatus)

Gill allocated the individual treatments by Swainson to the genera Synanceia
and Erosa of other authors. Trichophasia and Trachicephalus were synonymized
by Gill and he serves as first revisor selecting Trachicephalus over Trichophasia.
Problems remain only for Swainson’s subgenera Evrosa, Synanchia, and Bufich-
thys. Authors, e.g. Swain (1882), Jordan and Starks (1904), Jordan (1919),
and Whitley (1930), as well as Gill, consider Swainson’s Synanchia an emenda-
tion of Synanceia, although Bleeker (1874, pp. 4,11) indicated that the species
‘erosa could be placed in Synanchia of Swainson. Synanchia is an unjustified
emendation and cannot be used as a separate genus for the species ‘erosa’ as by
Bleeker (see Gill, 1905). It is clear from Swainson’s text (but not from p. 268)
that he intended first Erosa and then Bufichthys for the species ‘erosa. It
therefore appears that on page 268 Swainson switched the two headings Bufich-
thys and Synanchia (compare particularly his diagnosis and subgeneric cate-
gories on pp. 180-181 with p. 268).

Swain (1882, p. 277), in his review of Swainson’s genera, dealt only with
the entries on Swainson’s pages above 200 (7.e., p. 268 but not pp. 61 or 180—

Vor. XXXIX] ESCHMEYER & RAMA RAO: NEW STONEFISHES 343

181) and was therefore not aware of the switched headings. Unlike Gill, Swain
referred Bufichthys to Synancia |Synanceia| and selected ‘horrida’ as the type-
species of Bufichthys. But Bufichthys p. 268 is really Synanceia while Bufich-
thys on p. 181 is really Erosa. Swain’s selection of ‘korrida’ as the type of
Bufichthys of p. 268 does not affect the type-species for Bufichthys on p. 181.
The type-species of Bufichthys is really ‘erosa’ by monotypy. (It does appear
that Swain was in fact the first to select the type-species ‘korrida’ for Synanci-
dium Miiller; see Swain’s footnote 4 on p. 277; Jordan, 1919, also regarded
‘horrida’ as the type-species of Synancidium Miller.)

Bufichthys Swainson (on p. 181) is the original generic description of
Bufichthys, and Gill (1905, p. 223) serves as the first revisor selecting the
genus Erosa Swainson (p. 61) over Bufichthys. This interpretation seems to
have been the aim of Swainson and corresponds to current usage.

GENERIC DIAGNOsIS. Dorsal fin normally with 13-14 spines and 4%2-5 to
74-8 soft rays (last ray usually double); anal fin normally with 3 spines and
4%-5 to 6'~7 soft rays (last soft ray usually double); pectoral fin rays 11-19;
pelvic fin rays I+ 5, or I+ 4 in some specimens and in one species; mouth
vertical; eyes directed mostly upwards; vertebrae usually 24; swimbladder
absent; body covered with warts.

Remarks. The limits of the genus Syvanceia are discussed in the subfamily
section and reasons for placing some of the nominal genera in the synonymy of
Synanceia are discussed above.

The genus Emmydrichthys Jordan and Rutter was based on a specimen of
S. verrucosa with an abnormal dorsal fin. Deleastes was established by Seale
for a species which he thought had shorter pelvic fins, pelvic fins located more
posteriorly, and a smoother skin (see our account of S. verrucosa).

Synanceia nana Eschmeyer and Rama Rao, new species.
(Figures 1-2; tables 1, 3-5.)
No literature applies to this species.

HoLotyPE. USNM 209417 (a specimen 73.2 mm. in standard length), Red
Sea, Israel, northwest coast of the Gulf of Aqaba, bay at El Himeira, depth of
capture to 18 meters, Victor G. Springer and assistants, 1100-1230 and 1315—
1415 hours, 16 July 1969.

ParATYPES. CAS 14991 (1, 47.8), taken with the holotype; CAS 14992 (2,
21.3-39.7) and HUJ uncatalogued (1, 70.5), Red Sea, Israel, Gulf of Aqaba,
bay at El Himeira, in 9-12 meters, Victor G. Springer and assistants, 0945—
1215 hours, 8 September 1969; USNM 209418 (1, 24.0), Red Sea, Israel, Gulf
of Aqaba, between Marset Mahash el Ala and Marset Abu Samra, 32 kilometers
as road goes south of Marine Laboratory, in depths to 3.5 meters, Victor G.
Springer and assistants, 1030-1300 hours, 2 September 1969; USNM 209419

[Proc. 4TH Serr.

ACADEMY OF SCIENCES

CALIFORNIA

344

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Vor. XXXIX] ESCHMEYER & RAMA RAO: NEW STONEFISHES 345

Ficure 2. Drawing of head of Synanceia nana, based mostly on the holotype, 73 mm.
Sl.

(1, 52.8), Red Sea, Gulf of Suez, Et-Tur, Sinai Peninsula, 100 kilometers north
of Sharm el Sheikh as road goes, depths to 9 meters, Victor G. Springer and
assistants, 1015-1315 hours, 27 September 1969; CAS 14993 (1, 71.5), Red
Sea, Gulf of Suez, off Port Safaga, 27°16’15’”N., 33°47’30”E., in 3 meters, H. A.
Fehlmann and H. K. Badwi, 6 January 1965; AMNH 18385 (1, 102), Saudi
Arabia, Persian Gulf, Tarut Bay, near Ras Tanura spit, R. Bowen, July 1947.

DISTINGUISHING FEATURES. Dorsal XIV—XV (usually XIV) + 5; anal III
+ 4 to 6 (usually ITI + 5); pectoral 14-15(16), usually 14; pelvic I+ 5; head
broad, depressed, with pits; no large pit below eye; deep depression between
eyes; rectangular depression on occiput.

DEscriPpTION. Dorsal fin with 14 spines (15 in one specimen); all spines
short, nearly the same length, with thick skin covering the venom glands. Dor-
sal soft rays 5, all unbranched. Anal fin with 3 spines, first very short. Anal
soft rays 4—6, usually 5, last single, all unbranched. Pectoral fin rays 14-15 (16
on one side in 1 specimen), usually 14, all unbranched. Pelvic fin with 1 spine
and 5 unbranched soft rays. Gill rakers rudimentary, total 7-10, 0-2 on upper
arch, 7-8 on lower arch. Lateral line with 10-12 tubes (including one on
caudal), lateral line pores paler than nearby ‘warts’ in smaller specimens. Ver-
tebrae 24 (7 specimens), or 23? (1), or 25 (1).

[Proc. 4TH SER.

CALIFORNIA ACADEMY OF SCIENCES

346

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Vor. XXXIX] ESCHMEYER & RAMA RAO: NEW STONEFISHES 347

Body shape and coloration as in figure 1. Head (fig. 2) broad, depressed.
Eyes elevated, directed up and out. Posterior interorbital area as a deep pit,
bordered behind by a ridge which forms anterior edge of shallow rectangular
occipital pit. No deep pit before and below eyes. Low crest on upper posterior
corner of orbit. Most head spines difficult to distinguish; preorbital bone with
main spine pointing down and slightly forward, two lumps or ridges on anterior
edge. Supplemental and 4 preopercular spines present, all short. Other spines
absent or developed as lumps or ridges. Body covered with warts. Small teeth
on jaws and vomer; none on palatines.

Head and body pallid to tan, with darker brown areas, most notably dark on
back between posterior part of spinous dorsal fin and anterior half of anal fin
and at base of caudal fin. Pale irregular spots on a tan background on outside
and inside of pectoral fin, fin pale in middle and dark distally. Other fins,
except spinous dorsal, dark distally with tips of rays or margins white. Smallest
specimens with posterior part of body and caudal and anal fin darker than in
larger specimens.

Described from nine specimens. Probably a small species, largest available
102 mm. in standard length.

CoMPARISONS. This species most resembles S. verrucosa in body and head
shape but differs from it by having a lower pectoral ray count (14-15 versus
18-19) and a different dorsal ray count (usually XIV + 5 versus usually XIII
+ 6%). Synanceia nana lacks a pit below the eye, unlike S. verrucosa, S. platy-
rhynchus, and especially S. horrida. Synanceia nana has a rectangular depres-
sion on the occiput which is absent in S. verrucosa. Synanceia nana has a higher
pectoral ray count than S. alula (14 or more versus 11). Other differences are
given in the key or may be found by comparing ‘Distinguishing features’ sec-
tions.

REMARKS. The single specimen from outside the Red Sea (Saudi Arabia,
AMNH 18385, 102 mm. S.L.) has higher counts of dorsal, anal, and pectoral
rays and one more vertebra (table 1), and is darker brown in coloration than
the Red Sea specimens. It also has the tips of the inner pelvic rays joined to
each other by skin rather than having the tips of the inner rays attached to the
body. It is about 30 mm. longer in standard length than our largest Red Sea
specimen. We feel it is referable to S. nana, but study of additional specimens
from outside the Red Sea would be desirable.

DISTRIBUTION. Synanceia nana is known from the type material from the
Gulf of Suez and the Gulf of Aqaba in the Red Sea and from Saudi Arabia.
Depths of capture range from about 3 to 10 m.

Name. The specific name ‘nana’ is a Latin noun meaning dwarf or pygmy,
alluding to the small size of this species.

348 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Synanceia alula Eschmeyer and Rama Rao, new species.
(Figures 3-4; tables 2-5.)

Synanceia verrucosa, KEEGAN et al., 1964, fig. 26 (good photograph of a specimen from the
Solomon Islands).

HoLotyPe. SU 14673 (a specimen 85.0 mm. in standard length), Nicobar
Islands, Nancouri Island, 8°N., 93°40’E., shore collection, R.J.M.S. Investigator
station 615, 27 October 1921.

PARATYPES. ZSI 289/2 (1, 72.0), taken with the holotype; USNM 209420
(2, 30.0-48.5), Solomon Islands, New Georgia, Munda Pier, in old coral, col-
lected by W. Chapman, 7 May 1944; USNM 209421 (1, 27.8), Solomon Is-
lands, New Georgia, Munda lagoon, collectors Chapman and Cheyne, 19 June
1944.

DISTINGUISHING FEATURES. Dorsal XIII + 5'-6'%; anal normally III +
4%2-5' (last soft ray usually double); pectoral 11; pelvic I+ 4; head broad,
depressed, large pit below eye.

DescripTIon. Dorsal fin with 13 spines; spines nearly same length, first
shortest, second and third longest, covered by thick skin; venom glands promi-
nent, on distal half of spines. Dorsal soft rays 5’2—-6'2, branched distally, last
%2 ray virtually a distinct separate ray (without dissection soft rays probably
would be counted as 6-7). Anal fin with 3 spines (4 in one specimen) and 5%
(normally) or 4'2 soft rays; first anal spine about half length of second; last
anal soft ray usually double (but appearing as two close-set rays); anal rays
branched distally except in small specimens. Pectoral rays 11, all rays branched
at tips in largest 2 specimens. Pelvic fin with 1 spine and 4 soft rays, but 3 soft
rays on left side of holotype. Gill rakers rudimentary, total 7-8, O-1 on upper
arch, 7 on lower arch. Lateral line tubes difficult to distinguish from ‘warts,’
about 11 lateral line scales. Vertebrae 24 (4 specimens).

Body shape and coloration as in figure 3. Head (fig. 4) broad, depressed;
eyes elevated, with crest at upper posterior corner. Large pit below eye, largest
in largest specimens; deep pit below parietal and nuchal spines. Occipital pit
shallow or nearly absent. Most head spines developed as lumps or ridges, poorly
defined. Preorbital bone prominent, main spine as a broad lump pointing down,
with lateral ridges pointing anteriorly. Supplemental preopercular spine absent
or fused with first preopercular spine; four preopercular spines present. Small
teeth on jaws and vomer, none on palatines.

Head and body tan or brown, with darkest areas on back and between pos-
terior portion of spinous dorsal fin and anal fin; a broad dark band at base of
caudal fin. Dorsal fin brown in holotype (possibly discolored), paler in other
specimens.

Described from 5 specimens. Probably a small species, largest available 85.0
mm. in standard length.

349

NEW STONEFISHES

AMA RAO:

A

ESCHMEYVER & R

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350 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Ficure 4. Drawing of head in holotype of Synanceia alula, 85 mm. S.L.

TABLE 2. Counts and measurements for type specimens of Synanceia alula (measurements
in mm., percent standard length in parentheses).

Paratypes
Holotype

SU ZSI USNM USNM USNM

14673 289/2 209420 209420 209421
Standard length 85.0 72.0 30.0 48.5 27.8
Dorsal rays XII+5% #£XiIlI+6% #$;XIII+5% £XiIII+6% #«4XiIII-+5%
Anal rays TI-+5 T1I-+5% IV+4'% IlI+5% I1I+4%
Pectoral rays ialeila Talal 11,11 ileal iis
Pelvic rays I+3,1+4 I+4,1I+4 I+4,I+4 I+4,1+4 I+4,1+4
Vertebrae 24 — 24 24 24
Head length 35.4 (42) = 11.8 (39) 19.2 (40) 11.5 (41)
Body depth 31.8 (37) — 11.0 (37) 17.0 (35) 10.0 (36)
Orbit diameter 4.2 (05) —- 1.6 (05) 2.4 (05) 1.7 (06)
Snout length 8.3 (10) aa 3.0 (10) 4.8 (10) 2.8 (10)
Interorbital width 4.6 (05) oa Zeon (Op) 4.0 (08) 1.8 (06)
Jaw length 16.8 (20) — 5.5 (18) 9.0 (19) 4.7 (17)
Predorsal fin length 23.4 (28) a 8.3 (28) 13.5 (28) 7.8 (28)
Pectoral fin length* 39.5 (47) = 13.3 (44) 22.4 (46) 12.0 (43)

Pelvic fin length 23.5 (28) —- 8.8 (30) 14.8 (30) 8.0 (29)

* from base of lower ray to tip of fin

351

Vor. XXXIX] ESCHMEYER & RAMA RAO: NEW STONEFISHES

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352 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

CoMPARISONS. Synanceia alula resembles S. horrida in many respects, par-
ticularly by having a deep pit below the eye, but S. alula differs from S. horrida
and all other species of the genus in having a very low pectoral ray count (11
versus 14 to 19). Other differences are given in the key or may be found by
comparing ‘Distinguishing features’ sections.

DISTRIBUTION. Synanceia alula is known only from the type specimens from
the Solomon Islands in the western Pacific Ocean and from the Nicobar Islands
in the northern Indian Ocean. The specimen figured by Keegan eé al. (1964)
also was from the Solomon Islands.

Name. The specific name ‘alula’ (al’-t-la), meaning little wing, is a Latin
feminine noun, the diminutive of a/a (wing), alluding to the very low pectoral
ray count in this species.

Synanceia horrida (Linnaeus).
(Figures 5-6; tables 3-5.)

(A partial synonymy pertinent to the scope of the study.)

Scorpaena horrida LINNAEUS, 1766, p. 453 (original description; type locality East Indies).

Scorpaena “alepidota”’ Biocu, 1787, pp. 106-108, pl. 183 ([see ‘Remarks’ below]).

Synanceia horrida, BLocH and SCHNEIDER, 1801, pp. xxxvii, 194, 573 (misspelled genus as
Synanceja on p. 194, corrected on p. 573; brief description and synonymy) ; CUVIER in
Cuvier and Valenciennes, 1829, pp. 440-446 (lengthy description; review of earlier
literature) ; BLEEKER, 1849, pp. 4, 9 (synonymy; brief description; range); 1852a, pp.
230, 237, 242 (compiled range) ; 1874, pp. 4, 11-13, fig. 1 on pl. 1 (synonymy; descrip-
tion; distribution) ; 1879, fig. 7 on pl. CCCCXVII (figure from Bleeker, 1874) ; Herre,
1951, pp. 479-480 (synonymy; description; Philippines) ; p—E BEAUFORT in Weber and
de Beaufort, 1962, pp. 95-97 (synonymy; description; distribution).

Synanceia grossa GRAY, 1830, pl. 97 (plate only, plus caption; type locality Singapore).

Synanceia trachynis RICHARDSON, 1842, pp. 385-389 (original description; type locality Port
Essington, Northern Territory, Australia).

Synancidium horridum, GUNTHER, 1860, pp. 144-146 (brief synonymy; description of skele-
ton; BMNH specimens); KNER, 1865, p. 119 (description; Java); Day, 1875, p. 162,
fig. 3 on pl. XXXIX (synonymy; description) ; FOWLER, 1928, pp. 297-298 (compiled).

Scorpaena monstrosa GRAY, 1854, p. 117 (after Gronovius and ‘horrida’ of Linnaeus).

Synanceja horrida, McCuLtocH, 1929, p. 392 (synonymy; Australia) ; WHITLEY, 1930, pp.
24-25 (synonymy; comparisons with S. trachynis).

Synanceja trachynis, WHITLEY, 1930, pp. 25-26 (synonymy; Australian records; venom) ;
1932a, pp. 306-309, figs. 1-2 on pl. 4 (description; fresh coloration; Australia) ; 1960,
pp. 1-6, 4 figs. (semi-popular account; habitat, coloration, venom); Muwnro, 1967, p.
540 (description; New Guinea).

Remarks. References on venom and related subjects, as well as specimen
figures, are given by Halstead (1970). Some Australian workers continue to
recognize S. trachynis as a species distinct from S. horrida; we have not made
an exhaustive study but our specimens seem to indicate that the two names
are synonyms.

Scorpaena alepidota of Bloch (1787) originated through an inadvertent

Vor. XXXIX] ESCHMEYVER & RAMA RAO: NEW STONEFISHES 353

error. As evidenced by style in Bloch’s work, a heading “‘Scorpaena horrida” be-
fore the words Scorpaena alepidota was omitted, giving the incorrect impression
that ‘alepidota’ was proposed as the scientific name. The plate was correctly
labeled ‘horrida.’ The name ‘alepidota’ has no separate status nomenclaturally.
MATERIAL EXAMINED. (No counts were made on specimens which have
standard length omitted.)
SINGAPORE

SU 30873 (5 specimens, 67.8-134 mm. S.L.), Herre 1934 Pacific Expedition,
A. W. Herre, 15 March 1934. SU 36001 (1, 151), Singapore market, A. W.
Herre, February 1937. SU 32128 (1, 159), 8 February 1899.

THAILAND

USNM 209422 (2, 147-190), Paton Bay, Patong Phuket, International Indian
Ocean Expedition, Anton Bruun Cruise No. 1, 22 April 1963. CAS 15073 (1,
102), Rayong Province, SE. of Ban Phe Fisheries Station, 12°35’40’N., 101°
25’43’E., Fehlmann and assistants, 28 April 1960. Plus ANSP 89676 (1).

PHILIPPINES

CAS 15074 (1, 136), Calaogao, Cauayan, Negros Island, collected among coral
stones at night, 10°N., 122°30’E., Q. Akala, 18 May 1960. SU 34132 (1, 196),
Sitankai, Sulu Island, Oriental Expedition 1936-37, A. W. Herre, 9 January
1937. SU 29780 (1, 158), Manila Bay, A. W. Herre, December 1933. SU 28363
(1, 95.0), Atimonan, Tayabas, Herre 1931 Philippine Expedition, A. W. Herre,
1931. SU 39137 (1, 111), Ragay Gulf, Luzon, A. W. Herre, 1940. Plus SU
28365 (2).
BATAVIA
ANSP 90442 (1), Baai.
NEw GUINEA

SU 26744 (1, 160), Waigiu Island (= Waigeo Island), A. W. Herre, 7 June
1929.
AUSTRALIA

USNM 174014 (5, 57.5-73.7), Northern Territory, reefs and tide pools off
south entrance to lagoon, Groote Eylandt, Arnhemland Expedition, R. R. Miller
and party, 19-25 March 1948.

DISTINGUISHING FEATURES. Dorsal XIII-XIV (usually XIII) + 6%; anal
II-III (usually III) +51; pectoral 15-17, usually 16; pelvic I + 4—5, usually
5; all fin rays usually branched; head large, depressed, with deep pits; eyes
elevated, with bony crest above posterior corner of orbit, joined between orbits;
occiput depressed, forming a deep saddle behind orbits; a deep, mostly round
pit below eye; deep pit below parietal and nuchal spines. Body covered with
thick skin and warts.

354 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Ficure 5. Synanceia horrida, USNM 174014, sub-adult, 67 mm. S.L., Australia.

Vor. XXXIX] ESCHMEYVER & RAMA RAO: NEW STONEFISHES 305

Ficure 6. Synanceia horrida, CAS 15074, head of an adult, 142 mm. S.L., Philippines.

DEscrRIPTION. Dorsal fin with 13 spines, occasionally 14; spines nearly
same length, second to fourth longest, covered by thick skin. Dorsal soft rays
6%, branched distally. Anal fin with 3 spines (2 spines in 2 of 19 specimens)
and 5% soft rays, spines progressing in length from first to third, soft rays
branched distally. Pectoral fin rays 15-17, usually 16, all rays branched at tips.
Pelvic fin with 1 spine and 4 or 5, usually 5, soft rays [one specimen (SU
28363) abnormally with I + 3 on right and I+ 5 on left]. Gill rakers rudimen-
tary, total 13-15 in larger specimens, difficult to distinguish on upper arch in

our specimens below 75 mm. S.L., larger specimens with 5—6 on upper arch, all

356 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Taste 5. Frequency distributions of pectoral rays and pelvic soft rays in species of the
subfamily Synanceiinae.

Pelvic soft rays

Pectoral rays (left side) (left side)
11 WW 13 14 15 16 17 18 19 3 4 5
S. alula 5 — —- — 1 5a
S. nana = =—_ = 8 if Bee? Ye 9
S. horrida — = = = 3 14 2 = = >. 3 17
S. platyrhynchus a 1 a= 2a nae 1
S. verrucosa’ ee 19 3 =. 1** 21
E. erosa ~- _- — 1 14 1 16 an
D. daruma — 1 — — _ = a
L. asteroblepa — —- 3y 1 —_
P. melanostigma — = — 3 5 — = ass Le ” - a
T. uranoscopus —- —- — 6 a ee 16)
* 16 on right
** 5 on right

y+ one with 13 on right
lleft and right pectoral ray counts for 24 additional specimens: 17+18 (1 specimen), 18418 (22),
19419 (1).

sizes with 8—9 on lower arch. Lateral line pores usually 10-12, difficult to dis-
tinguish from warts. Vertebrae 24.

Body shape and coloration as in figure 5. Head (fig. 6) broad, depressed;
eyes elevated slightly, accentuated by large, mostly circular deep pit below
eyes and large crests above eyes. Occipital area deep, forming saddle between
eyes and beginning of dorsal fin; this area smooth, without warts. Deep pit
also present behind eye below parietal and nuchal spines. Preorbital bone prom-
inent, with broad spinous point directed downwards, with lateral ridges pointing
anteriorly. Suborbital ridge with one large lump in middle. Supplemental pre-
opercular spine usually present as a lump, first three preopercular spines usu-
ally present, first and second the largest. Other head spines indistinct or de-
veloped as lumps or absent. Small teeth on jaws and vomer, none on palatines.

Coloration of a small specimen as in figure 5. In preservative mostly brown;
fins tend to be darker distally, except for spinous dorsal fin; caudal darkest at
base, in middle, and distally; remaining areas of caudal fin streaked with white.

CoMPARISONS. Synanceia horrida is most like S. alula in having a large pit
below the eyes, but S. Aorrida is easily separated from S. alula by having more
pectoral rays (15-17 versus 11), among other features. From the widespread
S. verrucosa, S. horrida can be distinguished by having a large circular pit below
the eye and high crests above the eyes which are joined, leaving no pit between
the eyes. Other differences are given in the key and other ‘Comparisons’
sections.

DiIsTRIBUTION. Synanceia horrida has a fairly wide range, occurring from

Vor. XXXIX] ESCHMEYVER & RAMA RAO: NEW STONEFISHES 357

India eastwards to Java, New Guinea, Australia, the Philippines, and China.
Unlike S. verrucosa it is apparently absent from the central Pacific and from
western India to the Red Sea and Africa. It appears to be a continental and
‘large island’ species living on sandy or muddy bottom among rocks. (A record
of this species from Saint Helena in the Atlantic (Giinther, 1860, p. 145) was
presumed (Eschmeyer, 1971, p. 503) to be based on incorrect locality informa-
tion accompanying the specimen.)

Synanceia platyrhynchus Bleeker.

(Figure 7; tables 3-5.)

Synanceia platyrhynchus BLEEKER, 1874, pp. 4, 11, 14-15, fig. 2 on pl. 1 (original description ;
type locality Amboina); 1879, fig. 2 on pl. CCCXVI (figure of type from Bleeker,
1874) ; DE BrEAurort im Weber and de Beaufort, 1962, p. 99 (examined type and one
additional specimen with no locality data; thought to be a hybrid between S. verrucosa
and S. horrida).

Synanceja (Nofua) platyrhynchus, WHITLEY, 1930, p. 24 (as type of a new subgenus).

MATERIAL EXAMINED. RMNH 5898 (1, 129 mm. S.L., 164 or 165 mm.
T.L., holotype of S. platyrhynchus), Amboina. We could not locate the second
specimen mentioned by de Beaufort 7m Weber and de Beaufort (1962, p. 99).

REMARKS. De Beaufort gave the length of the two specimens he examined
as 153 and 220 mm. and stated that one of them was the type. One of these
measurements must be in error as the type in Leiden is 165 mm. T.L., the same
length Bleeker gave in the original description.

The type specimen appears to us to be referable to S. horrida, but it seems
to be abnormal in that the crests above the eyes are more poorly developed and
are not joined between the eyes. This leaves a depression between the eyes
which approaches somewhat the condition in S. verrucosa, although in S. ver-
rucosa the pit is broader and the eyes farther apart. The type of S. platy-
rhynchus is very similar to S. alula with regard to head shape and location of
crests and pits, but S. alula has far fewer pectoral rays (11 versus 17). We
have kept S. platyrhynchus as a separate entry in this paper to draw attention
to it in hopes that additional specimens may be found if it is in fact a species
distinct from S. horrida.

Counting by the methods used in this paper, the type specimen has dorsal
rays XIII + 6%, anal rays III + 5%, and pelvic rays I+ 5. De Beaufort (in
Weber and de Beaufort, 1962, p. 99) discusses the type specimen in more detail.
A photograph of the type as it appears now is reproduced as figure 7.

Synanceia verrucosa Bloch and Schneider.
(Figure 8; tables 3-5.)

(A partial synonymy pertinent to the scope of the study.)

Synanceia verrucosa BLocH and SCHNEIDER, 1801, pp. XX XVII, 195, pl. 45 (original descrip-
tion; type locality India; spelling of genus on p. 194 corrected to Synanceia in Cor-

358

CALIFORNIA ACADEMY OF SCIENCES

os
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[Proc. 4TH SER.

Ficure 7. Synanceia platyrhynchus, RMNH 5898, holotype, 129 mm. S.L., Amboina.

Vor. XXXIX] ESCHMEYER & RAMA RAO: NEW STONEFISHES 359

rigenda, p. 573); GUNTHER, 1860, p. 146 (synonymy; brief description; BMNH speci-
mens); BLEEKER, 1874, pp. 4, 11, 15-17 (synonymy; description; range); Day, 1875,
pp. 162-163, fig. 4 on pl. XXXIX (synonymy; description; range); BLEEKER, 1879,
fig. 5 on pl. CCCCXVII (figure only) ; Herre, 1951, pp. 479-482 (synonymy; descrip-
tion; Philippines) ; DE BEAUFORT 7m Weber and de Beaufort, 1962, pp. 95, 97-99 (synon-
ymy; description; distribution).

Scorpaena bicirrata LACEPEDE, 1801, pp. 333, 349-350 (original description, from Commerson).

Scorpaena brachion LAcEPEDE, 1801, pp. 333, 351-352, pl. 12 opposite p. 360 (original de-
scription, based on drawing from Commerson; no type locality).

Scorpaena Bicapillata SHAw, 1803, p. 273, pl. 40 (for Lacépéde’s Scorpaena bicirrata; de-
scription from Lacépede).

Scorpaena Brachiata SHAW, 1803, p. 274 (for Lacépéde’s Scorpaena brachion).

Synanceia brachio Cuvier in Cuvier and Valenciennes, 1829, pp. 447-454 (new spelling for
S. brachion Wacépede; synonymy; long description; review of earlier literature) ;
BLEEKER, 1849, pp. 4, 9-10 (synonymy; description; range); 1852a, pp. 233, 236, 240
(listed; Ternate, Banda, Ceram, and Waigioe).

Synanceia bicapillata, Cuvier in Cuvier and Valenciennes, 1829, pp. 454-456 (synonymy ;
description; discussion) ; BLEEKER, 1849, p. 4 (compiled range); 1852a, pp. 230, 242
(listed; Molucca).

Synanceia sanguinolenta Cuvier in Cuvier and Valenciennes, 1829, p. 447, footnote (original
description from Ehrenberg MS and figure; no locality).

Synanceja verrucosa, FOWLER, 1928, p. 299 (synonymy; remarks; range) ; MCCULLocH, 1929,
392-393 (synonymy; range).

Synanceichthys verrucosus, BLEEKER, 1863, p. 234 (listed; Ternate); WHITLEY, 1932a, pp.
309-310 (rare on Great Barrier Reef; AMS specimens from other localities) ; Munro,
1967, p. 540 (description; New Guinea).

Emmydrichthys vulcanus JorpAN and Rutter in Jordan, 1896, pp. 221-223, 562, pl. 26
(original description; type of new genus; type locality Society Islands).

Synanceia thersites SEALE, 1901, pp. 121-122 (original description; type locality Marianas
Islands [holotype BPBM 256, not found; paratype ANSP 91726]).

Deleastes daector SEALE, 1906, pp. 80-81, fig. 22 (original description; type locality Tahiti;
holotype BPBM 1360 [not found]).

Remarks. All the nominal species listed above have been recognized as
synonyms of S. verrucosa by previous workers. Emmydrichthys vulcanus was
based on a specimen with an abnormal dorsal fin. In the original description of
Deleastes daector, Seale reported it differed from Synanceia by having shorter
pelvic fins, pelvic fins located more posteriorly, and a smoother skin. The loca-
tion and apparent size of the pelvic fins depend in part on the position of the
hyoid arch on preservation, and we find that either of two conditions is com-
mon: the depth between the rear of the head and the pelvic fin will be shallow
when the hyoid arch is not depressed or will be deep when the arch is depressed
downward. The presence of warts was found to be somewhat variable. A more
thorough study should be made.

MATERIAL EXAMINED. (No counts were made, except of pectoral rays, on
specimens which have standard length omitted.)

[Proc. 4TH SER.

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VoL. XXXIX] ESCHMEYVER & RAMA RAO: NEW STONEFISHES 361

WESTERN INDIAN OCEAN

CAS 14943 (1 specimen, 123 mm. S.L.), Kenya, Andromache Reef just south
of entrance to Port Kilindini of Mombasa Harbor, 4°05’05”S., 39°40’39”E.,
over flat reef with dead coral and sand, International Indian Ocean Expedition,
Anton Bruun Cruise No. 9, sta. HA-1, 15 November 1964; SU 37205 (1, 154),
southern Andaman Islands, south of Corbyn’s Cove, Port Blair, under a coral
stone in low water, D. D. Mukerji, 16 December 1933; Plus ANSP 107718 (1)
and ANSP 107702 (1), Sechelles; USNM 19981 (1), Mauritius.

CEYLON

CAS 14969 (3), Trincomalee, outside harbor at coral cave, inside base of Royal
Navy of Ceylon, depth to 1.5 meters, W. Smith-Vaniz, 27 June 1969.

OKINAWA
MSN <71553° (1).

PHILIPPINES
SU 28362 (2, 121-187), Sitankai, Sulu Province, Herre Philippine Expedition,
A. W. Herre, 12 August 1931. SU 28364 (1, 51.3), Dumaguete, Herre Philip-
pine Expedition, A. W. Herre, 11 June 1931. Plus SU 34135 (1).

MICRONESIA

Palau Islands: CAS 14957 (1, 97.5), Auluptagel Island, Crocodile Cove, 7°17’
N., 134°29’E., Brittan et al., 29 July 1956. CAS 14959 (1, 106), Angaur Island
in Garangaoi Cove, south of Cape Nagaramudel, 6°53’50”’N., 134°7'49”E.,
DeWitt and party, 22 October 1957. CAS 14956 (2, 119-133), Ngadarak Reef
SW. of Auluptagel Island, 7°17’48’N., 134°28’37”E., Rikrik, 8 July 1956.
Plus CAS 14953 (1), CAS 14954 (1), CAS 14958 (1), CAS 14960 (1), and
CAS 14961 (1). Mariana Islands: CAS 14963 (1, 39.1), Guam, ca. % mile
SW. of Agat village, sand flat off north side of Bangi point, 13°22’36’N.,
144°38’53”E., Fehlmann, 12 October 1958. CAS 14965 (3, 54.2-121), Guam
reef and sand flat north of Tringhera Beach in Agana Bay, 13°28’53”N., 144°
45’45’”E., Fehlmann and Bronson, 7 April 1959. ANSP 91726 (1, 160, para-
type of S. thersites), Guam, Agana, A. Seale, 12 July 1900. Plus CAS 14966
(2), CAS 14962 (1), and CAS 14964 (1). Caroline Islands: CAS 14947 (1,
126), Yap Island, inlet on east side of Yap Island, 9°29’48”N., 138°26'57”E.,
Bronson and Hermana, 27 December 1959. Plus CAS 14952 (3) and CAS
14948 (1).

MELANESIA
Solomon Islands: CAS 15076 (1, 47.8), Bougainville, east side of Puk Puk
Island, sea beach outside of Poison Lagoon, Te Vega Expedition, Cruise No. 6,

362 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Sta. 243, William P. Davis, 9 March 1965. SU 6034 (1, 154), Sikiana Island,
Stewart group, Crocker Expedition, 16 May 1933.

POLYNESIA

Samoa: CAS 2228 (1, 115), A. Seale, 25 June 1929. CAS 2225 (2, 94.3-168),
Pago Pago, A. Seale, May 1929. Plus SU 9041 (3) and CAS 14967 (1). Fiji:
SU 21021 (1). Tonga Islands: CAS 14968 (1). Society Islands: SU 5357 (1,
187, holotype of Emmydrichthys vulcanus). CAS 14949 (1, 135), Tahiti,
Tavavo, J. E. Randall, 21 April 1956. CAS 14955 (1, 47.4), Moorea, Faatoai
village at entrance to Papetoai Bay, J. E. Randall and party, 30 June 1956.
Plus CAS 14950 (3) and CAS 14983 (1). Tuamotu Islands: CAS 14946 (2)
and CAS 14945 (1).

NEw CALEDONIA
USNM 208132 (1), Noumea.

NEw GUINEA
USNM 30516 (1).

AUSTRALIA

CAS 13760 (1, 129), Capricorn Islands, One Tree Island, west side, reef flat,
caught under stone on reef crest, F. McMichael, 20 November 1969. Plus CAS
14944 (1), Fairfax Island.

RED SEA
HUJ uncataloged (101 mm. S.L.).

DISTINGUISHING FEATURES. Dorsal XIJ-XIV, usually XIII + 5'%-7%,
usually 612; anal III + 5’2-6%, usually 5%; pectoral 17-19, usually 18; pelvic
I+ 5; all soft rays usually branched, covered with thick skin; head depressed;
eyes only slightly elevated, far apart, and with a deep depression between;
occipital area elevated, bordered laterally by a pit lying behind each eye; small
pit below and before eyes, pit smaller than orbit. Body covered with thick
skin and warts.

DescriPTIon. Dorsal fin normally with 13 spines; spines nearly same
length, about third through fifth the longest, covered by thick skin. Dorsal
soft rays 52-77%, usually 62, branched distally. Anal fin with 3 spines and
5% (rarely 6'’2) soft rays, soft rays branched distally, spines progressing in
length from first to third. Pectoral fin rays 18-19, usually 18; all rays
branched distally but dissection may be necessary to discern in lower fleshy
pectoral rays, rays unbranched in very small specimens. Pelvic fin with 1
spine and 5, or rarely 4, soft rays. Gill rakers rudimentary, total 8-11, 1-4 on

VoLt. XXXIX] ESCHMEYER & RAMA RAO: NEW STONEFISHES 363

upper arch, 6-8 on lower arch. Lateral line difficult to distinguish from warts,
usually 8-10 pores present. Vertebrae 24.

Body shape and coloration as in figure 8. Head large, broad, and depressed;
eyes very slightly elevated, far apart, and with a deep pit between them; occipital
area elevated, without a pit; a pit present behind each eye lateral to occiput,
deepest below parietal and nuchal spinous ridge; a small, more or less U-
shaped pit, less than orbit diameter, below and in front of eyes. Preorbital
bone covered by thick skin, usually with two diverging spines over maxillary.
Suborbital ridge with one large lump in middle under eye. Supplemental pre-
opercular spine usually absent, first and second preopercular spines large, third
sometimes present, fourth and fifth absent. Other head spines indistinct, or
developed as lumps or ridges, or absent. Small teeth on jaws, none on vomer
or palatine.

Coloration variable. In preservative mostly brown (fig. 8). Pectoral, pelvic,
and caudal fins tipped with white. Caudal fin with a subterminal dark band.
Paler areas on body, usually well marked between soft dorsal and anal fin.

COMPARISONS. Synanceia verrucosa is most like S. nana in shape and loca-
tion of pits on the head, but S. verrucosa is easily separated from S. nana by
having more pectoral rays (18-19 versus 14). Synanceia verrucosa lacks vomer-
ine teeth, while the other species of the genus have small teeth on the vomer.
Synanceia verrucosa may be separated from the other species of the genus by
the characters presented in the key.

DISTRIBUTION. Synanceia verrucosa is the most widespread stonefish and is
known from throughout the Indo-West Pacific faunal region from the Red Sea,
eastern Africa east to Tahiti, and from Australia north to Japan. It is found
in shallow water among coral reefs and coral rubble.

Genus Erosa Swainson
Erosa SWAINSON, 1839, p. 61 (type-species Synanceia erosa Langsdorf, understood from
text).

Bufichthys Swatnson, 1839, p. 181 (type-species S. erosa, by monotypy [see ‘Nomenclatural
remarks’]).

Synanchia, SwAINSON, 1839, p. 268, not pp. 61 or 280-281 (misidentification or misprint
[see ‘Nomenclatural remarks’]).

Synanchia, BLEEKER, 1874, pp. 3-4 (as a genus for ‘erosa’; after Swainson, 1839, p. 268 [see
‘Nomenclatural remarks’]).

Erosa, JorDAN and Starks, 1904, p. 156 (described as a “new genus” after Swainson;
wrongly thought Swainson’s name had no nomenclatural standing).

NOMENCLATURAL REMARKS. See ‘Nomenclatural remarks’ under the genus
Synanceia.
GENERIC DIAGNOSIS. See ‘Distinguishing features’ under the species account.

364 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Erosa erosa (Langsdorf).
(Figure 9; tables 3-5.)

Synanceia erosa LANGSDORF in Cuvier and Valenciennes, 1829, pp. 459-460 (original descrip-
tion; type locality Japan); Cuvier, 1837, p. vii, fig. 3 on Ichthyology pl. 21 (color
plate); TEMMINCK and SCHLEGEL, 1843, p. 45, fig. 1 on pl. xvii (brief description;
Japan); BorsreMan, 1947, pp. 54-55 (Burger and Von Siebold specimens from Japan;
equals Erosa erosa).

Synanchia erosa, SWAINSON, 1839, p. 268 (name only; Synanchia in error for Bufichthys
[see ‘Nomenclatural remarks’ under genus Synanceia]).

Synancidium erosum, GUNTHER, 1860, p. 146 (brief description; BMNH specimens) ; STEIN-
DACHNER and DODERLEIN, 1884, p. 199 (short description; Tokyo and Kagoshima) ;
Nystrom, 1887, p. 19 (description; one specimen from Japan); ISHIKAWA and Mat-
SUURA, 1897, p. 49 (listed; Kagoshima, Japan).

Erosa erosa, JORDAN and Starks, 1904, pp. 156-158, fig. 16 (synonymy; description; varia-
tion in coloration; Japan); FRANz, 1910, p. 74 (specimens from Japan); JORDAN and
TuHompson, 1914, p. 276, fig. 46 (listed; Japan; figure from Jordan and Starks, 1904) ;
McCuttocu, 1921, pp. 177-178 (E. iridea a synonym; description; figure of holotype
of E. ividea); JoRDAN and Hupss, 1925, p. 275 (specimens from Japan) ; McCULLOcH,
1929, p. 392 (listed; HE. wvidea a synonym); ScHMuipT, 1931, p. 111 (specimen from
Japan) ; FowLer, 1938b, p. 199 (listed from Malaya); MAtTsuBarA, 1943, pp. 422-424
(synonymy; description; internal features; specimens from Japan).

Erosa fratrum Octtey, 1910a, p. 32 (original description; type locality Moreton Bay, coast
of southern Queensland, Australia); McCuLtocu, 1929, p. 392 (listed; Australia) ;
WHITLEY, 1964, p. 57 (listed; Australia).

Erosa iridea OcttBy, 1910b, p. 113 (original description; type locality 19 miles N., 30°W.,
from Double Island Point, in 33 fathoms, coast of southern Queensland, Australia).

MATERIAL EXAMINED. (Counts were not taken on specimens with standard
length omitted and locality information abbreviated.)

JAPAN
SU 7385 (1, 98.3), Misaki, Jordan and Snyder. SU 7389 (1, 85.5), Nagasaki,
Hizen, Jordan and Snyder. SU 7197 (3, 77.7-109), Odowara. FSUT 7550 (1,
94.7), Wakayama. FSUT 32013 (1, 111), Nagasaki. FSUT 32263 (1, 48.4),
Nagasaki. FSUT 34630 (1, 131), Kochi. FSUT 34633 (1, 69.0), Kochi. Plus
USNM 51342 (1), USNM 59714 (1), USNM 75919 (2), and USNM 57650
(ye

SOUTH CHINA SEA
CAS 14800 (1, 91.0), 20°04’N., 111°58’E., R. L. Bolin, 20 July 1958. CAS
14797 (1, 85.2), 20°02’30’N., 113°32’E., F. D. Ommanney, 25 June 1958.
CAS 14799 (1, 111), Formosa Strait, just south of Formosa Banks to Pesca-
dores Island, about 30-50 meters, trawl, F. B. Steiner, 5 May 1972.

PHILIPPINES
USNM 168213 (1), southern Luzon.

Vor. XXXIX] ESCHMEYER & RAMA RAO: NEW STONEFISHES 365

AMBOINA

ZMK uncatalogued (1, 30.3), Bugten, in about 50 fathoms, T. Mortensen, 21
February 1922.

AUSTRALIA

QMB 13/1571 (about 97.0, ?holotype of Erosa fratrum), Moreton Bay,
Queensland. AMS E2943 (1, 63.0, ?holotype of Erosa iridea), 19 miles N., 30°
W. from Double Island Point, in 33 fathoms, Queensland.

DISTINGUISHING FEATURES. Dorsal XIV + 5%4-6%, usually 6%; anal III
+ 5%-6%, usually 5%; pectoral 14-16, usually 15; pelvic I+ 4; all fin rays
usually branched; no palatine teeth; head very large, globular; mouth terminal,
only slightly oblique; eyes large and on lateral surface of head; a strong bony
ridge connects orbits, followed on occiput with a deep pit; warts behind head,
not well marked on body posteriorly.

DEscRIPTION. Dorsal fin with 14 spines, all about same length. Dorsal soft
rays 5% or 6%, usually 6’2 (without close examination would probably count 7
rather than 6% soft rays). Anal fin with 3 spines, first short, second twice first,
third longest. Anal soft rays 5’2—6', usually 5’2 (appearing as 6). Pectoral fin
short, 14-16 (usually 15) branched rays. Pelvic fin with 1 spine and 4 soft
rays. Gill rakers rudimentary, total 10-13, 3-4 on upper arch and 7-9 on lower
arch. Lateral line tubes 10-11, including one on caudal fin. Some warts on
body, most conspicuous behind head. Vertebrae usually 25 (24-26).

Body shape and coloration as in figure 9. Head round; mouth terminal,
slightly oblique; eyes not elevated, on side of head, pointing outwards. A strong
ridge connects the orbits posteriorly; ridge followed by deep square to rectangu-
lar occipital pit; pit also present before ridge connecting orbits but occupied by
ascending arms of premaxillaries. No pit below eyes or below parietal and nuchal
spines. Preorbital bone with 2 broad spines over maxillary. Suborbital ridge
without distinct spines, raised in middle. Five blunt preopercular spines present,
plus 2 lumps on dentary and 3 spines near bases of upper preopercular spines.
Large cleithral spine, two blunt opercular spines. Other spines indistinct, de-
veloped as lumps. Small teeth on jaws and vomer, none on palatines.

In preservative (fig. 9) body brown or black on a pallid background; area
behind head and below soft dorsal fin darkest. Pectoral fin streaked and mot-
tled with brown or black, most lines enclosing circles, a white patch at midheight
of upper pectoral rays; anal fin mostly pallid with streaks and circular rings of
dusky pigment; caudal fin pallid, crossed by about 6 vertical narrow dark bands
(or double bands). Body most noticeably pallid at bases of middle dorsal spines
and below middle of spinous dorsal fin.

CoMPARISONS. The genus Evosa is most like Dampierosa in body shape and
shares with it (but not with Synanceia) the following features: eyes lateral,
mouth only slightly oblique, head globular and not depressed. Fvosa differs

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Vor. XXXIX] ESCHMEYVER & RAMA RAO: NEW STONEFISHES 367

from Dampierosa in having a lower soft dorsal fin ray count (5%2-6% versus
9%), 14 rather than 12 dorsal spines, and 3 rather than 2 anal spines.

DISTRIBUTION. rosa erosa occurs from Japan south to Australia; specific
localities represented by our material and records in the literature are Japan,
the South China Sea area, the Philippine Islands, Amboina, and eastern
Australia.

Genus Dampierosa Whitley

Dampierosa WHITLEY, 1932b, p. 346 (type-species Dampierosa daruma by original designa-
tion; monotypic).

GENERIC DIAGNOSIS. See ‘Distinguishing features’ under the species account
below.

Dampierosa daruma Whitley.

(Figure 10; tables 3-5.)

Dampierosa daruma WHITLEY, 1932b, pp. 346-347, figs. 2-3 on pl. XXXVIII (original de-
scription; type locality northwestern Australia, dredged off Broome, 1931, collector R.
Bourne; holotype AMS IA5116); 1964, p. 57 (listed).

Erosa daruma, MEEs, 1960, p. 19 (one specimen from Roeburne, Western Australia; dorsal
XIII + 8; anal I+ 7 (or III +5); pectoral 12; placed in genus Erosa).

MATERIAL EXAMINED. AMS JA5116 (97.2 mm. S.L., about 119 mm. T.L.,
holotype), locality as given above. Plus one specimen briefly examined,
C.S.I.R.O. Marine Laboratory, Munroe Collection C2766 (53.0 mm. S.L.), from
Exmouth Gulf.

DISTINGUISHING FEATURES. Dorsal XII + 9% (?XIII + 8%); anal II +
6’2; pectoral 12; pelvic I+ 4; no palatine teeth; head large, globular; mouth
terminal, slightly oblique; strong bony ridge connects orbits, followed on occi-
put by a deep pit; body with warts.

DESCRIPTION. (Based only on the holotype. A more complete description
is given by Whitley.) Dorsal fin with 12 spines and 9'% soft rays (first soft
ray segmented slightly, counted by Whitley (1932b) as a spine). Anal fin with
2 spines and 6% soft rays. Dorsal and anal spines fairly weak. Pectoral fin
short, with 12 rays. Pelvic fin with 1 spine and 4 soft rays. Gill rakers as given
by Whitley: 7 or 8 short, rounded, thick gill rakers on first arch. Lateral line
tubes 13. Body covered with warts. Vertebral count not available.

Body shape and coloration as in figure 10. Head globular; mouth terminal,
slightly oblique; eyes not elevated, on side of head, directed outwards. A strong
ridge connects the orbits; ridge followed by deep occipital pit. No pit below
eyes or below parietal and nuchal spines. Preorbital bone with 2 large lumps
over maxillary. Suborbital ridge without spines. Preopercle spines prominent.
(No information available on other spines.) Small teeth on jaws and vomer,
none on palatines.

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Vor. XXXIX] ESCHMEVER & RAMA RAO: NEW STONEFISHES 369

Coloration as given by Whitley (1932b, p. 347) is as follows:

“General colour in alcohol dark purplish-brown, irregular in tone and broken up
by the lighter papillae and raised cephalic surfaces. Interorbital and pterotic regions
white. Light brown mottling on lower surface of head and on parts of the body
below the spinous and soft dorsal fins. Dorsal dark brown anteriorly, but mottled
yellowish on the middle and posterior spines. Soft dorsal dark brownish with a
narrow margin of yellow and a broad oblique median band of yellow. Anal similar
to soft dorsal. Pectoral dark brownish with a yellowish band partly encircling its
base, a broader band crossing the rays to form large ocelli below and a distal mar-
gin of yellowish. Ventrals dark brown with two bands of yellowish and a similarly
coloured spot on the proximal part of the last ray. Caudal dark brown, crossed by
a broad median band of yellowish and with a broad margin of the same colour.”

CoMPARISONS. See ‘Remarks’ below.

DISTRIBUTION. Dampierosa daruma apparently is known only from three
localities off northwestern Australia.

Remarks. The holotype was briefly examined by the first author. We
believe that additional study will show that Dampierosa should be regarded as
a synonym of Frosa, even though the species differ substantially in counts of
dorsal and anal fin rays. The head shape is nearly identical, sharing the pres-
ence of a strong ridge joining the orbits, with a small pit before, and a large
occipital pit after this ridge. The eyes and mouth are similarly located in the
single species in each genus. In most scorpaenid fishes fin ray counts are fairly
stable for a given genus, but in the stonefishes the counts are variable in appar-
ently closely related species, and, as discussed under the subfamily section,
there appears to be a trend towards an increase of dorsal spines at the expense
of the dorsal soft rays in species of this subfamily. The presence of 2 anal
spines in Dampierosa and 3 (the usual condition in scorpaenids) in Erosa is not
regarded as a major difference; in Dampierosa the third anal spine has become
a segmented ray. Mees (1960, p. 19) gives anal rays as 1+ 7 or III +5 for 1
specimen of Dampierosa. A clarification of the dorsal and anal fin ray counts
in available specimens of Dampierosa is needed, and internal features of Evosa
and Dampierosa should be compared.

Genus Pseudosynanceia Day
Pseudosynanceia DAy, 1875, p. 163 (type-species Pseudosynanceia melanostigma Day, by

monotypy).

GENERIC DIAGNOsIS. See ‘Distinguishing features’ under the species account
below.

Pseudosynanceia melanostigma Day.
(Figure 11; tables 3-5.)

Pseudosynanceia melanostigma Day, 1875, p. 163, fig. 6 on pl. LV (original description;
excellent figure of holotype; type locality Karachi, Pakistan; holotype ZSI 1761).

370 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Leptosynanceia melanostigma, DAY, 1888, p. 788 (placed in genus Leptosynanceia) ; BLEGVAD
and L@pPENTHIN, 1944, p. 193 (brief description; coloration; specimens from Iranian
Gulf) ; Kuarar, 1961, p. 125 (description; counts as follows: dorsal XV—XVI + 5-6,
anal III + 7, pectoral 14-15, ventral I-++ 2; wrongly states no lateral line; from Fao,
Iraq).

MATERIAL EXAMINED.

PERSIAN GULF

USNM 196472 (1, 51.6), ESE. of Abu Ali, C. E. Dawson, sta. 13, 13 October
1956.

WEST PAKISTAN

USNM 199671 (1, 90), near Karachi, received from El Husseini, 24 January
1966. ZSI 1761 (1, 127, holotype of Psewdosynanceia melanostigma), Karachi.
AMS B8183 (1, about 115), Karachi, purchased from Day in 1885 [This speci-
men listed as type in AMS records but it is not]. SU 62409 (1, 75.7), vicinity
of Karachi, M.A. El Husseini, 1963.

INDIA

CAS 14801 (1, 77.8), western India, north side of Okha Point, tide pool, Inter-
national Indian Ocean Expedition, Anton Bruun Cruise No. 1, 9 March 1963.
USNM 209423 (1, 53.9), western India, 22°54’N., 68°36’E., in 15.5 meters,
International Indian Ocean Expedition, Anton Bruun Cruise No. 4B, sta. 223A,
19 November 1963.

DISTINGUISHING FEATURES. Dorsal XV—XVII + 4-6, usually XVI + 4;
anal III + 7-8; pectoral 14-15; pelvic I+ 3; all soft rays including pectoral
rays unbranched; head depressed; eyes on dorsal surface, pointing mostly up-
wards; mouth superior; no deep pits on head; body without prominent warts.

DEscRIPTION. Dorsal fin with 15-17 spines, all spines slender but with
prominent venom glands, all about same length. Dorsal soft rays 4—6, last single.
Anal fin with 3 spines of nearly same length. Anal soft rays 7-8, last single.
Pectoral fin long, reaching to over anal fin spines, with 14-16 (usually 15) un-
branched rays. Pelvic fin small, with 1 spine and 3 unbranched soft rays. Gill
rakers rudimentary, total 7-10, with 1-3 on upper arch and 6—7 on lower arch.
Lateral line not visible to unaided eye (with magnification a subsurface canal
usually can be seen at the normal location of the lateral line or slightly above;
small raised papillae with pores are also scattered on the body, some forming a
row below the dorsal fin, another group usually can be seen above the anal fin,
parallel to the lateral line).

Body shape and coloration as in figure 11. Head depressed, with mouth
superior; eyes small, on dorsal surface, pointing mostly upwards, at least 4 eye
diameters apart. Head with scattered low ridges, no deep pits on head. Head
spines poorly developed. Preorbital bone with 2 lumps over maxillary. Four

371

Vor. XXXIX] ESCHMEYER & RAMA RAO: NEW STONEFISHES

(apis We] UO poz} wourtoeds)

‘Ue ISTYeA ISAM

“TS ‘wut 9/ “60rZ9 AS ‘I[Npe ‘vus7ysounjam piaauUDUksopnasg

‘TT aunony

372 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

preopercular spines present. Other spines developed as ridges or absent. Small
teeth on jaws and vomer, none on palatines. Vertebrae 26 (2 specimens) or
24° (Gl).

Coloration in preservative (fig. 11) brown or gray to black on a pallid back-
ground, lighter ventrally. Body darkest on caudal peduncle, then abruptly white
at base of caudal fin. A broad subterminal brown or black bar on caudal fin,
otherwise this fin pallid. A clear area on anterior part of soft dorsal fin extend-
ing onto body. Soft dorsal fin dark distally. Spinous dorsal fin with dusky pig-
ment concentrated near distal part of spines. Pectoral fin mostly pallid with a
broad dark bar distally, fin rays tipped with white. Pectoral axil mostly pallid,
with a few dark specks or reticulations usually present. Anal fin mostly dusky,
darkest posteriorly. Pelvic fin rays pallid below, dark distally, with tips of rays
pallid. In life, pallid areas are yellow, according to Blevad and L¢ppenthin
(1944) and Khalaf (1961).

CoMPARISONS. See the ‘Comparisons’ section for Leptosynanceia asteroblepa.

DisTRIBUTION. This species is restricted in distribution; it is known only
from the Persian Gulf to western India. It is a marine and estuarine species
living on mud bottom.

Genus Trachicephalus Swainson

Trachicephalus Swainson, 1839, pp. 181, 268 (type-species Synanceia elongatus by original
designation).

Trichophasia SWAINSON, 1839, p. 61 (type-species Synanceia elongatus implied from text [see
‘Nomenclatural remarks’ ].

Polycaulus GUNTHER, 1860, p. 175 (proposed as a replacement name for Trachicephalus
Swainson) ; BLEEKER, 1874, p. 19 (description; nomenclature).

Uranoblepus GiLL, 1861, footnote on p. 5 (proposed as a replacement name for Trachi-
cephalus Swainson).

NOMENCLATURAL REMARKS. As discussed by Gill (1905, p. 224) the replace-
ment names Polycaulus and Uranoblepus were proposed because of the similarity
of Trachicephalus and the earlier Trachycephalus (e.g., the reptile genus Trachy-
cephalus Tschudi, 1838). Under the Rules of Zoological Nomenclature the two
are etymologically different and Trachicephalus is not preoccupied.

Both Trichophasia and Trachicephalus were proposed by Swainson (1839)
for the same category as evidenced from the text (p. 61, pp. 180-181, and p.
268) (see also our ‘Nomenclatural remarks’ under the genus Synanceia). Gill
(1905, p. 224) appears to be the first to mention both names, and he selected
Trachicephalus over Trichophasia.

Dracnosis. See ‘Distinguishing features’ in the species account below.

Trachicephalus uranoscopus (Bloch & Schneider).
(Figure 12; tables 3-5.)

Synanceja uranoscopa BLocH and SCHNEIDER, 1801, p. 195 (original description; type locality

Vor. XXXIX] ESCHMEYER & RAMA RAO: NEW STONEFISHES 373

Tranquebar, India; spelling of genus on p. 194 corrected to Synanceia on p. 573) ; CUVIER
in Cuvier and Valenciennes, 1829, pp. 458-459 (brief description; remarks).

Uranoscopus indicus KuuHt and VAN HASSELT in Cuvier and Valenciennes, 1829, p. 456
(nomen nudum); BLEEKER, 1849, p. 10 (as synonym of Synanceia elongata).

Synancya elongata CUVIER in Cuvier and Valenciennes, 1829, pp. 456-458 (original descrip-
tion; type locality Pondichery, India, and Java) ; BLEEKER, 1849, pp. 4, 10 (synonymy;
brief description; range) ; 1861b, p. 72 (listed; Penang).

Synanceia elongata, CUviER, 1837, p. viii, fig. 3 on Ichthyology pl. 27 (color plate).

Trachicephalus elongatus, SWAINSON, 1839, p. 268 (as type of Trachicephalus).

Synanceia breviceps RICHARDSON, 1845, pp. 71-72 (original description; type locality China) ;
WHITEHEAD, 1969, pp. 205, 207 (types lost).

Aploactis breviceps, RICHARDSON, 1846, p. 212 (placed in Aploactis; location of specimens).

Synancia elongata, CANTOR, 1850, p. 1029 (synonymy; description; range).

Uranoscopus adhesipinnis BiytH, 1860, p. 142 (original description; type locality India,
from Calcutta fish market [some counts of fin rays low, probably inaccurate]).

Polycaulus elongatus, GUNTHER, 1860, p. 174 (synonymy; description; osteology; collection
data) ; KNErR, 1865, p. 120 (synonymy; description; range); BLEEKER, 1874, pp. 20-21,
fig. 1 on pl. 2 (description; synonymy; range) ; 1879, fig. 2 on pl. CCCCXV (figure from
Bleeker, 1874) ; SEALE, 1910, p. 286 (specimens from Borneo).

Polycaulis uranoscopus, DAY, 1875, p. 164, fig. 6 on pl. XXXIX (synonymy; description;
range; good figure); Herre and Myers, 1937, p. 34 (2 specimens [SU 30872] from
Singapore).

Trachicephalus uranoscopus, RUTTER, 1897, p. 81 (good description; specimens from China
[SU 1758]) ; Fow er, 1929, p. 613 (listed; 4 specimens from Hong Kong) ; 1931, p. 305
(specimens from Hong Kong; coloration; status of Trachicephalus).

Polycaulus uranoscopus, FOWLER, 1935, p. 153 (one specimen from Bangkok, Thailand;
“agrees with Bleeker’s figure of Polycaulus elongatus”) ; DE BEAUFORT in Weber and de
Beaufort, 1962, pp. 102-103, fig. 29 (synonymy; description; distribution).

Trachycephalus uranoscopus, FOWLER, 1938a, p. 36 (two specimens from Tai Po, China, and
two from Hong Kong).

MATERIAL EXAMINED.

CHINA

SU 1758 (6, 44.8-56.5), Swatow, A. M. Fielde. SU 25743 (2, 70.7—80.0), Can-
ton, A. W. Herre.

Honc KoNG

SU 9853 (2, 47.0-63.1), P. L. Jouy. SU 39608 (2, 63.5-67.2), Aberdeen Fish
Market, A. W. Herre, 1 June 1937. Plus USNM 143296 (1).

SINGAPORE
SU 30872 (2, 19.6-53.8), A. W. Herre, 3 March 1934. SU 34090 (7, 37.0-62.3),

A. W. Herre, 8 May 1937. SU 39472 (1, 59.0), Siglap, A. W. Herre, 18 October
1940. Plus USNM 142947 (1).

THAILAND

USNM 119657 (1, 40.5), Laem Sing, at mouth of Chanthaburi River, H. M.
Smith, 17 July 1928.

374 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Ficure 12. Trachycephalus uranoscopus, adult, India; figure from Day, 1875, fig. 6 on
pl. 39.

INDIA

SU 14671 (2, 36.0-45.0), Marmagao Bay, S. Kemp. SU 14672 (2, 68.8-72.2),
Madras, Ennur Fisheries Station, A. W. Herre, 6 January 1941; MNHN A905
(3, 52.3-54.0, syntypes of S. elongata), Coramandel Coast, collected by
Leschenault.

DISTINGUISHING FEATURES. Dorsal XI—XIII + 12-14, usually XII + 12;
anal II + 12-14, usually II + 13; pectoral 14-15; pelvic I+ 5; all soft rays
including pectoral rays unbranched; head depressed; eyes on dorsal surface of
head, pointing mostly upwards; mouth superior; no deep pits on head; body
without prominent warts.

DescriPTION. Dorsal fin with 12 (11-13) spines, all spines about same
length, flexible, difficult to distinguish from soft rays. Dorsal soft rays 12-14,
usually 12, with last single. Pectoral fin moderate, reaching to level of anal
spines, with 14-15 unbranched rays. Anal fin with 2 spines and 12-14, usually
13, soft rays, last soft ray single. Pelvic fin base very long, reaching to level of
vent, with 1 spine and 5 unbranched soft rays. Gill rakers rudimentary, total
7-8, 2-3 on upper arch, 5 on lower arch. Lateral line difficult to observe with-
out microscope, runs high up on flanks; total tubes about 12—14, anterior ones
more prominent and with small bilobed flaps; additional papillae (with pores? )
scattered on body, a row present on body above anal fin and another above
lateral line.

Body shape and coloration as in figure 12. Head depressed, with superior
mouth; eyes small and on dorsal surface of head, pointing upwards and out-
wards, about 3 eye diameters apart. Head with scattered low ridges; two ridges
run medially from eye, shallow occipital pit bordered by ridges, pit present be-
tween eyes but occupied by ascending arms of premaxillaries; another ridge
passes under eye; no deep pits on head. Head spines poorly developed. Pre-
orbital bone with 2 or 3 short spines over maxillary. Preopercular bone with 4
blunt spines. Other spines developed as lumps, ridges, or absent. Small teeth on
jaws and vomer, none on palatines. Vertebrae 28-30, mostly 29.

Body mostly tan or brown in preservative (figure 12). Lateral line pores and

VoL. XXXIX] ESCHMEYER & RAMA RAO: NEW STONEFISHES as

other pores on body sometimes surrounded by white. Distal portions of soft dor-
sal, anal, caudal, and pectoral fins darker brown except caudal fin tipped with
white and with pale area on dorsal and on ventral margin of caudal at about
middle of fin (better marked in small specimens).

ComPaARISONS. See the ‘Comparisons’ section under Leptosynanceia astero-
blepa.

DistriBuTion. Tvrachicephalus uranoscopus occurs from western India to
China and south in the Malay Archipelago to Borneo. This species is associated
with mud bottom areas and some captures have been in estuaries.

Genus Leptosynanceia Bleeker

Leptosynanceia BLEEKER, 1874, p. 17 (type-species Synanceia asteroblepa Richardson by
monotypy).

Dracnosis. See ‘Distinguishing features’ under the species account below.

Leptosynanceia asteroblepa (Richardson).
(Figure 13; tables 3-5.)

Synanceia asteroblepa RicHARDSON, 1845, pp. 69-71, figs. 1-3 on pl. 39 (original description;
type locality New Guinea; compared with species of Synanceia); BLEEKER, 1849, p. 4
(listed; New Guinea) ; 1852a, p. 242 (listed; New Guinea) ; 1852b, pp. 419-420 (descrip-
tion; rivers and estuaries of Borneo) ; GUNTHER, 1860, p. 147 (compiled) ; 1868, p. 265
(listed; Sarawak) ; VINCIGUERRA, 1926, p. 539 (synonymy; listed; muddy rivers of Sara-
wak; range).

Leptosynanceia asteroblepa, BLEEKER, 1874, pp. 17-18, fig. 2 on pl. 4 (as type of a new genus;
description; specimens from Borneo; good figure); 1878, p. 49 (listed; New Guinea) ;
1879, fig. 6 on pl. CCCCXVI (figure from Bleeker, 1874) ; Fowier, 1928, p. 298, fig. 51
(brief synonymy and description; L. greenmani as a synonym); HERRE and Myers,
1937, p. 34 (description; specimen from Sumatra, 100 miles west of Singapore [SU
30870]) ; Munro, 1967, p. 539 (in key; New Guinea).

Leptosynanceia greenmani Fowler, 1905, pp. 507-510, fig. 12 (original description; type
locality Borneo; compared with Leptosynanceia asteroblepa; [holotype ANSP 114884,
2 paratypes ANSP 114885-6]).

MATERIAL EXAMINED.
BORNEO

ANSP 114884 (1 specimen, 117 mm. S.L., holotype of L. greenmani), Baram,
Borneo, A. C. Harrison, Jr., and H. M. Hiller, 1897; ANSP 114885 (1, 103,
paratype of L. greenmani), and ANSP 114886 (1, 87.2, paratype of L. green-
mani), Baram, Borneo, W. H. Furness, 1898.

SUMATRA

SU 30870 (1, 49.7), 100 mi. west of Singapore, A. W. Herre, 27 March 1934.

DISTINGUISHING FEATURES. Dorsal XVI + 5; anal IIJ-IV + 5-6; pectoral
13—15, usually 14; pelvic I + 4; all soft rays including pectoral rays unbranched;

376 CALIFORNIA ACADEMY OF SCIENCES [PRroc. 4TH Serr.

FicureE 13. Leptosynanceia asteroblepa, adult, New Guinea; figure from Richardson,
1844, fig. 1 on pl. 39.

head depressed; eyes on dorsal surface, directed mostly upwards; mouth superior ;
no deep pits on head; body without prominent warts.

DEscRIPTION. Dorsal fin with 16 spines, all spines about same length, firm
but of small diameter, with small venom glands at about midheight of spines.
Dorsal soft rays 5, unbranched, last single. Anal fin with 3—4 spines and 5-6
unbranched soft rays, last single. Pectoral fin reaching to over anal spines,
upper rays the longest, with 13-15 rays, all unbranched. Pelvic fin short, with 1
spine and 4 unbranched soft rays. Gill rakers rudimentary, 3 on upper arch, 8
or 9 on lower arch (in one specimen). Lateral line runs high up on back, with
11 tubes, last on caudal fin; additional papillae (with pores?) visible on body
with microscope, a row of them present above anal fin.

Body shape and coloration as in figure 13. Head depressed, with mouth
superior; eyes small and on dorsal surface of head, eyes pointing mostly up-
wards, about 3 eye diameters apart. Head with scattered low ridges; shallow
pits present but none prominent. Most head spines poorly developed. Preorbital
bone with 2 lumps over maxillary. Preopercular bone with 5 well marked spines.
Opercular spines well defined. Other spines developed as lumps or ridges or
absent. Small teeth on jaws, none on vomer and palatines. Vertebrae 28 (1
specimen ).

Specimens available to us mostly faded. Fowler (1905) reports body pale
brown, scarcely paler below, head finely mottled or marbled with darker brown,
and back and sides with numerous large deep brownish blotches. Fins also simi-
larly marked. Soft dorsal, anal, and caudal darker distally but with a white
margin. Bleeker (1874) illustrates a brownish-yellow fish, mottled with large
dark areas, fins dark distally, tinge of pink on fins.

VoL. XXXIX] ESCHMEYVER & RAMA RAO: NEW STONEFISHES Si]

ComPaARIsoNs. The genera Pseudosynanceia, Trachicephalus, and Lepto-
synanceia each contain a single species. These species are more elongate (and
resemble uranoscopid fishes in outward appearance) than the other more globular
species treated in this paper. The pelvic rays are I + 3 in Pseudosynanceia, I +
4 in Leptosynanceia, and 1+ 5 in Trachicephalus. Pseudosynanceia resembles
Leptosynanceia, and differs from Trachicephalus, in having a high dorsal spine
count (usually XVI + 4 in Pseudosynanceia, XVI + 5 in Leptosynanceia, and
XII + 12 in Trachicephalus). Significant differences in anal counts are also
found (III + 7-8 in Pseudosynanceia, IJ + 12-14 in Trachicephalus, and usu-
ally IV + 5 in Leptosynanceia). These external differences are extensive and
serve to distinguish the species from each other. A more detailed study is needed,
but it would appear that these differences warrant separate genera for these
species.

DISTRIBUTION. We know this species only from Singapore, Sumatra, Borneo,
and New Guinea. It is reported to inhabit estuaries and rivers.

LITERATURE CITED

Acassiz, L.

1845. Nomenclator zoologicus, continens nomina systematica generum animalium tam
viventium quam fossilium, secundum ordinem alphabeticum disposita, adjectis
auctoribus libris, in quibus referiuntur, anno editionis, etymologia et familiis, ad
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BLEEKER, P.

1849. Bijdrage tot de kennis der Scleroparei van den Soenda-Molukschen Archipel.
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1852b. Zesde bijdrage tot de kennis der ichthyologische fauna van Borneo. Natuurkundig
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1861. Iets over de vischfauna van het eiland Pinang. Verslagen en Mededeelingen der
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1863. Onziéme notice sur la faune ichthyologique de l’ile de Ternate. Nederlandsch
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378 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

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Vor. XXXIX] ESCHMEYER & RAMA RAO: NEW STONEFISHES 379

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ISHIKAWA, C., AND K. MAtsutraA
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PROCEEDINGS

OF THE

CALIFORNIA ACADEMY OF SCIENCES

FOURTH SERIES

Vol. XXXIX, No. 19, pp. 383-410; 7 figs.; 1 table. October 24, 1973

GOGOLIA FILEWOODI, A NEW GENUS
AND SPECIES OF SHARK FROM
NEW GUINEA
(CARCHARHINIFORMES: TRIAKIDAE),
WITH A REDEFINITION OF THE FAMILY
TRIAKIDAE AND A KEY TO
TRIAKID GENERA

By

L. J. V. Compagno
Department of Biological Sciences, Stanford University,
Stanford, California 94305

INTRODUCTION

On the night of July 18-19, 1970, Mr. Komet Kisokau handlined a peculiar
small shark in Astrolabe Bay, off the Gogol River near Madang, New Guinea.
Mr. Kisokau and Mr. R. T. Gibson (Fisheries Officer, Madang, and a former
shark fisherman) recognized the unusual nature of the specimen and sent it to
Mr. L. W. Filewood (Biologist-In-Charge of the Department of Agriculture,
Stock and Fisheries, Konedobu, Papau-New Guinea), who is currently studying
the elasmobranchs of the New Guinean region. Mr. Filewood discovered that
the specimen represented a new genus and species of triakid and intended to
describe it in collaboration with the present writer. Unfortunately Mr. File-
wood was unable to participate in this task and generously relinquished both
it and the specimen to me.

Gogolia Compagno, new genus

TYPE SPECIES. Gogolia filewoodi Compagno, new species.
DERIVATION OF NAME. Gogolia (considered feminine), from the Gogol River
of Northern New Guinea.

[383]

[Proc. 4TH Ser.

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384

‘TOO-8S891 ‘J ‘OU Wnasny, URTTRAYsNY ‘o[euLoy y[Mpe “WU HEY ‘adA}O[OY Jo MarA [e19}e'T ‘satsads puv snues mou ‘ousedwog spoomapy{ vy0S0H “| aunorg

VoL. XXXIX] COMPAGNO: GOGOLIA FILEWOODI 385

GENERIC DEFINITION. (Terminology follows Compagno, 1970; Compagno
and Springer, 1971; and Compagno, 1973a.). Triakid sharks with a moderately
long head, length from snout tip to level of 5th gill openings about 1.4-1.5 times
the distance between the dorsal fin bases and about 22—24 percent of total length.
Head and snout acutely wedge-shaped and blunt-tipped in lateral view (figs. 1,
2F). Outline of snout bell-shaped in dorsoventral view (fig. 2D—E) and para-
bolic in front of nostrils. Snout very long, preoral length (distance from snout
tip to upper symphysis) about 1.6—1.7 times mouth width and 2.1—2.2 in head
length. Subocular ridges very strong and prominently visible in dorsal view of
head (fig. 2D: SOR) and blocking off view of eyes ventrally (fig. 2E). Eyes
very large, elliptical, and about twice as long as high. Nictitating lower eyelid
external and having a diagonal edge (fig. 2A: NLE). Subocular pouch shallow
and entirely covered with denticles. Gill openings very small, width of 3rd about
2.2—5.2 in eye length. Anterior nasal flaps very low and virtually vestigial (fig.
2B: ANF), not barbel-like or greatly expanded. Posterior ends of anterior na-
sal flaps well forward of mouth (fig. 2E). No nasoral grooves. Nasal cavities
communicating to the exterior only by narrow incurrent and excurrent apertures
(fig. 2B: INA and EXA), not exposed ventrally. Internarial width about 2.0
times nostril widths. Nostrils about 2.5 times farther from the snout tip than
from the mouth. Mouth arcuate in shape (fig. 2E). Mouth very short, its length
from level of upper symphysis to level of mouth corners 2.5—3.0 in width of
mouth between corners. Lower jaw very flat, with its sides hardly protruding
or not showing below ventral surface of head in lateral view (figs. 1, 2F). Upper
labial furrows extending anteriorly to level of upper symphysis.

Tooth-row counts 40—-41/35 (2). 2-3/2-4 series of teeth functional. Pre-
medial edges of most teeth convex and undifferentiated, but in adult the medials
and some upper anteroposteriors have premedial cusplets or strong serrations
(figs. 3C—D, 4A). Primary cusps very strong on all teeth except for the last
lower posteriors (fig. 4B: teeth nos. 14-15). Postlateral edges of crowns
strongly notched in all cusped teeth, these differentiated basally into postlateral
cusplets or blades. Roots of teeth relatively low (fig. 3C-H: RT). Teeth
strongly compressed, sharp-edged, and blade-like, but not molariform. Teeth
not enlarged at symphysis to form a knob and not extending onto edges and
ventral surface of lower jaw. Tooth-row groups poorly differentiated and includ-
ing medials (fig. 4: M) and anteroposteriors (fig. 4A, teeth nos. 1-20; and fig.
4B, teeth nos. 1-15) in both jaws. Medials have erect or semioblique cusps and
relatively higher and narrower crowns than the lower, broader, oblique-cusped
anteroposteriors. Dignathic heterodonty strong in anteroposteriors near sym-
physis, but much weaker near the ends of the dental bands. In an adult these

4TH SER.

[ Proc.

MY OF SCIENCES

CALIFORNIA ACADE

386

VoL, XXXIX] COMPAGNO: GOGOLIA FILEWOODI 387

parasymphysial anteroposteriors have premedial cusplets or serrations, narrow
cusps, prominent transverse ridges or striations, and more postlateral cusplets
in the upper jaw, while lower teeth have no premedial cusplets and very few
postlateral ones, broader cusps, and no ridges (compare fig. 3C-E with F-H).

Interdorsal ridge present in adult and absent in fetus. Caudal peduncle short,
its length from second dorsal insertion to upper caudal origin about % of sec-
ond dorsal base. Lateral trunk denticles from dorsum below first dorsal fin with
teardrop-shaped crowns that are considerably longer than wide and have rudi-
mentary lateral cusps or none (fig. 3A-B).

Longest distal radials of pectoral fin skeleton about 1.6 times as long as long-
est proximal radials (fig. 5|A-B: DRA and PRA). Anterior margins of pelvic
fins about % as long as pectoral anterior margins.

First dorsal origin far anterior, varying from over pectoral insertions to
about over pectoral midbases. Free rear tip of first dorsal slightly posterior to
pelvic origins. First dorsal base extremely long (figs. 1, 2F), its length almost
equal to length of dorsal caudal margin, 2.3-3.2 times the first dorsal height,
and 1.3—1.4 times the distance between dorsal bases. Second dorsal much smaller
than first, its height about 7 of first dorsal height and its base length less than
half of first dorsal base. Anal fin smaller than second dorsal, its height 0.5—
0.6 of second dorsal height and its base 0.5—0.7 of second dorsal base. Anal
insertion slightly anterior to second dorsal insertion by a distance about 0.1—0.2
of second dorsal base.

Caudal fin with a short but strong ventral caudal lobe in adult but a weak
one in a fetus (figs. 1, 2F). Postventral caudal margin shallowly indented but
not deeply notched. Length of terminal caudal sector from subterminal notch
to tip of caudal about 3.1—3.8 in length of dorsal caudal margin.

Cranium with a long rostrum, with the length of the medial rostral cartilage
from its base to the anterior edge of the rostral node about 1.4—-1.9 in nasobasal
length (distance from the base of the medial rostral cartilage to the postero-
ventral edge of the occipital centrum, here used as an independent variable for
cranial proportions). Bases of lateral rostral cartilages connected by a ridge
(fig. 6A, C: RRF) to the edge of the anterior fontanelle. Nasal capsules trans-

Ficure 2. Gogolia filewoodi. A-E, photographs from 739 mm. holotype. A, left eye in
lateral view. B, left nostril in ventral view. C, valvular intestine cut longitudinally to show
spiral valve. D, head, dorsal view. E, head, ventral view. F, photograph in lateral view of
224 mm. fetus, CAS—27588. All scale lines equal 1 centimeter. Abbreviations: ANF, anterior
nasal flap; EXA, excurrent aperture; INA, incurrent aperture; MNF, mesonarial flap; NLE,
nictitating lower eyelid; PNF, posterior nasal flap; SLE, secondary lower eyelid; SOP, sub-
ocular pouch; SOR, subocular ridge; SP, spiracle; UE, upper eyelid.

388 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

ee +
BG ™N BL Bi

Ficure 3. Gogolia filewoodi. A, crowns of lateral trunk denticles from dorsum below
first dorsal fin, 739 mm. holotype. B, same from 224 mm. fetus. C—E, teeth from 739 mm.
holotype. Labial (C), lingual (D), and lateral (E) views of tooth from 4th upper anteropos-
terior row (numbered from symphysis). Labial (F), lingual (G), and lateral (H) views of
tooth from 4th lower anteroposterior row. Abbreviations: BG, basal groove; BL, basal
ledge; CR, crown; PC, primary cusp; PLAS, postlateral attachment surface of root; PLC,
postlateral cusplets; PMAS, premedial attachment surface of root; PMC, premedial cusplets;
PME, premedial edge; RT, root; TG, transverse groove; TN, transverse notch; TR, trans-
verse ridges.

versely elongated and anteriorly flattened, not circular (fig. 6: NC). Nasal
fontanelles delimited anteriorly from nasal apertures (fig. 6B: NA and NF).
Nasal openings of orbitonasal canals inside nasal capsules and well behind pos-
terior edges of nasal fontanelles. Thickened ridges or ectethmoid condyles (fig.
6B—C: ECN) present on nasal capsules and articulating with the orbital pro-
cesses of the palatoquadrates (fig. 7B—-C). A depression or subethmoid fossa
(fig. 6B: SEF) present on the ventral surface of the cranium between the ecteth-
moid condyles. Basal plate very narrow across orbital notches (fig. 6B: NP),
its transverse width there about 3.0 in nasobasal length. Two pairs of arterial
foramina present on basal plate, one for the internal carotid arteries and one
for the stapedial or orbital arteries (fig. 6B: FC and FS). Least width across
supraorbital crests (transverse to the long axis of the cranium) about 2.4 in
nasobasal length. Postorbital processes short and shallowly notched distally
(fig. 6: PT). Suborbital shelves narrow and not laterally expanded, ear-shaped,
fenestrate, or notched. Greatest width across suborbital shelves (perpendicular
to the long axis of cranium) about 1.7 in nasobasal length.

Vor. XXXIX] COMPAGNO: GOGOLIA FILEWOODI 389

Total vertebral counts 180 (2). No stutter zone of alternating long and
short centra in vertebral column. Thoracic vertebral centra with strong diagonal
calcified lamellae (fig. 5D-E: DCL).

Other characteristics of Gogolia those of the family Triakidae and of the
single species (see below).

FAMILIAL CLASSIFICATION OF GOGOLIA

Gogolia is a genus that, with Hemitriakis and Jago, stands midway between
the families Carcharhinidae and Triakidae (or their equivalents) as defined
by Garman (1913), White (1936, 1937), Bigelow and Schroeder (1948), and
Garrick and Schultz (1963). According to these workers, triakids have several
functional series of small, molariform or blade-like and multicuspidate teeth
and ‘nictitating folds,’ whereas carcharhinids have not more than 1 or 2 series
of small to large, blade-like teeth and ‘nictitating membranes.’ The failure of
these characters to distinguish the two families is discussed elsewhere (Com-
pagno, 1970). Gogolia itself has a ‘nictitating fold’ but has small, blade-like
teeth in 2—4 functional series.

Previously I resolved the problem of triakid-carcharhinid intergradation by
the unsatisfactory expedient of merging the two families (Compagno, 1970).
Subsequent work on a revision of carcharhinoid genera indicated that a group
of genera partially corresponding to the family Triakidae of White (1937) could
be separated from the Carcharhinidae.

Part of the difficulty in separating the families Carcharhinidae and Triaki-
dae of White (1937) and Bigelow and Schroeder (1948) was that each of the
two families contained genera closer to the type genus of the other family than
to its own type genus. Thus Triaenodon has many external, cranial, pectoral
fin skeleton, and cephalic myological characters that ally it closely with typical
carcharhinids such as Carcharhinus, but the strong cusplets on its teeth led many
workers to place 77iaenodon incorrectly in the Triakidae. Galeorhinus has often
been placed in the Carcharhinidae but it similarly has many characters that
indicate close affinities with the Triakidae. Also, Galeorhinus is linked with the
genera Tviakis and Mustelus by a group of roughly intermediate forms (Gogolia,
Hypogaleus, lago, Furgaleus, and Hemitriakis).

The reinstated family Triakidae includes the genera Mustelus Linck, 1790;
Galeorhinus Blainville, 1816; Triakis Miller and Henle, 1838a; Scylliogaleus
Boulenger, 1902; Hemitriakis Herre, 1923; Furgaleus Whitley, 1951; Hypo-
galeus J. L. B. Smith, 1957a; Jago Compagno and Springer, 1971; Allomycter
Guitart, 1972; and Gogolia. Triaenodon Miller and Henle, 1837 is transferred
to the Carcharhinidae (see also Gohar and Mazhar, 1964, and Kato, Springer,
and Wagner, 1967, for discussion of the familial position of Triaenodon). The
“scyliorhiniform triakoids” (Compagno, 1970) have often been placed in the

390 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

family Triakidae or its equivalents and included in the genus Tviakis. How-
ever, these genera, including Proscyllium Hilgendorf, 1904, Eridacnis H. M.
Smith, 1913, Ctenacis Compagno, 1973a, and Gollum Compagno, 1973a, are
sufficiently distinct from triakids to require a separate family, Proscylliidae
(elevation of subfamily Proscylliinae Fowler, 1941). Leptocharias A. Smith, in
Miiller and Henle, 1838a is difficult to place but probably is best placed in its
own family, Leptochariidae (elevation of tribe Leptochariana Gray, 1851) owing
to its morphological distinctness from other carcharhinoids. Leptocharias has
often been grouped with the triakids in proximity to Triaenodon, but these two
genera are not closely related.
The Triakidae can be defined as follows:

Family TRIAk1paE Gray, 1851, new rank

Subfamily Musteli BonAPARTE, 1838, p. 199 (Family Squalidae. Type genus: Mustelus Cuvier,
1817, equals Mustelus Linck, 1790.). Preoccupied in Mammalia by families Mustelini
Fischer, 1817 and Mustelidae Swainson, 1835 (Type genus: Mustela Linnaeus, 1758) ;
references for mammalian families in Simpson (1945).

Family Galei MULLER and HENLE, 1839, p. 57 (Type genus: Galews Cuvier, 1817, a junior
synonym of Galeorhinus Blainville, 1816; not Galeus Rafinesque, 1809.).

Family Scylliodontes MULLER and HENTE, 1839, p. 63 (Not based on a type genus.).

Tribe Triakiana Gray, 1851, p. 39 (Family Squalidae. Type genus: Triakis Miller and Henle,
1838a.). The family Triakidae was independently proposed by White (1936a, 1936b,
1937).

Family Galeorhinoidae Git, 1862b, p. 393 (Type genus: Galeorhinus Blainville, 1816.).

Family Scylliogaleidae WHITLEY, 1940, p. 68 (Type genus: Scylliogaleus Boulenger, 1902.).
The family Scylliogaleidae was independently proposed by Smith (1957b).

Family Emissolidae WHITLEY, 1940, p. 68 (Type genus: Emissola Jarocki, 1822, probably
a junior synonym of Mustelus Linck, 1790.).

Family Eugaleidae Gurr, 1962, p. 428 (Type genus: Eugaleus “Rafinesque, 1810”, equals
Gill, 1864, not Rafinesque, 1809-1810; a junior synonym of Galeorhinus Blainville, 1816.).

Carcharhinoid sharks with the head not expanded laterally into a wing-like
blade. Eyes high on sides of head, with their ventral edges above the nostrils.
Eye length 1.5—2.5 or more times eye height. Nictitating lower eyelid variably
external, transitional, or internal (Compagno, 1970). Small to moderately
large spiracles present, with their greatest widths about 3—10 in eye length. La-
bial furrows long and present on both jaws. Labial cartilages well developed.

Tooth-row groups poorly differentiated or absent, with medials often present
but no symphysials, anteriors, or well defined posteriors. Posterior teeth not
comb-like. Gynandric heterodonty absent or poorly developed as far as is known
(data not available for Scylliogaleus, Gogolia, and Allomycter). Tooth-row
counts 18—94/27-94. Teeth with strong basal ledges and grooves.

Precaudal pits absent. Pectoral fin skeleton with its radials projecting half
of pectoral anterior margin length or less into fin (fig. 5A). Distal pectoral

VoL. XXXIX] COMPAGNO: GOGOLIA FILEWOODI 391

radials with parallel edges and truncate or distally rounded tips. Longest distal
radials 1-2 times as long as longest proximal radials.

First dorsal fin with distinct apex and separate anterior and posterior mar-
gins, not arcuate-edged and keel-like. First dorsal base anterior to pelvic bases.
Length of first dorsal base usually much shorter than dorsal caudal margin, but
subequal to it in Gogolia. No undulations in dorsal caudal margin.

Neurocranium with bases of lateral rostral cartilages well separated. Nasal
capsules not greatly depressed or transversely expanded (fig. 6). Large nasal
fontanelles present and broadly continuous anteriorly with the nasal apertures
(fig. 6B: NA and NF). Nasal openings of orbitonasal canals inside the nasal
cavities, located at the posterior edges of the nasal fontanelles or behind them.
Ectethmoid condyles, when present, without foramina for the anterior facial
veins (ectethmoid foramina). Internasal septum a high, compressed plate. Ar-
terial foramina on the basal plate usually include two pairs of stapedial and in-
ternal carotid foramina (fig. 6B: FS and FC), but Furgaleus has a single pair
of common foramina apparently formed by the merging of the stapedials and
internal carotids on each side. Parietal fossa single. Supraorbital crest present
ies 6205SC).

Wedge-like intermedialia strongly developed in vertebral centra of adults
and subadults (fig. 5E) but often absent or poorly developed in fetuses or new-
born specimens (fig. 5D).

Valvular intestine with a spiral valve of 5-11 turns. Levator palatoquadrati
muscles small, with their origins not expanded anterior to the postorbital pro-
cesses (fig. 7C: MLP).

Triakidae is distinguished from the families Scyliorhinidae, Pseudo-
triakidae, and Sphyrnidae of Bigelow and Schroeder (1948) by many charac-
ters that need not be detailed here. The Proscylliidae as delimited above differ
from the Triakidae in having rudimentary nictitating lower eyelids; very short
labial furrows (sometimes absent in Eridacnis); comb-like posterior teeth;
transverse grooves absent from teeth; distal pectoral radials much shorter than
the proximal radials; and wedge-like intermedialia absent from the vertebral
centra. The Leptochariidae differ from the Triakidae in having teeth without
transverse grooves; strongly developed gynandric heterodonty; nasal openings
of the orbitonasal canals not in the nasal cavities but posterior to them and pene-
trating the suborbital shelves; no supraorbital crest; and 14—-16 turns in the
spiral intestinal valve.

The family Carcharhinidae is restricted to the ‘advanced carcharhinids”’
of Compagno (1970) and includes Carcharhinus Blainville, 1816; Scoliodon
Miller and Henle, 1837; Galeocerdo Miiller and Henle, 1837; Triaenodon Miil-
ler and Henle, 1837; Loxodon Miiller and Henle, 1838a; Hypoprion Miiller and
Henle, 1839; Prionace Cantor, 1849; Aprionodon Gill, 1862a; Jsogomphodon

392 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Gill, 1862b; Lamiopsis Gill, 1862b; Rhizoprionodon Whitley, 1929; and
Negaprion Whitley, 1940. The “intermediate carcharhinids” of Compagno
(1970) are sufficiently distinct from the Carcharhinidae to rate a family, Hemi-
galeidae (family Hemigalei or Hemigaleus of Hasse, 1879), that includes the
genera Hemipristis Agassiz, 1843; Hemigaleus Bleeker, 1852 (including Nego-
galeus Whitley, 1931); Dirrhizodon Klunzinger, 1871 (including Heterogaleus
Gohar and Mazhar, 1964); Chaenogaleus Gill, 1862b; and Paragaleus Budker,
1935. Dirrhizodon is usually synonymized with the fossil Hemipristis but is
separable by differences in tooth histology (Compagno, 1973b).

The Carcharhinidae and Hemigaleidae differ from the Triakidae in having
stronger monognathic heterodonty; precaudal pits present; pectoral fin skeleton
extending into the distal parts of the fins; distal pectoral radials with pointed
tips and tapering edges; longest distal radials over 3 times the length of the
longest proximal radials; lateral undulations present in the dorsal caudal margin
(irregular in Scoliodon and in some specimens of Triaenodon) ; nasal fontanelles
either separated from the nasal apertures by a cartilaginous bridge (hemigaleids,
Galeocerdo and Loxodon) or absent; no supraorbital crest; and levator palato-
quadrati muscles greatly expanded anteriorly, with their origins extending in
front of the postorbital processes and onto the sides and dorsal surface of the
cranial roof (Moss, 1972, and unpublished data). The Hemigaleidae addition-
ally differ from the Triakidae in having two pairs of foramina for the lateral aor-
tae and efferent hyoidian arteries on the basal plate. Carcharhinids differ from
triakids also in having weak or no basal ledges on their teeth; ectethmoid foram-
ina present on the ectethmoid condyles; and scroll intestinal valves.

COMPARISON WITH OTHER GENERA

Gogolia is separable from all other triakids by its unusually high ratio of
preoral length to mouth width (this is below 1.5 in other genera); very short
mouth (approached in this only by Furgaleus) ; very small gill openings; hetero-
donty pattern in adult (fig. 4); extreme anterior position of its first dorsal or-
igin (ago is similar in this but other genera have the first dorsal origin posterior
to the pectoral insertions) ; very long and low first dorsal fin (other genera have
the first dorsal base 7% of the dorsal caudal margin or less and the fin height
over half of the base length); and high total vertebral counts (Tviakis acuti-
pinna has 175-176 centra, but other triakids usually have fewer than 160).

Gogolia is distinguished from other triakids in a key to the genera of Tri-
akidae provided below. In this key the genera Galeorhinus, Hypogaleus, and
Hemitriakis are restricted to the species listed in Compagno (1970). Tvriakis
comprises the species placed in it by Compagno (1973a): T. scyllium Miler
and Henle, 1839, including Hemigaleus pingi Evermann and Shaw, 1927; T.
megaloptera (A. Smith, 1849), including Mustelus nigropunctatus J. L. B.

VoL. XXXIX] COMPAGNO: GOGOLIA FILEWOODI 393

Smith, 1952; 7. semifasciata Girard, 1854, including Mustelus felis Ayres, 1854;
T. maculata Kner and Steindachner, 1866, including Mustelus nigromaculatus
Evermann and Radcliffe, 1917; and 7. acutipinna Kato, 1968. The poorly
known genus Allomycter is tentatively placed in the Triakidae and included in
the key. Allomycter (with a single species, A. dissutus) was described from
photographs of a unique specimen of shark from Cuba, but this specimen was
lost before its describer could examine it (Guitart, 1972).

FAMILY TRIAKIDAE. KEY TO GENERA

la. Nostrils with broad nasoral grooves. Anterior nasal flaps very large, meeting each other

at midline of snout and overlapping mouth posteriorly. Scylliogaleus Boulenger, 1902.
1b. Nostrils without nasoral grooves. Anterior nasal flaps small to absent, when present sepa-
RALcOmnOmMEea cheOthersand snoOtereaciim ey TM OUbH ses ee De
2a. Preoral length 1.6-1.7 times mouth width. First dorsal fin with its base length almost
equal to length of dorsal caudal margin and 2.3—3.2 times the first dorsal height

ep eE ae momma CeR a SEs pe SE ies op ote Pl Gogolia Compagno, new genus.
2b. Preoral length less than 1.5 times mouth width. First dorsal with its base length 74 or
less of the dorsal caudal margin length and about 1.2-1.7 times the first dorsal height — 3.
3a. Anterior nasal flaps absent. Nasal cavities widely open to the exterior...
Enna rae ees SP a ee eS Allomycter Guitart, 1972.
3b. Anterior nasal flaps present. Nasal cavities not widely open to the exterior but communi-

cating with it through restricted incurrent and excurrent apertures 4.
4a. Teeth markedly unlike in upper and lower jaws. Anterior nasal flaps formed into slender
VDA ie SEUSS aa, cag a aE 2 CO ele Ae Furgaleus Whitley, 1951.
4b. Teeth not markedly unlike in upper and lower jaws. Anterior nasal flaps not barbel-
like _— Beane SL Sieg at Oe ae ae ee

5a. Ofgin ‘oft first omar far Forward, over Pheer. pies Se ee es ee
eS ES a Iago Compagno and Springer, 1971.
5b. Origin of first dorsal farther back, over inner margins of pectorals or posterior to
(HES, ne I I a Oe Sa ee meee es 6.
6a. Second dorsal fin about as large as anal fin, with its height 24 or less of first dorsal
height and its base about half as long as first dorsal base. Terminal sector of caudal
about half as long as dorsal caudal margin Galeorhinus Blainville, 1816.
6b. Second dorsal fin noticeably larger than anal fin, its height 42 to virtually equal to first
dorsal height and its base 73 to about equal to second dorsal base. Terminal sector of

cancel looks Ta As) loins As, clayseill Gauto bil Twoehyeabo ie
7a. Nostrils narrow and far apart, internarial space about 2.5-3.0 times nostril width. Teeth
formed into compressed cutting blades. Medials ee at symphysis of both jaws 8.

7b. Nostrils wider and closer together, internarial space about 1.0-2.0 times nostril width.
Teeth molariform to semibladelike, not greatly compressed. No differentiated medials
SEE, SYA OU OR PIS a Oa 2 ee ee Se ce Ag BA Oe 9.
8a. Eyes fusiform in shape and over 2 times as long as high. Mouth broadly arched. Second
dorsal over 7% of height of first dorsal. Ventral caudal lobe very short.
ee en A Ne ew a Re esse Hemitriakis Herre, 1923.
8b. Eyes pear-shaped and less than twice as long as high. Mouth triangular. Second dorsal
about half as high as first dorsal. Ventral caudal lobe very long.
we Ne ee Hy pogaleus \)\, iB. smith, 1957.
9a. Snout bluntly rounded in Bec eniel view, thick and blunt in lateral view. Mouth

394 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

arcuate in shape, lower jaw with convex edges. Teeth of lower jaw hardly overlapping
or not extending onto its ventral surface, not enlarged at symphysis or forming a knob
Be ee a ae ee Triakis Miller and Henle, 1838.
9b. Snout bluntly parabolic to subtriangular in dorsoventral view, bluntly to narrowly
pointed in lateral view. Mouth subtriangular or triangular in shape, lower jaw with
straight or nearly straight edges. Teeth of lower jaw prominently overlapping onto its
ventral surface, more or less enlarged at symphysis to form a knob resembling those of
FaySuok thessenus) cy 77CioO Oise Mustelus Linck, 1790.

Gogolia is closest to Galeorhinus, Hypogaleus, and Hemitriakis within the
Triakidae and is intermediate in some characters between Hemitriakis and the
other genera. Thus Gogolia resembles Hemitriakis in having strong subocular
ridges, external nictitating lower eyelids, shallow subocular pouches, and poorly
developed lateral cusps on the trunk denticles, but agrees with Galeorhinus and
Hypogaleus in eye shape, greatly reduced anterior nasal flaps and protruding
mesonarial flaps, and size disparity between first and second dorsal fins. The
premedial cusplets on the upper parasymphysial teeth of Gogolia recall similar
cusplets on upper and lower anteroposteriors of the Eocene fossil ““Galeus recti-
conus” (as described and illustrated by Leriche, 1905). However, the teeth of
“G. recticonus” have heavier cusps, larger and more regular premedial cusplets,
less compressed crowns and roots, and stronger transverse notches than in those
of Gogolia. Living Galeorhinus species also have a few irregular premedial cus-
plets on anteroposteriors closest to the symphysis, but these are less prominent
than the premedial cusplets of Gogolia and “G. recticonus.”

Gogolia filewoodi Compagno, new species.

Hototype. A pregnant adult female, 739 mm. total length, with 2 fetuses
(1 retained by Mr. L. W. Filewood), Australian Museum (Sydney) no. I.
16858—001. The remaining fetus, a 224 mm. male, is cataloged as CAS—27588.

TyprE LocaLity. Astrolabe Bay, Northern New Guinea, about 1 mile north
of the Gogol River mouth at 40 fathoms depth; about 5° 18’ S. Latitude and
145° 50’ E. Longitude.

SPECIES NAME. Named for Mr. L. W. Filewood in recognition of his work
on the poorly known elasmobranch fauna of New Guinea.

MEASUREMENTS AND PROPORTIONS. These are given as percentages of total
length for the adult female holotype and the male fetus (table 1).

DESCRIPTION (based on the adult female and fetal male). Head broad,
width at spiracles about half head length. Head moderately depressed and trape-

>

Ficure 4. Gogolia filewoodi. Set of detached teeth from left jaw half of 739 mm. holo-
type in labial view, with anteroposterior teeth numbered sequentially from medial row at
symphysis (M) to last row at end of dental band. A, upper teeth. B, lower teeth.

; og eam =r re mee ome ma)
| SS ae
: 8 Siow ; .
Sr SDEDERSDSDOLIERIOID

396 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

TasLe 1. Measurements of two specimens of Gogolia filewoodi.

Measurement Adult female % Male fetus %
Total length: 739 mm. — 224 mm. —
Tip of snout to:
Nostrils. 45 6.1 15 6.7
Upper symphysis. 72 9.8 25 tee?
Eyes. 57 Tell 20 8.9
Spiracles. 94 WAT 33 14.7
Ist gill openings. 133 18.0 46 20.6
5th gill openings. 158 25 53 DB
Pectoral origins. 152 20.1 51 22.8
1st dorsal origin. 173 23.4 62 Died
2nd dorsal origin. 434 Sail 138 61.6
Pelvic origins. 336 45.5 106 47.3
Anal origin. 445 60.2 140 62.5
Upper caudal origin. 560 75.8 172 76.8
Vent. 353 47.8 106 47.4
Distance between:
Vent and caudal tip. 386 52.2 118 52.6
1st and 2nd dorsal bases. 119 16.1 35 15.6
Pectoral and pelvic bases. 162 21.9 44 19.7
Pelvic and anal bases. 81 11.0 28 1225
2nd dorsal base and upper caudal origin. 58 7.9 14 6.3
Anal base and lower caudal origin. 58 7.9 14 6.3
2nd dorsal and anal origins. 17 2:3 3 3}
2nd dorsal and anal insertions. 11 125 2 0.9
Eyes:
Length. 26 3.5 13 5.8
Height. 13 1.8 6 27
Transverse distance between anterior corners. 52 7.0 20 8.9
Nostrils:
Width. 16 De 6 2.7
Internarial space. 33 45 12 5.6
Spiracles:
Diameter. 4 0.5 2 0.9
Mouth:
Length. 14 1.9 6 Dil
Width. 42 il 15 6.7
Labial furrows:
Length upper furrows. 17 2.3 6
Length lower furrows. 9 1.2 3 1.3
Gill opening widths:
Ist. wl 1S 3 3
2nd. 12 1.6 3 1.3
3rd. 12 1.6 DS) wei!
4th. 10 1.4 2 0.9
5th. 8 ail 115) 0.7

VoL. XXXIX] COMPAGNO: GOGOLIA FILEWOODI 397

TaABLe 1. Continued.

Measurement Adult female %o Male fetus %

Head:

Width at anterior corners of eyes. 66 8.9 25 11.2

Height at anterior corners of eyes. 35 4.7 11 4.9
Trunk:

Width at pectoral insertions. 68 9.2 18 8.0

Height at pectoral insertions. 78 10.6 20 8.9
Caudal peduncle:

Width at 2nd dorsal insertion. 24 Bee 6 DSi

Height at 2nd dorsal insertion. 26 3.5 7 Sail
Pectoral fins:

Length anterior margins. 118 16.0 31 13.8

Length posterior margins. 84 11.4 18 8.0

Length bases. 44 6.0 13 5.8

Distance from origin to free rear tip. 81 11.0 24 10.7

Length inner margins. 45 6.1 14 6.3
Claspers:

Length outer margins. — — 3 FS
Pelvic fins:

Length anterior margins. 46 6.2 13 5.8

Length bases. 43 5.8 10 4.5

Distance from origin to free rear tip. 59 8.0 17 6.7

Height. 33 4.5 7 3.1

Length inner margins. 30 4.1 8 3.6
1st dorsal fin:

Length anterior margin. 121 16.4 32 14.3

Length base. 157 Dis 49 21.9

Height. 67 9.1 15 6.7

Length inner margin. 32 4.3 8 3.6
2nd dorsal fin:

Length anterior margin. 76 10.3 22 9.8

Length base. 72 9.7 20 8.9

Height. 46 6.2 11 4.9

Length inner margin. 23 Sail 6 Dei
Anal fin:

Length anterior margin. 51 6.9 13 5.8

Length base. 50 6.8 10 4.5

Height. Dil Sail 6 Deh

Length inner margin. 19 2.6 6 2
Caudal fin:

Length dorsal margin. 167 22.6 51 20.9

Length preventral margin. 57 ed 18 8.0

Length lower postventral margin. 21 2.8 5 DD)

Length upper postventral margin. 77 10.4 21 9.4

Length subterminal margin. 32 4.3 11 4.9

Length terminal margin. 58 7.9 10 4.5

Length terminal sector. 64 8.7 16 He

398 CALIFORNIA ACADEMY OF SCIENCES [PRroc. 4TH SER.

zoidal in transverse section at eyes. Outline of head in lateral view slightly con-
vex ventrally and undulated dorsally, with a slight concavity in front of eyes
and a convexity above them (figs. 1, 2F). Head outline with prominent notches
just anterior to nostrils when viewed dorsoventrally (fig. 2D—E). Dorsal sur-
face of suborbital ridges with depressions over the nostrils and in front of eyes.
Eyes dorsolaterally situated on head, without eye notches. Ends of nictitating
lower eyelid attached to upper eyelid (fig. 2A). Secondary lower eyelid sharp-
edged and extending below the entire length of the eye. Kinetics and morphol-
ogy of the nictitating lower eyelids, secondary lower eyelids, subocular pouches,
and levator nictitans muscles (see below) indicate that Gogolia may not be able
to completely close its eyes with its nictitating lower eyelids. Spiracles large,
oval, and about % as long as eyes. Spiracles positioned posterior to eyes by
about their own lengths and about opposite the nictitating lower eyelids. No
papillose gill rakers on gill arches. Small lobes or mesonarial flaps (fig. 2B:
MNF) present on the bases of the anterior nasal flaps above their tips and pro-
truding from the nasal apertures (as in Galeorhinus). Low posterior nasal flaps
present (fig. 2B: PNF). Mouth width about % of width of head at mouth
corners. No large buccal papillae in mouth. Upper labial furrows about twice
as long as lower ones, with their anterior ends slightly in front of eye pupils.

Teeth showing strong gradient monognathic heterodonty in anteroposterior
teeth of both jaws of adult. Teeth become proportionately longer, lower, smaller,
and more oblique-cusped towards the ends of the dental bands (fig. 4). Post-
lateral cusplets are reduced and replaced by postlateral blades in this direction
in both jaws, while premedial cusplets are replaced by undifferentiated premedial
edges in the upper jaw. Ontogenic heterodonty apparently similar to that of
Hemitriakis (Compagno, 1970), with the fetus lacking cusplets and having post-
lateral blades only on all of its anteroposterior teeth. Teeth not forming a pave-
ment and hardly protruding when mouth is closed.

Body moderately slender, with trunk high and subtriangular in section at
first dorsal base in adult but lower in fetus. Caudal peduncle short and nearly
oval in section, not greatly compressed and without lateral dermal keels. No
predorsal or postdorsal ridges present. Lateral trunk denticles with strong me-
dial cusps and a pair of medial ridges in fetus and adult, but with lateral ridges
also present in adult (fig. 3A-B). Denticle crowns have reticulated depressions
on their dorsal surfaces, much as in Jago (Compagno and Springer, 1971, fig. 6)
and many other carcharhinoids.

Pectoral fins broad and subangular, with slightly convex anterior margins,
narrowly rounded apices, slightly concave posterior margins, and broadly
rounded free rear tips and inner margins. Pectorals slightly smaller than first
dorsal. Origins of pectorals about under 3rd gill openings. Apex of pectoral
posterior to its free rear tip when fin is elevated and appressed to body.

VoL. XXXIX] COMPAGNO: GOGOLIA FILEWOODI 399

5mm.

el

Ficure 5. Gogolia filewoodi. A, pectoral fin skeleton of 739 mm. holotype, traced from
radiograph (details of the axial segments of the metapterygium could not be distinguished),
with outline of pectoral fin indicated in black. B, dissection of pectoral fin skeleton from
224 mm. fetus. C, hyobranchial skeleton (excluding hyobranchial rays and extravisceral
cartilages) from 224 mm. fetus. D-E, transverse sections of MP vertebral centra from below
first dorsal fin base, with sections cut through the apices of the calcified double cones of each
centrum; calcified structures are in heavy black and cartilage stippled. D, centrum from 224
mm. fetus. E, centrum from 739 mm. holotype. Abbreviations: BBC, basibranchial copula ;
BH, basihyoid; CB 1-5, ist to 5th ceratobranchials; CDC, calcified double cone; CH, cera-
tohyal; DCL, diagonal calcified lamella; DRA, distal radial segments; EB 1-4, Ist to 4th
epibranchials; HB 2-4, 2nd to 4th hypobranchials (1st absent); HMD, hyomandibula; IN,
intermedialia; IRA, intermediate radial segments; MS, mesopterygium; MT, basal segment
of metapterygium (lightly calcified in fetus) ; MTH, basal segment of metapterygium (heav-
ily hypercalcified and irregular in shape in adult); MTS, distal segments of metapterygium
(metapterygial axis); NAR, neural arch; NCA, neural canal; NO, notochordal canal; P,
propterygium; PB 1-4 +, 1st to 4th pharyngobranchials (4th one probably a compound
element) ; PRA, proximal radial segments.

400 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

KS

ONF
ECN

Ficure 6. Gogolia filewoodi. Neurocranium dissected from 224 mm. fetus, in dorsal
(A), ventral (B), and lateral (C) views. Abbreviations: AF, anterior fontanelle; AFV,
position of anterior facial vein in nasal fontanelle; ASC, anterior semicircular canal; BP,
basal plate; ECN, ectethmoid condyle; F IJ, optic nerve foramen; F III, oculomotor nerve
foramen; F IV, trochlear nerve foramen; F IX, glossopharyngeal nerve foramen; F X, vagus
nerve foramen; FC, foramen for internal carotid artery; FCV, foramen for anterior cerebral
vein; FEN, foramen for endolymphatic duct; FES, foramen for efferent spiracular artery;
FM, foramen magnum; FOC, orbitocerebral foramen for superficial ophthalmic nerve; FOE
and FOI, preorbital and orbital foramina for supraorbital canal through supraorbital crest,

VoL. XXXIX] COMPAGNO: GOGOLIA FILEWOODI 401

Pectoral fin skeleton with 1 radial on propterygium, 4-5 on mesopterygium,
and about 15-16 on metapterygium (including segmented axis of metapteryg-
ium). Radials mainly divided into 3 segments: proximal, intermediate, and
distal. Metapterygium (exclusive of segmented axis) densely hypercalcified
and irregularly expanded in adult but not in fetus (fig. 5A—-B).

Pelvic fins triangular, short, and about as large as anal fin. Pelvic anterior
margins slightly convex, apices subacute, posterior margins straight, free rear
tips pointed, and inner margins nearly straight. Inner margins, free rear tips,
and posterior margins of pelvics forming a broad triangle.

First dorsal broadly triangular, with convex to undulated anterior margin,
narrowly rounded apex, concave posterior margin, acute and attenuate free rear
tip, and concave inner margin. Midpoint of first dorsal base varying from about
equidistant between pectoral and pelvic bases (fetus) to about 1.6 times farther
from pelvic bases than pectorals (adult). First dorsal insertion slightly anterior
to pelvic origins by a distance somewhat less than the lengths of the pelvic bases.
First dorsal insertion far posterior to apex of first dorsal.

Second dorsal much narrower-based and more acutely triangular than first
dorsal, with undulating anterior margin, broadly rounded apex, moderately con-
cave posterior margin, acutely pointed and attenuate free rear tip, and concave
inner margin. Second dorsal origin anterior to anal origin by a distance less than
% of second dorsal base length.

Anal fin a low triangle, with undulated to slightly convex anterior margin,
round apex, shallowly concave posterior margin, acute free rear tip, and nearly
straight posterior margin.

Dorsal caudal margin convex to undulated, without a crest of enlarged den-
ticles. Preventral margin slightly convex, tip of ventral lobe narrowly rounded,
upper and lower postventral margins and subterminal margin nearly straight,
and terminal margin slightly concave. Subterminal margin about 0.6—1.1 times

=

containing superficial ophthalmic nerve; FPC, orbitocerebral foramen for deep ophthalmic
nerve; FPE and FPI, preorbital and orbital foramina for canal transmitting deep ophthalmic
nerve through supraorbital crest; FPN, fenestra for the perilymphatic canal; FS, foramen for
the stapedial or orbital artery; HF, hyomandibular facet; IOC, interorbital canal; LR,
lateral rostral cartilage; MR, medial rostral cartilage; NA, nasal aperture; NF, nasal fon-
tanelle; NP, orbital notch; O, orbit; OC, occipital condyle; OCN, occipital centrum; ONF,
orbital foramen for orbitonasal canal (nasal foramen or ectethmoid chamber for the canal
is inside nasal capsule) ; OR, opisthotic ridge; ORF, orbital fissure; OT, otic capsule; PR,
preorbital process; PRF, parietal fossa; PSC, posterior semicircular canal; PT, postorbital
process; RF, rostral fenestra; RN, rostral node; RRF, ridge between base of lateral rostral
cartilage and edge of anterior fontanelle; SC, supraorbital crest; SEF, subethmoid fossa; SR,
sphenopterotic ridge; SS, suborbital shelf.

[Proc. 4TH SER.

CALIFORNIA ACADEMY OF SCIENCES

402

VoL. XXXIX] COMPAGNO: GOGOLIA FILEWOODI 403

terminal margin, relatively shorter in adult than fetus. Vertebral axis of caudal
noticeably raised.

Total vertebral counts 180 (2). Counts of monospondylous precaudal (MP)
centra 27.2 percent, counts of diplospondylous precaudal (DP) centra 35.0-36.7
percent, and counts of diplospondylous caudal (DC) centra 35.0-36.7 percent
of total counts. Ratios of DP/MP counts 1.3-1.4, DC/MP counts 1.3—-1.4, A
ratios (length of penultimate MP centrum/length of first DP centrum x 100)
150-160, and B ratios (length/width of penultimate MP centrum x 100) 67—
68 (methods of counting and ratios follow Springer and Garrick, 1964, and
Compagno, 1970). Transition between MP and DP centra over region of pelvic
fin bases. Last few MP centra before MP—DP transition not greatly enlarged.

Vertebral calcification patterns studied from transverse sections of MP cen-
tra below first dorsal base (terminology from Ridewood, 1921). Diagonal cal-
cified lamellae very long in adult but short in fetus. Dorsal, ventral, and lateral
intermedialia are present in the adult but not in the fetus (fig. 5D-E). Noto-
chordal canal very small at apices of calcified double cones in adult (unlike
adults of Jago) but large in fetus.

Neurocranium was dissected in the fetus (fig. 6) and studied by stereo-
radiographs in the adult. Cranial terminology is modified from Gegenbaur
(1872), Allis (1923), Holmgren (1941), Gilbert (1967), and Compagno (1973a
and b). The cranium of G. filewoodi is most similar to those of Hemitriakis,
Galeorhinus, and Hypogaleus, and less like those of other triakids (the neuro-
cranium was dissected and examined by the writer from all triakid genera except
Allomycter). Rostral cartilages slender and not hypercalcified in adult, fused
at their tips to form a fenestrate rostral node. Nasal capsules slightly wider than
long, greatest transverse width across them 1.1—1.3 in nasobasal length. Anterior
fontanelle horizontally oval, with its width about 3.4-3.7 in nasobasal length.

Ficure 7. Gogolia filewoodi. Head of 224 mm. fetus in dorsal (A), ventral (B), and
lateral (C) views, dissected to show relation of neurocranium to jaws, jaw and hyoid muscles,
and cranial nerve roots. Head outline in black. Abbreviations: AF, anterior fontanelle;
EM, epibranchial myomeres; HMD, hyomandibula (distal end); LC, labial cartilages; LCP,
cushioning ligament between ectethmoid condyle and orbital process of palatoquadrate; LR,
lateral rostral cartilage; MAM, adductor mandibulae muscle; MC, Meckel’s cartilage; MCM,
coracomandibular muscle; MIH, interhyoid muscle; MIM, intermandibular muscle; MLH,
levator hyomandibuli muscle; MLP, levator palatoquadrati muscle; MLS, levator labii supe-
rioris or preorbitalis muscle; MR, medial rostral cartilage; N II, optic nerve; N III, oculo-
motor nerve; NA, nasal aperture; NB VII, buccal ramus of facial nerve; NC, nasal capsule;
NH VII, hyomandibular ramus of facial nerve; NM V, mandibular ramus of trigeminal
nerve; NMX V, maxillary ramus of trigeminal nerve; NP V, deep ophthalmic ramus of tri-
geminal nerve; NS V + VII, supraorbital trunk or superficial ophthalmic ramus of trigemi-
nal and facial nerves; OPQ, orbital process of palatoquadrate; PQ, palatoquadrate; PT,
postorbital process; SC, supraorbital crest; SP, spiracle; SS, suborbital shelf.

404 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Dorsal lip of fontanelle not anteriorly flared and without an epiphysial foramen.
Cranial roof broadly arched between orbits. Orbital notches very prominent (fig.
6B: NP). No keels on basal plate. Internal carotid foramina closer to each
other than to the stapedial foramina. Edge of supraorbital crests arcuate in
dorsal view. Orbit oval in lateral view, with contents indicated in fig. 6C. No
ledges between nasal capsules and suborbital shelves. Otic capsules not greatly
expanded or inflated, their lengths about 3.3 in nasobasal length and the great-
est transverse width across them about 1.9 in nasobasal length. Hyomandibular
facets small and horizontally elongate. Occipital condyles short, with a single
occipital centrum between them (fig. 6B—C: OC and OCN).

The hyobranchial skeleton was studied in the fetus by dissecting it out,
staining it with methylene blue by Van Wijhe’s method (Gray, 1954), and
clearing it with the tricresyl phosphate-tributyl phosphate clearing solution of
Groat (1941). It resembles comparable structures in other carcharhinoids ex-
amined (Leptocharias and Scoliodon) but has an unusually large and broad
basihyoid and basibranchial copula (fig. 5C: BH and BBC). The hyomandibu-
lae are short and moderately stout, the ceratohyals long and stout, and 3 pairs
of hypobranchials (apparently nos. 2—4) are present.

Jaws and musculature associated with them are relatively weak (terminology
for muscles follows Lightoller, 1939). Palatoquadrates and Meckel’s cartilages
flat and weak, with low orbital processes on palatoquadrates. The orbital pro-
cesses articulate with the ectethmoid condyles on the nasal capsules and are
attached to them by cushioning ligaments (fig. 7B—C: LCP and OPQ). Leva-
tor labii superioris or preorbitalis muscles small and single-headed, their origins
on the sides of the nasal capsules lateral to the ectethmoid condyles and their
insertions on the adductor mandibulae muscles near the mouth angles. Levator
palatoquadrati muscles very slender, originating at the bases of the postorbital
processes and inserting on the posterodorsolateral ends of the palatoquadrates.
Levator hyomandibuli muscles very slender, originating on the sides of the otic
capsules below the sphenopterotic ridges and inserting on the dorsolateral ends
of the hyomandibulae. Levator nictitans short and poorly differentiated (not
shown in fig. 7). Coracomandibular (fig. 7B: MCM), coracohyoid, and cora-
cobranchial muscles very large, but branchial constrictors could not be found in
the specimen examined.

Stomach divided into a moderately large fundus and a long, slender pylorus.
The fundus extends posteriorly about half the length of the pleuroperitoneal
cavity then reverses direction as the pylorus nearly to the base of the liver, where
it connects with the valvular intestine. The intestine (fig. 2C) has 7 turns in
its spiral valve. The rectum has a bulbous rectal gland attached to it by a stalk.
Liver moderately large, irregularly bilobate, and extending posteriorly over
*, of the length of the pleuroperitoneal cavity. Liver not obscuring other vis-

VoL. XXXIX] COMPAGNO: GOGOLIA FILEWOODI 405

cera entirely in ventral view. Spleen elongate and not nodular, with an irregu-
larly thickened part opposite the end of the fundus and a slender anterior part
extending along the pylorus. Pancreas small, located at junction of spiral in-
testine and pylorus. Ovary present on right side only, with follicles up to 6 mm.
in diameter present. Both oviducts are developed, each having a moderately
large nidimental gland.

Development ovoviviparous. The 2 fetuses (probably full-term or nearly so)
were carried one to an oviduct and lacked placental connections to the mother.

Color of adult dark gray tinged with brown on dorsum, fins, and underside
of head, light gray or gray-cream on abdomen and flanks. Dorsal fin edges dusky
or blackish and posterior margin of pectorals light. Fetus is gray-brown above,
lighter below, and dusky finned.

SUMMARY

Gogolia filewoodi, new genus and species, is described from two specimens
of triakid shark from northern New Guinea. The genus differs from other tri-
akids in having an unusually high ratio of preoral length to mouth width, very
short mouth, small gill openings, extremely long first dorsal base, and high ver-
tebral counts. The family Triakidae is resurrected, redefined, and limited to
the genera Mustelus, Galeorhinus, Triakis, Scylliogaleus, Hemitriakis, Fur-
galeus, Hypogaleus, Iago, Allomycter, and Gogolia. The family Carcharhinidae
is confined to the genera Carcharhinus, Scoliodon, Galeocerdo, Triaenodon,
Loxodon, Hypoprion, Prionace, Aprionodon, Isogomphodon, Lamiopsis, Rhizo-
prionodon, and Negaprion. Hemipristis, Hemigaleus, Dirrhizodon, Chaeno-
galeus, and Paragaleus are referred to the family Hemigaleidae, Leptocharias
tentatively to its own family, Leptochariidae, and the ‘scyliorhiniform triakoid’
genera Proscyllium, Eridacnis, Ctenacis, and Gollum to the family Proscylliidae.

ACKNOWLEDGMENTS

I would especially like to thank L. W. Filewood (Department of Agriculture,
Stock and Fisheries, Konedobu, Papua-New Guinea) for making it possible for
me to describe his unique new shark. The following individuals were very help-
ful in providing specimens, information, and facilities used in this study: Phillip
C. Heemstra (Academy of Natural Sciences of Philadelphia); John R. Paxton
(Australian Museum, Sydney); W. I. Follett, William N. Eschmeyer, George
S. Myers, Warren C. Freihofer, and Lillian J. Dempster (Department of Ichthy-
ology, California Academy of Sciences, San Francisco); C. G. Alexander (Cali-
fornia State University at San Francisco); Shelton P. Applegate (Division of
Vertebrate Palaeontology, Los Angeles County Museum of Natural History) ;
J. M. Moreland (Dominion Museum, Wellington, New Zealand); Stewart
Springer (Mote Marine Laboratory, Sarasota, Florida); Tyson R. Roberts and
Myvanwy M. Dick (Museum of Comparative Zoology, Harvard University) ;

406 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Susumu Kato (National Marine Fisheries Service, Tiburon Fisheries Laboratory,
Tiburon, California); John Bass and John Wallace (Oceanographic Research
Institute, Durban, South Africa); the late J. L. B. Smith (Rhodes University,
Grahamstown, South Africa); Carl L. Hubbs, Richard H. Rosenblatt, and
Leighton R. Taylor, Jr. (Scripps Institution of Oceanography, La Jolla, Cali-
fornia); Leslie W. Knapp (Smithsonian Oceanographic Sorting Center, Wash-
ington, D. C.); Robert P. Dempster (Steinhart Aquarium, California Academy
of Sciences); Victor G. Springer and Stanley H. Weitzman (Division of Fishes,
U. S. National Museum, Washington, D. C.); Reeve M. Bailey (University
of Michigan Museum of Zoology, Ann Arbor); J. A. F. Garrick (Victoria Uni-
versity, Wellington, New Zealand); and R. J. McKay (Western Australian
Museum, Perth, Australia). Through the invaluable aid of these people I was
able to examine and dissect representatives of all genera of the families Proscyl-
liidae, Leptochariidae, Triakidae (except Allomycter), Hemigaleidae, and Car-
charhinidae and compare them with one another and with Gogolia.

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PROCEEDINGS

OF THE

CALIFORNIA ACADEMY OF SCIENCES
3 FOURTH SERIES

_ Vol. XX XIX, No. 20, pp. 411-500; 72 figs. December 19, 1973

MUSCULAR ANATOMY OF THE FORE-
-LIMB OF THE SEA OTTER

EY LE TE ALS. BS

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PROCEEDINGS
OF THE ee

CALIFORNIA ACADEMY OF SCIE

FOURTH SERIES

Vol. XXXIX, No. 20, pp. 411-500; 72 figs. December 19, 1973

MUSCULAR ANATOMY OF THE FORE-
LIMB OF THE SEA OTTER
(ENHYDRA LUTRIS)

By
L. D. Howard
Hopkins Marine Station of Stanford University, Pacific Grove, California 93950

INTRODUCTION

The present study was undertaken to make available for future reference
the detailed muscular anatomy of the forelimb of this remarkable marine mam-
mal, Enhydra lutris.

The sea otter has a limited southern range of approximately 140 miles along
the coast of California from the Monterey Peninsula on the north to Morro Bay
on the south. It is estimated by aerial and shoreline survey, that 1,000 individ-
uals make up the population in this area.

The sea otter rarely comes on shore in this area, preferring the sanctuary of
the abundant kelp beds for its home. Food preference is exclusively the many
invertebrates found in this area, such as the various kinds of crabs, shellfish,
and sea urchins. Since this food is mainly on the ocean floor, the animals dive
to obtain it but return to the surface to consume it while floating on their backs
using the chest as a dinner table. Divers, observing otters gathering their food,
note that it is carried either in a bimanual manner or under one arm in the axil-
lary area where a fold of skin assists in securing the object.

On occasions, an animal will bring up a rock and, placing it on its abdomen,
use it as an anvil against which it can break shellfish. In so doing the animal
holds the food between its forepaws and rapidly strikes it against the rock. In
view of this habit, the otter has been classified as a tool user.

We are concerned, therefore, with the work this animal has to do with the

[411]

412 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

forelimbs in an aquatic environment, and the structural anatomy he has with
which to accomplish these tasks.

Over the past several years, the author has observed these animals in their
natural habitat, noting the high degree of specialized use of the forelimbs in
eating, grooming, and caring for the young. Based on these observations it would
appear that the ‘hand’ of this animal should be extremely dextrous, yet when
observed at close range, the anatomical characteristics show marked resemblance
to the forepaw of mammals which use the forelimb mainly for land propulsion.

Many references in the literature describe how the otter uses its forelimbs.
It seemed of interest, therefore, to study the ‘hand’ in some detail from an ana-
tomical standpoint. To do so necessitated the study of the entire forelimb since
the hand, to be effective, must be positioned by the forelimb and positional stabil-
ity maintained when the hand itself is in use.

The material for this study consisted of one fresh specimen on which the
range of motion measurements were made, and one preserved specimen used for
recording the detail of the muscular anatomy. The method used for this latter
study was to make a series of anatomical dissections, photograph each, and then
from the projected slides make accurate line drawings. In this way, the relative
size and relationship of each muscle could be portrayed.

The forelimb muscles can be grouped in various ways: by their innervation,
by their action (such as extensors and flexors), by their origin (pre- and post-
axial), or by where they make their attachments. Examples of the last would
be muscles connecting the trunk to the limb, muscles connecting the scapula to
the humerus, etc.

In this treatise a fixed method of presentation is not used, but rather the
muscles are described as they are encountered in a dissection going from the
proximal to the distal areas. Each muscle is numbered and retains this number
wherever it appears. A gross description of each muscle is given with its origin
and insertion and a notation as to its action or actions if the muscle were to be
activated as a single unit. This situation, of course, never happens in the living
specimen, as each voluntary motion of any part of the limb is a complicated
interplay between the many muscles, some relaxing, some serving to stabilize
the joints, and some serving as prime movers for any given motion.

For further clarification, plates of the forearm skeleton have been added
and the areas of origin and insertion of muscles designated.

OBSERVATIONS OF JOINT MOBILITY

In a fresh specimen, by means of passive motions, some idea of the range
of motion of the forelimb could be observed and for some joints the range of
motion could be actually recorded.

VoL. XXXIX] HOWARD: SEA OTTER FORELIMB

ro
me» .

e . e*
a

Ficure 1. An anterior or palmar view of the forefoot to show the disposition of the
palmar pads. The major crease lines are transverse and quite deep. Distally, shallow incom-
plete crease lines suggest interdigitation, but such is not the case to any practical degree.

The pads appear to bulge slightly and are turgid, and the covering skin is thick with a peb-
bly roughness.

414

CALIFORNIA ACADEMY OF SCIENCES

[Proc. 4TH Serr.

VoL. XXXTX] HOWARD: SEA OTTER FORELIMB 415

Ficure 3. In this posterior or dorsal view of the forefoot the hair has been trimmed
away to show the position of the claws which, in the unflexed hand, are visible only in this
view. Since the claws are closely associated with, and move only with, the terminal phalanx,
their position indicates a marked degree of hyperextension of the terminal digital joints.

As there is no clavicle in this animal, the pectoral girdle, which consists of
the scapula only, is extremely mobile on the chest wall, being limited only by
the muscles attaching the girdle to the trunk. Thus, in a transverse plane, the
pectoral girdle could be moved to near the midline dorsally and ventrally, and
in the longitudinal plane an equal, if not slightly greater, range of motion was
present. A good range of motion in the shoulder joint was also present but tech-

416 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

nically it was not possible to get accurate measurements in degrees of abduction,
adduction, flexion, extension, and rotation.

For the elbow, extension was to an angle of 145° and flexion to an angle
of 40°.

For the forearm, pronation was to 30° beyond the midposition and supina-
tion to 45° beyond the midposition.

The wrist is very mobile. Palmar flexion of the wrist was to an angle of 82°
beyond the straight line of the forearm, and dorsiflexion to an angle of 40°.
Good lateral motion of the wrist was also present, radioflexion being to an angle
of 30° and ulnaflexion to an angle of 50°.

For digits 2, 3, 4, and 5, the average range of motion of the individual joints
as measured from the line of the metacarpal was as follows:

Extension Flexion
Metacarpal phalangeal joint 80° 45°
Proximal interphalangeal joint 525 45°
Distal interphalangeal joint Ih7i0)= 205
For digit 1:
Metacarpal phalangeal joint 80° 65°
Interphalangeal joint 170" 50°

From the above, it is to be noted that the digital joints have a greater range of
extension than of flexion, which, in itself, would indicate the sacrificing of a
gripping mechanism in favor of ambulatory functions.

This situation was even more evident when traction was made on the digi-
tal extensors and the flexor tendons in the forearm.

The digits of the ‘hand’ are not individualized, being held closely together
by webbing and the first digit, or thumb, is in the same plane as the other digits.
Without an established thumb web the first digit is unopposable and lateral
motions of the other digits are of insignificant amount because of the webbing.
In fact, a better term than “webbing” would be total syndactylism.

An effort was made to see if objects could be held in the hand by a grasp
mechanism. When the long flexors were pulled upon strongly in the forearm
an ordinary pencil, for example, could not be grasped firmly in the hand. Larger
objects could be held but rather precariously in the absence of an opposable
thumb. Part of the problem, as far as gripping objects is concerned, appears to
be due to the large pads on the palmar surface which, on grasp, appear to fill
the palm. When the digits were flexed to their limits, the terminal digital pads
failed to touch the palmar pad by about 12 mm. However, there is sufficient
flexion of the digits to allow the claws to be used against the flat opposing sur-
face. Since palmar flexion of the wrist is extremely good, large objects of 8 or
more cm. in diameter could be held between the hand and distal forearm.

VoL. XXXIX] HOWARD: SEA OTTER FORELIMB 417

«

ae
SHAS
za

Ficure 4. This figure is a direct line copy of an x-ray of the hand and wrist after soft
iron wires were placed in the creases of the palmar pad. This was done to show the relation-
ship of these crease lines to the underlying joints. The pad itself lies distal to the most proxi-
mal wire (heavy line). From this it is to be noted that the volar pad covers only the distal
one-half of the metacarpals plus the phalanges. The main transverse creases are at metacarpal
phalangeal joint level. Therefore when walking on this pad the otter does not walk on the
flat of the hand as it appears, but rather on the metatarsal heads and the phalanges. This
is readily understandable in view of the limited dorsiflexion of the wrist and the great degree
of hyperextension of the metacarpal phalangeal joints.

418 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Ficure 5. The grasp mechanism of the individual hand is markedly limited as shown in
this figure. The long flexor tendons of the digits were pulled upon forcibly in an effort to
demonstrate grasp of a pencil. This precarious grasp depended upon sufficient flexion of the
finger joints to allow the claw of the fifth digit to become effective. Without a good claw
(and the claws are frequently worn down to mere nubbins) it is doubtful that either of the
objects shown in figures 5 and 6 could be held in the one hand. See also figure 6.

On the few occasions that the animal was observed on land, the forelimb
functioned admirably for terrestrial progression. Underwater observers report
that the forelimbs are not used for swimming, but are active in food gathering.

In light of these observations it is understandable why the animal performs

VoL. XXXIX] HOWARD: SEA OTTER FORELIMB 419

Ficure 6. In this figure, a medium-sized crab leg has been substituted for the pencil
of figure 5.

most of the forelimb functions in a bimanual manner. Anatomically, the hand is
small, the digits are not individualized, and grasp function is poor. Therefore,
to obtain the observed dexterity the forelimbs must be used together. One hand
opposing the other provides what we would interpret as a ‘thumb service.’

One is reminded of a bilateral human amputee who has lost both hands at
wrist level, and in the absence of a prosthesis everything that can be done is
done by opposing the two forearm stumps.

Thus it is interesting to speculate whether or not an otter could survive if
one hand was lost. The probability seems good that he could, using the forearm
stump against the opposite hand. However, to lose one extremity at elbow
level would probably prove disastrous.

The anatomical study which follows was made on sea otter ##222 (State of

420 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

California Fish and Game). The otter was an adult male with an overall body
length (tip of nose to tip of tail) of 125 cm. The length of the upper arm was
approximately 14 cm. and the lower arm was of equal length. The length of the
forefoot (hand) was 7 cm. and the width was 5 cm. It must be kept in mind
that only a single specimen was used and therefore variations and anomalies
which are bound to occur between individual specimens are not recorded.

MUSCLES OF THE FORELIMB OF THE SEA OTTER

ALPHABETICAL LISTING OF MUSCLES

Muscle No. Muscle No.
Abductor digiti quinti 49 Flexor digitorum sublimis ulnaris 48
Abductor digiti secundi 57 Flexor pollicis brevis 55
Abductor pollicis brevis 54 Infraspinatus 24
Abductor pollicis longus 38 Interossei 59
Acromiodeltoideus 17 _—Latissimus dorsi 15
Acromiotrapezius 2 Levator anguli scapulae 11
Adductor digiti quinti 52 Lumbricali 53
Adductor digiti secundi 57. Omotrachleon 6
Adductor pollicis 56 Opponens digiti quinti 50
Anconeus externus 23 + Palmaris longus 40
Biceps brachii 28 Pectoantebrachialis 2
Brachialis 29 + Pectoralis major 13
Brachioradialis 30 ~=©Pectoralis minor 14
Clavobrachialis 5 Pronator quadratus 46
Clavotrapezius 3 Pronator teres 44
Cleidomastoideus 4 Rhomboideus capitis 8
Dorsoepitrochlearis 19 Rhomboideus major 7
Extensor carpi radialis brevis 32 Rhomboideus profundus 9
Extensor carpi radialis longus 31 Serratus magnus 10
Extensor carpi ulnaris 35 Spinodeltoideus 18
Extensor digitorum communis 33 Spinotrapezius 1
Extensor digitorum lateralis 34 Subscapularis 27
Extensor pollicis et indicis longus 36 Supinator Si]
Flexor carpi radialis 43 Supraspinatus 26
Flexor carpi ulnaris 39 Teres major 16
Flexor digitorum profundus, central head 45 Teres minor 25
Flexor digitorum profundus, humeral head 42 Triceps brachii caput lateralis 21
Flexor digitorum profundus, ulna head 41 Triceps brachii caput longus 20
Flexor digiti quinti brevis 51 ‘Triceps brachii caput medialis 22
Flexor digitorum sublimis radialis 47

DESCRIPTIONS OF MUSCLES

1. Spinotrapezius.

This muscle is the most posterior of the three muscles which make up the
trapezius muscle complex. The spinotrapezius, with the acromiotrapezius and

VoL. XXXIX] HOWARD: SEA OTTER FORELIMB 421

Ficure 7. Lateral view of the right side of the trunk to show the superficial trapezius
muscle complex, which consists of three muscles, the spinotrapezius (1), the acromiotrapezius
(2), and the clavotrapezius (3). These muscles are vago-accessory, presumably having mi-
grated posteriorly from the pharyngeal area. Together they form a broad dorsal sheet of
muscle suspending the pectoral girdle and the forelimb from the middorsal line of the body.
The scapula is outlined to show its relationship to the musculature. Key: 1, spinotrapezius;
2, acromiotrapezius; 3, clavotrapezius; 6, omotrachlean (levator scapulae ventralis); 15,
latissimus dorsi; 18, spinodeltoideus; 19, dorsoepitrachlearis; 20, triceps brachii caput longus;
24, infraspinatus; A, spinous process of first dorsal vertebra; B, spinous process of eighth
dorsal vertebra; C, middorsal line-cervical area; D, site of tuberosity on spine of scapula;
E, lambdoid ridge of skull; S, outline of scapula.

the clavotrapezius, forms a broad superficial muscle sheath joining the axial
skeleton to the scapula and forelimb. The muscle is thin and triangular in shape
with the base along the middorsal line.

Orictn. Arising by fleshy fibers along the middorsal line and spinous pro-
cesses from T-2 to T-8, the muscle triangulates toward the tuberosity of the
scapular spine, where a short, flat tendon develops.

422 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Ficure 8. Lateral view of the right scapular area. The acromiotrapezius muscle (2) has
been detached from its origin and reflected downward and posteriorly to show the detail of
its insertion on the scapular spine and acromium process. Note the unattached raphe area
(B). The deeper rhomboid muscles, (7), (8), (9), lying beneath the acromiotrapezius muscle
are now exposed. Key: 1, spinotrapezius; 2, acromiotrapezius; 3, clavotrapezius; 6, omo-
trachlean (levator scapulae ventralis); 7, rhomboideus major; 8, rhomboideus capitis; 9,
rhomboideus profundus; 15, latissimus dorsi; 19, dorsoepitrachlearis; 24, infraspinatus; 26,
supraspinatus; A, scapula, spine; B, raphe—unattached to spine of scapula; C, scapula, acro-
mion process; D, middorsal line, cervical area; S, outline of scapula.

INSERTION. The tendon inserts on the tuberosity of the spine of the scapula
along the anterior and distal surface.

Action. The principal action is to support the scapula and forelimb and
move these structures toward the middorsal line.

2. Acromiotrapezius.

This is the central muscle of the trapezius muscle complex being situated
between the spinotrapezius posteriorly and the clavotrapezius anteriorly. The
muscle is thin and broad, and rhomboid in shape.

Ortcin. Arising by fleshy fibers from a broad base along the middorsal

VoL. XXXIX] HOWARD: SEA OTTER FORELIMB 423

line extending from the midcervical area anteriorly to the spinous process of
T-2 posteriorly. The muscle passes sheetlike toward the spine of the scapula.
Here, the muscle fibers end in a crescent-shaped tendinous raphe in the central
area and in short tendon fibers at the proximal end and fleshy fibers at the dis-
tal end.

INSERTION. The proximal tendon fibers insert on the tuberosity of the
spine of the scapula just distal to the tendon of the spinotrapezius. The distal
fleshy fibers insert on the full length of the acromion process of the scapula.
The tendinous raphe spans between the two insertions and is unattached to the
spine of the scapula, although paralleling it. The distal insertion is crossed
superficially by the omotrachleon muscle. The tendinous raphe gives origin
to the spinodeltoideus muscle.

Action. This muscle assists in the support of the scapula and forelimb
from the axial skeleton and can assist in rotating the scapula counterclockwise.

3. Clavotrapezius.

The anterior muscle of the trapezius muscle complex is also thin and broad
and closely associated along its posterior border with the acromiotrapezius.
The muscle is somewhat triangular in shape with a broad base dorsally.

OrIcGIN. The muscle arises by fleshy fibers from the middorsal line adjacent
to the origin of the spinotrapezius in the cervical area. The origin extends an-
teriorly the length of the remaining cervical area to reach the skull at the exter-
nal occipital crest. The origin then continues laterally along the lambdoid ridge
of the skull approximately one-third the distance to the jugular process. From
their origin the muscle fibers pass ventrally toward the shoulder, at which point
the muscle divides into an anterior one-third and posterior two-thirds.

INSERTION. The anterior one-third of the muscle continues toward the cla-
vicular area inserting into the deep fascia at this site since there is no clavicle.
The posterior two-thirds of the muscle divides about equally into two parts.
The anterior portion joins with a remaining portion of the cleidomastoid muscle
at this level to continue distally and eventually reaches the ulna. The posterior
portion joins with the muscle fibers from the cleidomastoid muscle to form the
clavobrachialis muscle, which in turn inserts more distally along the pectoral
ridge of the humerus.

Action. With this somewhat complicated insertion the muscle acts, in ad-
dition to supporting the pectoral girdle and drawing it upwards and forward,
to assist in flexion of both the shoulder and elbow.

4. Cleidomastoideus.

This long, thick, straplike muscle joins the skull to the upper extremity.
Oricin. Arising by fleshy fibers from the anterior and inferior surface of

424 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Ficure 9. Anterolateral view of the right shoulder. The clavotrapezius—cleidomas-
toideus—and clavobrachialis complex. This somewhat complex muscle arrangement, occur-
ring over the point of the shoulder, is depicted in figures 9, 10, and 11. The essential feature
is an interchange of muscle fibers between the clavotrapezius (3) and the cleidomastoideus
(4) resulting in the formation of the clavobrachialis muscle (5).

This view shows a division of the clavotrapezius (3) into an anterior one-third which
spreads out to insert into the pectoral fascia, and a posterior two-thirds being joined by
fibers of the cleidomastoideus (4) to form the clavobrachialis muscle (5). Not shown in this
plate is the continuation of the cleidomastoideus (4) and its being joined by muscle fibers
from the clavotrapezius (3); see figure 11 for this detail. Key: 3, clavotrapezius; 4, cleido-
mastoideus; 5, clavobrachialis; 6, omotrachlean (levator scapulae ventralis); 13, pectoralis
major; 26, supraspinatus; 27, subcapularis; A, pectoral fascia; B, sternomastoideus muscle;
S, scapula, superior border.

VoL. XXXIX] HOWARD: SEA OTTER FORELIMB 4

bo
On

Ficure 10. Anterolateral view of the right shoulder area identical to figure 8, except
that the clavotrapezius muscle (3) has been detached from its origin and reflected laterally
and posteriorly. The sternomastoideus muscle (C) has been reflected anteriorly. The muscle
interchange to form the clavobrachialis (5) is again shown. Not shown (as in figure 9) is
the continuation of the cleidomastoideus muscle (4). Key: 3, clavotrapezius; 4, cleidomas-
toideus; 5, clavobrachialis; 6, omotrachlean (levator scapulae ventralis); 9, rhomboideus
profundus; 13, pectoralis major; 26, supraspinatus; 27, subscapularis; A, mastoid process
of the skull; B, transverse process of the first cervical vertebra; C, sternomastoideus muscle;
S, scapula, superior margin.

the mastoid process of the skull, the muscle passes in uniform diameter under
the clavotrapezius and over the omotrachleon toward the shoulder at which
point the muscle splits into a posterior one-fourth and an anterior three-fourths.

INSERTION. The posterior one-fourth joins with the muscle fibers from

426 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

M

Sem

FicurEe 11. An enlarged view of the right shoulder area and upper humerus to show
the continuation of the cleidomastoideus muscle (4) and its receiving of muscle fibers from
the clavotrapezius (3) (depicted by dotted line). Also shown is the insertion of the clavo-
brachialis (5) and the acromiodeltoideus (17). Key: 3, clavotrapezius; 4, cleidomastoideus;
5, clavobrachialis; 17, acromiodeltoideus; 30, brachioradialis; H, humerus.

VoL. XXXIX] HOWARD: SEA OTTER FORELIMB 427

the clavotrapezius to form the clavobrachialis muscle. The anterior three-
fourths joins with the remaining muscle fibers of the clavotrapezius. Thus re-
inforced, this combined muscle continues distally into the anterior cubital area
of the elbow where it joins with the pectoantebrachialis muscle which, in turn,
inserts into the ulna.

AcTION. With this somewhat complicated insertion it would be difficult
to analyze the specific action of this muscle. From its position the muscle could
assist in both flexion of the elbow and shoulder and drawing of the forelimb
forward and dorsally.

5. Clavobrachialis.

This is a rather small flat muscle passing over the shoulder area.

Oricin. This muscle is made up of fibers from the clavotrapezius and the
cleidomastoideus as described in the description of each. The newly formed
muscle converges to a longitudinal linear configuration.

INSERTION. The muscle fibers insert into the humerus along the pectoral
ridge at its junction with the deltoid ridge. The insertion is adjacent to and
just posterior to the insertion of the pectoralis major muscle and just distal to
the insertion of the spinodeltoideus and acromiodeltoideus muscles.

AcTION. This small muscle, by virtue of its position, could assist in abduc-
tion and flexion of the humerus.

6. Omotrachleon (levator scapulae ventralis).

This is a long, flat, ribbonlike muscle connecting the scapula to the spine.

ORIGIN. This muscle arises by fleshy fibers from the anteromedial surface
of the transverse process of the first cervical vertebra. At its point of origin
it overlies the origin of the rhomboideus profundus muscle. The muscle passes
obliquely posteriorly across the neck toward the acromion process of the scapula.

INSERTION. About one-fourth of the fibers split off anteriorly just before
reaching the acromion process of the scapula. These fibers join the panniculus
carnosus muscle of the forelimb. The remaining fibers insert into the posterior
border of the acromion process of the scapula.

AcTIon. This muscle serves to support the scapula and rotate it counter-
clockwise.

7. Rhomboideus major.

This thin, flat, triangular muscle lies beneath the trapezius muscle complex
and connects the scapula with the axial skeleton.

OricIn. Fleshy fibers arise from the middorsal line of the cervical area
starting just posterior to the external occipital crest and continue posteriorly
the full length of the cervical spine into the thoracic area to terminate at the

428 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

FicurE 12. Lateral view of the right shoulder and neck area to show origin and insertion
of omotrachlean muscle (levator scapulae ventralis) (6) and its relation to the other muscles
in the area. The rhomboideus major (7) and the rhomboideus capitis (8) have been reflected
dorsally. The clavotrapezius (3) detached at its origin, is reflected anteriorly for clear
exposure of the underlying musculature. The spinotrapezius has been removed. Key: 2, acro-
miotrapezius; 3, clavotrapezius; 4, cleidomastoideus; 6, omotrachlean (levator scapulae ven-
tralis) ; 7, rhomboideus major; 8, rhomboideus capitis; 9, rhomboideus profundus; 17, acro-
miodeltoideus; 18, spinodeltoideus; 19, dorsoepitrochlearis; 20, triceps brachii caput longus;
24, infraspinatus; 27, subscapularis; A, to the panniculus carnosus; B, transverse process
of first cervical vertebra; C, sternomastoid muscle; S, scapula, acromion process.

dorsal spine of T-3. From its origin the muscle triangulates toward the posterior
portion of the vertebral border of the scapula.

INSERTION. Fleshy and tendinous fibers insert along the vertebral border
of the scapula, starting at the inferior angle and extending anteriorly to the
origin of the scapular spine. At this point, a tendinous raphe is formed which

VoL. XXXIX] HOWARD: SEA OTTER FORELIMB 429

E

yi a I ad

Ficure 13. Dorsolateral view of right cervical and scapular to show the rhomboid
muscle complex. This group of three muscles, the rhomboideus major (7), capitis (8), and
profundus (9), form a sheet-like layer beneath the trapezius muscle group. In this plate
the spinotrapezius (1) and the acromiotrapezius (2) have been detached at their origins
and reflected ventrally. The clavotrapezius has been removed. As noted from their origins
and insertions, these three rhomboid muscles act to support the pectoral girdle drawing it
forward and rotating it clockwise. Key: 1, spinotrapezius; 2, acromiotrapezius; 4, cleido-
mastoideus; 6, omotrachlean (levator scapulae ventralis); 7, rhomboideus major; 8, rhom-
boideus capitis; 9, rhomboideus profundus; 24, infraspinatus; 25, supraspinatus; A, mastoid
process of skull; B, deep cervical musculature; C, lambdoid ridge of skull; D, spinous pro-
cess of first dorsal vertebra; 5, middorsal line, cervical area; S, outline of scapular.

spans the area of origin of the spine, then joins the remaining fibers to insert
along the anterior ridge of origin of the scapular spine. This latter portion over-
lies and more-or-less joins with the insertion of the rhomboideus capitis.

Action. This muscle pulls the scapula forward as in shrugging the shoulder.
It can also rotate the scapula clockwise.

8. Rhomboideus capitis.

The second of the rhomboideus muscle group is a thin, flat, elongated muscle
connecting the skull with the scapula. The muscle parallels to a degree the in-
ferior border of the rhomboideus major and underlies the trapezius musculature
in this area.

Oricin. Arising by fleshy fibers from the lateral one-third of the lambdoidal

430 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Ficure 14. Dorsolateral view of cervical and scapular area identical with figure 13
except that the rhomboideus major (7) has been detached from its origin and reflected
posteriorly at its insertion. The origin and insertion of the rhomboideus capitis (8) is shown.
Key: 1, spinotrapezius—reflected; 2, acromiotrapezius—reflected; 4, cleidomastoideus;
6, omotrachlean; 7, rhomboideus major—reflected; 8, rhomboideus capitis; 9, rhomboideus
profundus; 24, infraspinatus; 26, supraspinatus; A, mastoid process of the skull; S, outline
of scapula.

crest and the jugular process of the skull, the muscle courses inferiorly to pass
over the supraspinatus fossa of the scapula where it widens in a flare-type man-
ner terminating in a thin, flat, tendinous, sheath.

INSERTION. This flat, thin tendon inserts along the spine of the scapula
more or less fusing with the tendon of the rhomboideus major but also continues
its insertion along the vertebral margin of the scapula superiorly approximately
two-thirds of the distance to the first angulation of this margin.

ActTIon. The muscle draws the scapula forward and acts to rotate the scap-
ula clockwise.

9. Rhomboideus profundus.

The third and deepest of the rhomboid muscles underlies, for the most part,
the rhomboideus capitis and connects the spine to the scapula.

VoL. XXXIX] HOWARD: SEA OTTER FORELIMB 431

Ficure 15. Lateral view of right scapula. The rhomboideus major (7) and the rhom-
boideus capitis (8) have been reflected posteriorly to show the detail of insertion of the
rhomboideus profundus (9) on the scapula. Again, the spinotrapezius (1) and the acro-
miotrapezius (2) have been reflected as in the previous plate. Key:° 1, spinotrapezius—re-
flected; 2, acromiotrapezius—reflected; 7, rhomboideus major—reflected; 8, rhomboideus
capitis—reflected; 9, rhomboideus profundus; 24, infraspinatus; 26, supraspinatus; S, out-
line of scapula.

OricIn. This muscle arises by fleshy fibers from the inferior surface of the
transverse process of C-1 vertebra overlying the origin of the omotrachleon mus-
cle. From this point the strap-type muscle passes obliquely inferiorly to underlie
the rhomboideus capitis as it reaches the scapula area. Near its insertion the
muscle spans out ending in a thin tendinous sheath in a manner similar to the
rhomboideus capitis.

INSERTION. The tendon of the rhomboideus profundus inserts with that of
the rhomboideus capitis but extends more superiorly along the vertebral margin
of the scapula to the first angulation of the vertebral border.

432 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

AcTIon. This muscle’s action is the same as that of the rhomboideus capitis,
drawing the scapula forward and rotating it clockwise.

10. Serratus magnus.

This large, flat, interdigitated muscle connects the rib cage with the ver-
tebral margin of the scapula, and is the posterior portion of a common suspen-
sory sheet of muscle which includes superiorly the levator anguli scapulae. The
muscle is best observed as the vertebral margin of the scapula is spread out
from the rib cage.

OriciIn. Arising by fleshy digitations from ribs 1 to 6; starting on rib 1
near the spine the origin continues in an oblique line to rib 6 at about the level
of the midlateral line of the chest. The muscle triangulates from its origin to-
ward the vertebral margin of the scapula and, in doing so, the digitation is
joined into a flat muscular sheath, the superior margin of which becomes con-
tinuous with the levator anguli scapulae.

INSERTION. The serratus magnus portion of this muscle sheath inserts
on the vertebral margin of the scapula starting at the inferior angle and extend-
ing superiorly to the level of origin of the scapular spine where it joins the leva-
tor anguli scapulae. The insertion is deep to that of the rhomboideus major.

Action. This muscle acts to hold the scapula against the chest wall and
serves to suspend the trunk in weight bearing. It can also move the scapula in
the line of its fibers.

11. Levator anguli scapulae.

This muscle is, in effect, the superior continuation of the serratus magnus
and connects the cervical spine to the scapula.

OricIN. Fleshy fibers arise from the transverse processes and intervening
fascia of C-3 to C-7. A sheet of muscle is formed which triangulates toward
the vertebral margin of the scapula. The inferior border approximates the
superior border of the serratus magnus distally to form a common muscle.

INSERTION. The levator anguli scapulae portion of the muscle sheath in-
serts superiorly to the serratus magnus in a similar manner along the vertebral
margin of the scapula from the origin of the scapular spine halfway to the
superior angle of the scapula.

Action. In view of the muscle’s more dorsal origin, the action is to sus-
pend the scapula and rotate the scapula clockwise. It also assists the serratus
magnus muscle in holding the vertebral margin of the scapula to the chest wall.

12. Pectoantebrachialis.

This muscle is the most superficial of the pectoral muscle group. It is a
large, strong, triangular muscle connecting the trunk to the forearm.

VoL. XXXIX] HOWARD: SEA OTTER FORELIMB 433

Ficure 16. Dorsal view of right-shoulder area. The trapezius muscles have been re-
moved. The rhomboideus major (7) is shown reflected outward from the vertebral margin
of the scapula. The scapula has been spread outward from the body to disclose the serratus
magnus (10) and the levator anguli scapulae (11) muscles. These broad, flat but thick and
strong muscles act as a sling support for the trunk between the forelimbs. The two muscles
form a common and continuous muscle sheath, their division being based on the site of ori-
gin, the levator anguli scapulae (11) from the cervical area, serratus magnus (10) from the
rib cage. Key: 7, rhomboideus major—reflected; 10, serratus magnus; 11, levator anguli
scapulae; 27, subscapularis; A, fifth rib; B, deep cervical musculature; C, middorsal line
cervical area; D, transverse process of third cervical vertebra; S, scapula, vertebral margin.

OricIn. Fleshy fibers arise from the superior part of the sternum start-
ing at the upper border of the manubrium and reaching to between ribs 1 and
2. The muscle then courses obliquely inferiorly over the pectoral musculature
toward the anterior cubital area where the muscle converges and, joined by

434 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Ficure 17. Anterior view of the right humerus and upper chest with the forelimb ro-
tated externally. The cleidomastoideus (4) and clavotrapezius (3) are reflected laterally.
The sternomastoid muscle (A) is detached at its insertion and reflected superiorly. The pec-
toral muscle complex is thus revealed. This complex consists of three flat, thick, and strong
muscles arising from the anterior upper chest, connecting the trunk to the upper extremities.
The most superficial is the pectoantebrachialis (12) which overlies the pectoralis major (13),
which in turn overlies the pectoralis minor (14). This strong muscle group flexes and in-
ternally rotates the humerus. In addition, action of the pectoantebrachialis (12) is to assist
in flexion of the elbow. Further detail of the relationship of these three muscles is shown
in figures 18, 19, and 20. Key: 3, clavotrapezius—reflected; 4, cleidomastoideus—reflected ;
5, clavobrachialis—reflected; 12, pectoantebrachialis; 13, pectoralis major; 14, pectoralis
minor; 19, dorsoepitrochlearis; 20, triceps brachii caput longus; 22, triceps brachii caput
medialis; 26, supraspinatus; 28, biceps brachii; 30, brachioradialis; A, sternocleidomastoid
muscle—reflected; B, sternum, manubrium; C, sternum, corpus; D, extensor muscles of
the forearm; E, flexor muscles of the forearm; H, humerus; U, ulna.

fibers from both the cleidomastoideus laterally and pectoralis major medially,
forms a strong tendon.

INSERTION. Passing deep to the tendon of insertion of the biceps, the
heavy tendon of the pectoantebrachialis inserts on the medial ridge of the ulna
at the conoid process just distal to the joint surface.

AcTION. Strong flexion of the elbow is accomplished plus assistance in
flexion of the humerus.

VoL. XXXIX] HOWARD: SEA OTTER FORELIMB 435

Ficure 18. Anterior view of the right upper chest and humerus with the forelimb ro-
tated externally. This is essentially the same view as figure 17 but with the pectoantebrachi-
alis (12) reflected laterally to show the pectoralis major (13) and the lower fibers of the mus-
cle joining with the pectoantebrachialis (12) which in turn joins the cleidomastoideus (4)
for common insertion into the proximal ulna. The lower part of the pectoralis minor has
been omitted in this drawing. Key: 4, cleidomastoideus; 5, clavobrachialis; 12, pectoante-
brachialis—reflected; 13, pectoralis major; 14, pectoralis minor; 19, dorsoepitrochlearis;
20, triceps brachii caput longus; 22, triceps brachii caput medialis; 26, supraspinatus; 28,
biceps brachii; 30, brachioradialis; A, sternocleidomastoid—reflected; B, sternum, manu-
brium; C, sternum, corpus; D, extensor muscles of forearm; E, flexor muscles of forearm;
H, humerus; U, ulna.

13. Pectoralis major.

This strong, broad, rhomboid muscle of the pectoral group underlies the pec-
toantebrachialis and connects the sternum to the humerus. The muscle is heavi-
est superiorly.

OricIn. There is a continuous origin of fleshy fibers from the top of the
manubrium to the fourth rib level inferiorly. This forms the base of the muscle
which then continues transversely as a broad sheet to the humerus. In so do-
ing, the muscle overlies the pectoralis minor and bridges the anterior chest.
Short thin tendinous fibers develop on the underside as the humerus is ap-
proached.

436 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Ficure 19. An anterior view of the right shoulder and upper arm musculature in a
limited area. The pectoantebrachialis (12) has been detached from its origin and reflected
inferiorly to show its contribution of muscle fibers to the pectoralis major (13) near its distal
insertion on the humerus, and also the joining of the cleidomastoideus (4) with the pecto-
antebrachialis (12) to form a common tendon. Key: 4, cleidomastoideus; 5, clavobrachialis;
12, pectoantebrachialis; 13, pectoralis major; 14, pectoralis minor; 26, supraspinatus; 27,
subscapularis; S, scapula, supraglenoid area of superior border.

INSERTION. The insertion of this muscle extends practically the entire
length of the humerus, starting superiorly at the greater tuberosity at the mar-
gin of insertion of the supraspinatus tendon and extending distally along the
lateral side of the pectoral ridge and humeral crest to the cubital fossa area.

VoL. XXXTIX] HOWARD: SEA OTTER FORELIMB 437

De

Ficure 20. Anterior view of right shoulder area with the forelimb adducted and exter-
nally rotated. The pectoantebrachialis (12) and the pectoralis major (13) have been de-
tached from their origins and reflected laterally, thus exposing the insertion of the under-
lying pectoralis minor (14). With adduction of the forelimb the latissimus dorsi (15) comes
into view and overlies the triceps muscle group. Note raphe-type insertion of the upper part
of the pectoralis major (13). Also shown are the pectoralis major (13) fibers joining the
pectoantebrachialis (12). Key: 12, pectoantebrachialis; 13, pectoralis major; 14, pectoralis
minor; 15, latissimus dorsi; 26, supraspinatus; 27, subscapularis; 28, biceps brachii; A,
raphe-type insertion of pectoralis major; B, forearm flexor musculature; H, humerus; S,
scapula, superior margin; U, ulna.

At the upper end, a tendinous raphe is present, and at the lower end some fleshy
and tendinous fibers join the pectoantebrachialis muscle.

Action. This muscle is a strong flexor and internal rotator of the humerus,
and it also assists in adduction of the forelimb.

14. Pectoralis minor.

This deepest muscle of the pectoral group underlies the pectoralis major
and joins the trunk to the humerus.

438 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Ficure 21. Lateral view of right thoracic and shoulder area to show origin of large and
strong latissimus dorsi (15) muscle from the trunk, and its relation to the scapula and ad-
jacent musculature. The spinotrapezius muscle which overlies the dorsal and anterior area
of the latissimus dorsi (15) has been removed. Key: 7, rhomboideus major; 15, latissimus
dorsi; 19, dorsoepitrochlearis; 24, infraspinatus; A, spinous process of eighth dorsal vertebra;
S, outline of scapula.

Oricin. Arising by fleshy fibers from the border of the sternum, rib levels
2 to 7, this thin flat rectangular muscle courses toward the humerus under the
pectoralis major and over the biceps muscle. There is a tendency towards
clefting of the muscle in its upper part between ribs 2 and 3. Near the humerus
a broad thin tendon develops.

INSERTION. The tendon inserts on the humerus just medial and parallel to
the pectoralis major along the length of the pectoral ridge. The uppermost and
more fleshy fibers insert on the greater tuberosity of the humerus just medial
to the tendon of insertion of the supraspinatus. At the lower or most distal end
the tendon is joined by fleshy and tendinous fibers of the anterior half of the
split latissimus dorsi muscle for common insertion.

Action. Along with the pectoralis major this muscle flexes, internally ro-
tates, and adducts the humerus.

VoL. XXXIX] HOWARD: SEA OTTER FORELIMB 439

Ficure 22. Anterior view of the right shoulder and humerus with the upper extremity
in external rotation. The pectoral muscles (13) and (14) are reflected laterally exposing the
biceps brachii (28) and the teres major (16). The biceps brachii is displaced a bit laterally.
The latissimus dorsi (15) is now viewed as the two halves reach their insertion on the
humerus. A common tendon of insertion is formed with the teres major and the pectoralis
major. Key: 13, pectoralis major; 14, pectoralis minor; 15, latissimus dorsi; 16, teres major;
19, dorsoepitrochlearis; 20, triceps brachii caput longus; 22, triceps brachii caput medialis;
26, supraspinatus; 27, subscapularis; 28, biceps brachii; A, tuberosity of the humerus; H,
humerus, S, scapula, superior margin.

15. Latissimus dorsi.

This is a large, flat, strong muscle, triangular in shape, covering most of
the thorax from T-3 to T-12 and joining the trunk to the humerus.

Oricin. Arising by fleshy fibers from the middorsal line and spinous processes
from T-3 to T-9 and then from the lumbar fascia to T-12 level inferiorly and
ribs 10 and 11 level midlaterally. This large sheetlike muscle triangulates to-
ward the inferior angle of the scapula, which it crosses to disappear under the
dorsoepitrochlearis. At about this point the muscle divides into a dorsal, or
proximal, two-fifths and a ventral, or distal, three-fifths, but both divisions con-

440 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

5 Cim

Ficure 23. Anterior view of the right shoulder and humerus with upper extremity in
external rotation. This is essentially the same view as in figure 22 except that the biceps
brachii (28) is now displaced medially to disclose the detail of insertion of that portion of
the latissimus dorsi (15) which joins the pectoralis minor (14). The pectoral muscles (13)
and (14) are again reflected laterally. Key: 13, pectoralis major; 14, pectoralis minor; 15,
latissimus dorsi; 16, teres major; 19, dorsoepitrochlearis; 20, triceps brachii caput longus;
22, triceps brachii caput medialis; 26, supraspinatus; 27, subscapularis; 28, biceps brachii;
A, tendon fibers begin here; H, humerus; S, scapula, superior border.

tinue under the triceps musculature to gain access to the medial side of the hu-
merus.

INSERTION. At this point, the proximal two-fifths portion narrows and flat-
tens into a tendon which passes between the biceps brachii and the humerus to
join that of the teres major for a common insertion on the humerus along the
pectoral ridge and just under the insertion of the pectoralis minor. The distal
three-fifths of the muscle passes as fleshy fibers anterior to the biceps brachii,
then, spreading into flat tendinous fibers, joins the deep side of the tendon of
the pectoralis minor for a common insertion with this tendon.

VoL. XXXIX] HOWARD: SEA OTTER FORELIMB 441

yf

— —o——08 ut

Ficure 24. Lateral view of the right shoulder and forelimb. The relationship of the
acromiodeltoideus (17) and the spinodeltoideus (18) is shown. Also, the insertion of the
levator anguli scapulae (11) on the acromion process of the scapula is seen. The clavobrachi-
alis (5) has been reflected anteriorly to show its tendinous insertion on the humerus. The
relationship of the other muscles in this lateral view is to be noted. Key: 1, spinotrapezius;
2, acromiotrapezius; 3, clavotrapezius; 4, cleidomastoideus; 5, clavobrachialis; 7, rhom-
boideus major; 11, levator anguli scapulae; 15, latissimus dorsi; 16, teres major; 17, acro-
miodeltoideus; 18, spinodeltoideus; 19, dorsoepitrochlearis; 20, triceps brachii caput longus;
21, triceps brachii caput lateralis; 23, anconeus externus; 30, brachioradialis; 31, extensor
carpi radialis longus; 32, extensor carpi radialis brevis; 33, extensor digitorum communis;
34, extensor digitorum lateralis; 35, extensor carpi ulneris; A, forearm fascia; B, panniculus
carnosa; C, scapula, acromion process; H, humerus; U, ulna; S, scapula, vertebral border.

442 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Ficure 25. Enlarged localized lateral view of the right shoulder area to show detail of
spinodeltoideus (18) joining acromiodeltoideus (17) to gain insertion on the humerus. To
show this adequately the acromiodeltoideus (17) has been rotated anteriorly, thus exposing
the undersurface where the juncture of the two muscles occurs. Key: 5, clavobrachialis;
11, levator anguli scapulae; 16, teres major; 17, acromiodeltoideus; 18, spinodeltoideus;
19, dorsoepitrochlearis; 20, triceps brachii caput longus; 21, triceps brachii caput lateralis;
30, brachioradialis; H, humerus; S, scapula, acromion process.

Action. This muscle is a strong extensor and internal rotator of the hu-
merus.

16. Teres major.

This short but strong muscle connects the inferior border of the scapula with
the humerus.

OricIn. Muscle fibers arise from the inferior angle and dorsal one-half of
the inferior border of the scapula with some tendinous fibers from the fascia
overlying the infraspinatus muscle in this area to form a somewhat cone-shaped
muscle paralleling the inferior scapular border passing directly toward the hu-

VoL. XXXIX] HOWARD: SEA OTTER FORELIMB 443

FicurE 26. The triceps muscle complex is a group of strong muscles arising from the
scapula or humerus with insertion on the olecranon process of the ulna to give extension of
the elbow joint. The triceps musculature proper consists of the triceps brachii caput longus
(20), lateralis (21), and medialis (22). The caput medialis has basically three divisions,
having a somewhat complicated configuration and relationship to the caput lateralis (see
figure 27). This triad of muscles is reinforced by the large dorsoepitrochlearis (19) which,
due to its site of origin from the scapular margin, is responsible for the posterior webbing
between the humerus and the scapula. Also, the small anconeus externus (23) is an accessory
to this group (see figure 29).

This figure is a posterolateral view of the upper right extremity to show detail of the
triceps brachii caput longus (20) and lateralis (21), and the dorsoepitrochlearis (19). Note
a few tendon fibers (C) from the dorsoepitrochlearis (19) joining the tendon of the triceps
brachii caput longus (20). Key: 11, levator anguli scapulae; 15, latissimus dorsi; 16, teres
major; 17, acromiodeltoideus; 18, spinodeltoideus; 19, dorsoepitrochlearis; 20, triceps brachii
caput longus; 21, triceps brachii caput lateralis; A, deep fascia of the forearm; B, forearm
extensor musculature; C, tendon fibers, from (19) entering tendon of (20); H, humerus;
S, outline of scapula, axillary border; U, ulna, olecranon process.

merus. Near the humerus the muscle is joined by muscle and tendon fibers of the
superior portion of the latissimus dorsi. A flat strong tendon forms which passes
under the biceps brachii.

INSERTION. This tendon now inserts into the humerus on the medial side
and nearly parallel to the insertion of the pectoralis minor.

Action. Adduction and internal rotation of the humerus is accomplished
by this muscle.

444 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Ficure 27. Medial view of the right forelimb and shoulder area with the forelimb in
full external rotation to show the triceps brachii caput medialis (22), its various parts, and
their relationships to each other and to the caput lateralis (21). The caput longus (20) of
the triceps has been completely removed. The biceps brachii (28) is displaced anteriorly to
visualize the deeper triceps brachii caput medialis (22).

Note the three (22) labels which mark the three parts of the medial head. The upper
(22) label is the intermediate part (anconeus internus). The middle (22) label is the long
part (anconeus medialis). The lower (22) label is the medial part (anconeus medialis). Key:
21, triceps brachii caput lateralis; 22, triceps brachii caput medialis; 25, teres minor; 27,
subscapularis; 28, biceps brachii; 29, brachialis; 30, brachioradialis; A, site of origin of
triceps brachialis, caput longus; B, medial epicondylar ridge of humerus; C, forearm flexor
musculature; H, humerus, articular surface of head; S, scapula, infraglenoid area; U, ulna,
olecranon process.

VoL. XXXIX] HOWARD: SEA OTTER FORELIMB 445

Ficure 28. The shoulder cuff muscles of the scapula consist of the subscapularis (27),
supraspinatus (26), infraspinatus (24), and teres minor (25). All four of these muscles
arise from the surface of the scapula and terminate in a tendinous cuff about the shoulder
which inserts on the humerus, stabilizing and activating this joint. The tendons are inti-
mately associated with the shoulder joint capsule.

This right lateral view of the scapula and shoulder area has all muscles removed that
attach to the scapula except for those arising from the scapular fossae (lateral surface). Key:
21, triceps brachii caput lateralis; 24, infraspinatus; 25, teres minor; 26, supraspinatus; 30,
brachioradialis; A, extensor musculature of the forearm; H, humerus, greater tuberosity;
S, scapula, acromion process.

17. Acromiodeltoideus.

Joining the scapula to the humerus, this small but stout triangular muscle
overlies the point of the shoulder.

OriciIn. From the full width of the most distal margin of the acromion
process muscle fibers arise on the undersurface and tendinous fibers on the
outer surface to form a short muscle which then passes over the lateral aspect
of the shoulder joint to the upper humerus.

INSERTION. With tendinous fibers developing on the undersurface but
fleshy fibers persisting on the outer surface, the muscle inserts into the distal
area of the deltoid ridge after first receiving the tendinous and the fleshy fibers

446 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Ficure 29. Anterolateral view of the right scapula and shoulder to show the detail of
insertion of the supraspinatus (26) in relation to the bicipital groove and infraspinatus (24)
and subscapularis (27) muscles. Key: 21, triceps brachii caput longus; 24, infraspinatus;
26, supraspinatus; 27, subscapularis; 28, biceps brachii; 30, brachioradialis; A, extensor
musculature of the forearm; B, biceps brachii tendon in bicipital groove of humerus; C,
humerus, lesser tuberosity; H, humerus, greater tuberosity; S, scapula, acromion process.

VoL. XXXIX] HOWARD: SEA OTTER FORELIMB 447

Ficure 30. Medial view of the right scapula and shoulder to show the detail of sub-
scapularis muscle (27). The biceps brachii (28) has been displaced somewhat anteriorly and
the upper tendon lifted out of the bicipital groove of the humerus. Key: 21, triceps brachii
caput lateralis; 22, triceps brachii caput medialis; 26, supraspinatus; 27, subscapularis; 28,
biceps brachii; 29, brachialis; A, biceps brachii tendon out of bicipital groove; H, humerus,
lesser tuberosity; S, scapula, axillary border.

of the spinodeltoideus muscle. The insertion is linear between the insertion of the
pectoralis muscles and the origin of the brachioradialis.

Action. This muscle can abduct the humerus and possibly give slight ex-
ternal rotation as well.

18. Spinodeltoideus.

This small triangular muscle joins the spine area of the scapula to the hu-
merus via the acromiotrapezius and courses with convergence toward the acro-
miodeltoideus muscle.

INSERTION. Both fleshy and tendinous fibers join the undersurface of the
acromiodeltoideus and proceed to a common insertion with this muscle on the
humerus just proximal to the insertion of the clavobrachialis.

Action. Acting with the acromiodeltoideus some degree of abduction, ex-
tension, and external rotation of the humerus is accomplished.

448 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

|

“ rare A ee

Ficure 31. Anteromedial view of the right forelimb to show origin and insertion of the
biceps brachii muscle (28). The supraspinatus muscle (26) and the subscapularis muscle
(27) have been retracted sufficiently to disclose the upper border of the glenoid. The pecto-
antebrachialis (12) has been excised near its insertion. Key: 12, pectoantebrachialis; 22,
triceps brachii caput medialis; 26, supraspinatus; 27, subscapularis; 28, biceps brachii; 29,
brachialis; 30, brachioradialis; A, greater tuberosity of the humerus; B, lesser tuberosity of
the humerus; C, forearm flexor musculature; D, forearm extensor musculature; S, scapula,
supraglenoid area; R, radius; U, ulna.

VoL. XXXIX] HOWARD: SEA OTTER FORELIMB 449

Ficure 32. Lateral view of the right upper extremity to show the brachialis and the
brachioradialis muscles. The triceps mechanism has been displaced posteriorly somewhat
and the more superficial muscles of the upper arm and shoulder area have been removed.
The superficial sensory branch of the radial nerve is shown displaced from its normal position
between the brachioradialis (30) and the radial wrist extensors (31) (32). Key: 21, triceps
brachii caput lateralis; 22, triceps brachii caput medialis; 24, infraspinatus; 25, teres minor;
26; supraspinatus; 29, brachialis; 30, brachioradialis; 31, extensor carpi radialis longus; 32,
extensor carpi radialis brevis; 33, extensor digitorum communis; 34, extensor digitorum
lateralis; 35, extensor digitorum ulnaris; A, radial nerve; H, humerus; R, radius (radial sty-
loid) ; S, scapula, acromion process.

19. Dorsoepitrochlearis.

This strong, somewhat flat and triangular, fleshy muscle connects the scap-
ula to the forearm on the extensor surface, thus becoming a functional part of
the triceps muscle complex.

450 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

5iem

Ficure 33. Posterior view of the right upper arm to show origin and insertion of anco-
neus externus (23). Triceps brachii caput lateralis (21) has been detached near its insertion
and displaced medialward. The triceps brachii caput longus (20) has been removed except
for its tendon of origin and insertion. The scapula is located presenting the axillary border.
Key: 20, triceps brachii caput longus; 21, triceps brachii caput lateralis; 22, triceps brachii
caput medialis; 23, anconeus externus; 24, infraspinatus; 27, subscapularis; 30, brachioradi-
alis; 31, extensor carpi radialis longus; 32, extensor carpi radialis brevis; 33, extensor digi-
torum communis; 34, extensor digitorum lateralis; H, humerus, lateral epicondylar crest;
R, radius, radial head outline; S, scapula, infraglenoid area; U, ulna, olecranon process.

VoL. XXXIX] HOWARD: SEA OTTER FORELIMB 451

Ficure 34. Lateral view of right forearm and elbow area to show relationship of the
superficial dorsal musculature. Note the transverse dorsal carpal ligament at wrist level (A).
Key: 21, triceps brachii caput lateralis; 23, anconeus externus; 30, brachioradialis; 31, ex-
tensor carpi radialis longus; 32, extensor carpi radialis brevis; 33, extensor digitorum com-
munis; 34, extensor digitorum lateralis; 35, extensor carpi ulnaris; 36, extensor pollicis et
indicis longus; 38, abductor pollicis; 39, flexor carpi ulnaris; A, transverse dorsal carpal
ligament; B, radial nerve, distal portion displaced from its bed; R, radius, outline of radial
head; U, ulna, olecranon process.

OricIn. The muscle arises by fleshy and tendinous fibers from the dorsal
three-fifths of the inferior border of the scapula adjacent to and overlying the
origin of the teres major muscle. From its origin, the muscle triangulates toward
the point of the elbow paralleling the long head of the triceps brachii and over-
lying the latissimus dorsi.

INSERTION. At the elbow a flat tendon forms which overrides the olecranon
process. Except for a few short tendinous fibers which join the tendon of the
long head of the triceps brachii, this muscle inserts into the deep fascia of the
upper part of the forearm, mostly on the medial side.

AcTION. Strong extension of the elbow is accomplished.

20. Triceps brachii caput longus.

This strong spindle-shaped muscle with a tendon at each end joins the scapula
to the ulna, and is part of the triceps muscle complex.

OricIN. Arising as a heavy tendon with but few fleshy fibers from the in-
ferior surface of the neck of the scapula adjacent to the glenoid, the muscle
courses distally down the posterior surface of the humerus between the dorso-

452 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Ficure 35. An anterolateral view of the right upper extremity to show the relationship
of the brachialis (29) to the pronator teres (44) and the supinator (37) muscles. The radial
carpal extensors (31) and (32) have been reflected at their origin and the brachioradialis
(30) has been excised except for its insertion on the distal radius. The muscles overlying the
brachialis (29) have been removed. Key: 29, brachialis; 30, brachioradialis; 31, extensor
carpi radialis longus; 32, extensor carpi radialis brevis; 33, extensor digitorum communis;
34, extensor digitorum lateralis; 35, extensor carpi ulnaris; 37, supinator, 38, abductor pol-
licis longus; 44, pronator teres; R, radius, radial styloid area; U, ulna, coronoid area.

VoL. XXXIX] HOWARD: SEA OTTER FORELIMB 453

epitrochlearis and the triceps brachii caput lateralis directly to the olecranon
process of the ulna.

INSERTION. A heavy tendon terminating this muscle inserts into the ole-
cranon process of the ulna posteriorly and medially. Near its actual bony inser-
tion it receives the tiny tendinous slip from the dorsoepitrochlearis muscle.

AcTIon. Strong extension of the elbow results from contraction of this mus-
cle.

21. Triceps brachii caput lateralis.

This is another strong fusiform muscle of the triceps muscle complex. This
muscle connects the upper humerus with the ulna.

OricIn. Fleshy and tendinous fibers arise from the medial surface of the
neck of the humerus and adjacent shoulder capsule. The muscle thus formed
courses distally to the lateral side of the elbow adjacent to the triceps brachii
caput longus and overlies the origin of the brachialis muscle. In its midportion
it receives muscle fibers from the medial head of the triceps brachii, and again
in its distal portion more fibers from the medial head join in a tendinous raphe.

INSERTION. The muscle terminates at the elbow in a short broad tendon
which inserts on the posterolateral side of the olecranon process of the ulna ad-
jacent to and just distal to the insertion of the long head of the triceps brachii.

AcTION. This muscle joins the other members of the triceps muscle com-
plex to produce strong extension of the elbow joint.

22. Triceps brachii caput medialis.

This muscle is the lesser of the three triceps muscles and consists of three
distinct parts, all of which are closely approximated to the humerus. The long
(anconeus posterior) and the intermediate (anconeus internus) portions are
more closely associated. The medial division (anconeus medialis) is more apart,
much shorter, and much more distal.

OricIn. The long portion arises from the posteromedial surface of the up-
per humerus by thin tendinous fibers from as far proximal as the level of the
lesser trocanter. The intermediate portion arises to the lateral side of the long
portion but takes origin by fleshy and tendinous fibers more proximally from
the posteromedial surface of the neck of the humerus and adjacent capsule of the
shoulder joint. Both portions progress distally and unite into a common single
muscle belly after the intermediate portion first gives off muscle fibers to the
caput lateralis triceps brachii. The medial portion (anconeus medialis) arises
distally on the humerus from the posterior surface of the medial epicondylar
flare and the fleshy fibers approach the elbow.

INSERTION. The tendons of the caput medialis insert on the medial side
of the olecranon process.

454 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

12

Ficure 36. Anterior view of the right forearm in midposition to show the pronator
(44) supinator (37) relationship. Insertions of the brachialis (29) and the pectoantebrachi-
alis (12) are shown on the coronoid area of the ulna. Also seen is the insertion of the lower
biceps tendon on the tubercle of the radius. The radial carpal extensors (31) (32) and the
brachioradialis (30) muscles are reflected. Key: 12, pectoantebrachialis; 28, biceps brachii;
29, brachialis; 30, brachioradialis; 31, extensor carpi radialis longus; 32, extensor carpi ra-
dialis brevis; 33, extensor digitorum communis; 36, extensor pollicis et indicis longus; 38,
abductor pollicis longus; 44, pronator teres; 43, flexor carpi radialis; H, humerus; R, ra-
dius, tubercle.

VoL. XXXIX] HOWARD: SEA OTTER FORELIMB 455

Action. This muscle joins with others of the triceps muscle complex to
give strong extension of the elbow joint.

23. Anconeus externus.

This short fleshy triangular muscle bridging the elbow joint is, in a sense,
a counterpart of the anconeus internus (medial part of the caput medialis of
the triceps brachii).

OriciIn. This muscle arises from the posterior surface of the lateral epi-
condylar flare of the distal humerus by fleshy fibers which then course distally
to the lateral side of the olecranon.

INSERTION. Fleshy and tendinous fibers insert laterally on the olecranon
along the semilunar notch.

Action. This muscle probably assists in extension of the elbow.

24. Infraspinatus.

This flat muscle occupying the infraspinatus fossa of the scapula joins the
scapula to the humerus.

Oricin. Fleshy fibers arising from the infraspinatus fossa of the scapula
triangulate toward the upper end of the humerus. The inferior part of the mus-
cle shows a tendency to linear deviation from the remainder of the muscle mass
to be designated as the teres minor muscle. At the shoulder the two are joined
to form a heavy flat tendon forming part of the tendinous cuff of the shoulder
joint.

INSERTION. The tendon fibers insert into the lower posterior area of the
greater tuberosity adjacent to and just below the tendon of the supraspinatus.

AcTIon. The action is to abduct and externally rotate the humerus.

25. Teres minor.

This small muscle lying along the inferior scapular border is hardly distin-
guishable as separate from the infraspinatus muscle.

Oricin. The common origin from the infraspinatus fossa with the infra-
spinatus muscle and a common tendon with this muscle indicates how close the
association actually is.

INSERTION. Tendon fibers join with those of the infraspinatus muscle to
insert on the greater tuberosity of the humerus. The most distal fibers are those
of the teres minor.

AcTIon. The muscle contraction produces external rotation of the humerus
with some assistance in abduction as well.

26. Supraspinatus.

This large muscle is a strong, flat, fleshy muscle connecting the scapula to
the humerus over the top of the shoulder joint.

456 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

FicurE 37. Lateral view of the right upper arm with the forearm in midposition to
show origin in relationship of forearm extensor musculature. All muscles arising from the
lateral epicondylar area have been reflected at their origins except for the supinator (37).
Key: 29, brachialis; 30, brachioradialis; 31, extensor carpi radialis longus; 32, extensor
carpi radialis brevis; 33, extensor digitorum communis; 34, extensor digitorum lateralis;
35, extensor carpi ulnaris; 36, extensor pollicis et indicis longus; 37, supinator; 38, abductor
pollicis longus; 44, pronator teres; H, humerus, connective tissue overlying the bone; R,
radius; U, ulna.

Oricin. Muscle fibers arise from the entire supraspinatus fossa and con-
verge toward the shoulder joint where a strong flat tendon develops bridging this
joint as part of the tendinous cuff.

INSERTION. Tendon fibers enter the greater tuberosity of the humerus ad-

VoL. XXXIX] HOWARD: SEA OTTER FORELIMB 457

Ficure 38. Lateral view of the right upper extremity with the forearm in midposition.
All forearm extensor musculature has been removed to show origin of supinator muscle (37).
The interossei membrane is shown spanning between the radius and ulna. Key: 29, brachi-
alis; 30, brachioradialis; 37, supinator; 44, pronator teres; A, interossei membrane; H,
humerus, lateral epicondyle.

jacent and superior to the tendon of the infraspinatus and extend to the margin
of the bicipital groove.
AcTIoN. This muscle initiates abduction of the humerus.

27. Subscapularis.

This large triangular muscle lying on the underside of the scapula connects
this bone with the humerus.

OricIn. Arising as fleshy and tendinous fibers from the entire subscapular
surface, the muscle triangulates toward the glenoid where, beyond the scapular
spine, it parallels and approximates the supraspinatus muscle. The flat, heavy
tendon which then forms bridges the shoulder joint as part of the tendinous cuff
of this joint.

INSERTION. The tendon fibers insert into the full width of the lesser tuber-
osity of the humerus.

Action. This muscle is a strong internal rotator of the humerus.

458 CALIFORNIA ACADEMY OF SCIENCES [PRroc. 4TH SER.

5 cm

Ficure 39. An enlarged view of the radial side of the wrist of the right upper extremity.
Detail of insertion of the radial carpal extensors (31) and (32) and the abductor pollicis
longus (38) is shown. Note dorsal ligamentous structures which hold tendons in
position when wrist is dorsiflexed. Key: 30, brachioradialis; 31, extensor carpi radialis
longus; 32, extensor carpi radialis brevis; 33, extensor digitorum communis; 36, extensor
pollicis et indicis longus; 38, abductor pollicis; 43, flexor carpi radialis; A, metacarpal-1;
B, metacarpal-2; C, metacarpal-3; D, radial sesamoid; R, radius, styloid area.

Note. The tendons of the subscapularis, supraspinatus, and infraspinatus
(with the teres minor) are closely associated with the superior portion of the
shoulder joint capsule and form a strong tendinous cuff about the joint. The
articular surface of the humerus is thus held snuggly against the glenoid sur-
face of the scapula.

28. Biceps brachii.

This single-bellied, spindle-shaped muscle with a well developed tendon at
each end joins the scapula to the radius.

OrIcIN. From the tubercle on the neck of the scapula at the superior bor-
der of the glenoid a heavy tendon arises which passes through the shoulder joint
proper to enter the bicipital groove at the upper end of the humerus. From
here the tendon expands into a spindle-shaped muscle lying along the anterior
surface of the humerus to reach the anterior cubital fossa. The lower tendon
now forms and passes across the elbow joint to the proximal radius.

INSERTION. The distal tendon fibers insert into the bicipital tubercle of
the radius.

Action. This muscle has the combined action of flexion of the elbow and
supination of the forearm.

29. Brachialis.

This strong thick muscle underlies the brachioradialis muscle in the upper
one-half of the humerus, and connects the humerus with the ulna.

VoL. XXXIX] HOWARD: SEA OTTER FORELIMB 459

Ficure 40. An enlarged view of the dorsal and the distal right forearm and hand to
show relationship of extensor musculature of the wrist and digits. For digit 2, the extensor
aponeurosis overlying the proximal phalanx has been split longitudinally and reflected (A)
to show relationship to the sesamoid and proximal joint (metacarpal-phalangeal joint).
Also, note ligamentous attachment (B) of sesamoid to base of proximal phalanx, thus caus-
ing the sesamoid to move with the proximal phalanx on flexion and extension. Distally, the
extensor digitorum communis (33) and the index portion of the extensor pollicis et indicis
(36) of the extensor aponeurosis have been divided transversely to show insertion into base
of dorsum of the middle phalanx (C). For digit 4, the extensor aponeurosis and common
digital extensors have been split longitudinally (E) and spread apart to show relationship
to sesamoid. The insertion of the extensor carpi ulnaris (35) into the base of the fifth meta-
carpal and the abductor pollicis (38) into the base of the first metacarpal are shown. Also
note the transverse overlying ligaments at wrist level. Key: 30, brachioradialis; 31, extensor
carpi radialis longus; 32, extensor carpi radialis brevis; 33, extensor digitorum communis;
34, extensor digitorum lateralis; 35, extensor carpi ulnaris; 36, extensor pollicis et indicis
longus; 38, abductor pollicis; A, split extensor aponeurosis of digit 2; B, ligament attaching
sesamoid to proximal phalanx; C, insertion of long extensor tendon on digit 2; D, sesamoid
bone; E, split extensor tendon of digit 4; F, base of metacarpal 1; G, base of metacarpal 5.

460 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

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Ficure 41. Medial view of right upper extremity with forearm in midposition, thus
presenting its volar or flexor surface. This figure shows the origin and relationship of the
superficial forearm flexor musculature. The abductor digiti quinti has been displaced out-
ward from the shaft of the fifth metacarpal. Key: 13, pectoralis major, insertion; 29, brachi-
alis; 30, brachioradialis; 37, supinator; 40, palmaris longus; 41, flexor digitorum profundus
(ulnar head); 44, pronator teres; 49, abductor digiti quinti; A, elbow joint; B, pisiform
bone; H, humerus, medial epicondylar foramen.

Oricin. Muscle fibers arise from almost the entire posteromedial surface
of the humerus, from the neck of the humerus above to the lateral epicondylar
ridge below. Entering the area of the cubital fossa a tendon is formed which
bridges the elbow joint.

INSERTION. The tendon of the brachialis muscle inserts on the medial side
of the proximal ulna adjacent to the insertion of the pectoantebrachialis tendon.

Action. This muscle is a strong flexor of the elbow joint.

30. Brachioradialis.

This is a long elbow-spanning fleshy muscle connecting the proximal hu-
merus with the distal radius.

OricIn. Arising as thin flat tendinous fibers from the surface of the upper
part of the brachialis muscle and from along the deltoid ridge of the humerus
from the greater tuberosity to the humeral crest, a fleshy muscle belly promptly
develops which spans the elbow joint anterolaterally then tapers gradually fol-
lowing the radial border of the forearm to the area of the radial styloid.

INSERTION. Tendon fibers developing on the underside of the muscle distally
insert into the bone of the radial styloid.

VoL. XXXIX] HOWARD: SEA OTTER FORELIMB 461

Ficure 42. Volar view of right forearm in its distal portion and volar medial view of
proximal portion as the forearm is in a slightly pronated position. The palmaris longus
(40) and the flexor carpi ulnaris (39) have been divided and reflected to show origins and
relationships of the deeper flexor musculature. The musculature of the flexor digitorum
profundus can, by origin, be divided into three distinct portions, the more superficial humeral
head (42) and the ulnar head (41), and the deeper central head (45). Not visible in this
view, the central head has an ulnar, radial, and interosseus portion which will be shown
later. Distally, all of the profundi group are joined together at the wrist as a broad tendi-
nous structure from which separate the five profundi tendons, one passing to each digit.
At wrist level the tendon passes under the deep carpal ligament. Also distally, and in addition
to the main flexor tendon, the profundi give rise to the sublimi and lumbrical musculature
which will be described later. Key: 30, brachioradialis; 39, flexor carpi ulnaris; 40, pal-
maris longus; 41, flexor digitorum profundus, ulnar head; 42, flexor digitorum profundus,
humeral head; 43, flexor carpi radialis; 44, pronator teres; 49, abductor digiti quinti; U,
ulna, olecranon area.

Action. This strong muscle has the double action of flexing the elbow
and supinating the forearm.

31. Extensor carpi radialis longus.

This muscle is the more radial of the two radial carpal extensors.

OricIn. This muscle arises in common with the extensor carpi radialis
brevis by muscular fibers from the proximal three-fourths of the lateral epi-
condylar ridge. The common muscle extends about half-way down the forearm
before dividing into the longus and brevis portions. In the midforearm the
longus portion lies between the brachioradialis muscle and the extensor digi-
torum communis muscle. At wrist level, the tendon passes through a groove
on the dorsal surface of the radius adjacent to the styloid process. The over-
lying transverse dorsal carpal ligament forms a compartment for the tendon.

INSERTION. Bridging the wrist joint the tendon inserts at the base of meta-
carpal 2 into a tubercle on the dorsum and slightly to the radial side.

Action. This muscle is a strong dorsiflexor and radial flexor of the wrist
joint.

462 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Ficure 43. Volar view of right forearm with most of the flexor musculature reflected
to show the deeply situated flexor digitorum profundus (central head) (45), with its fibers
joining the ulna humeral divisions at near-wrist level. Also shown in this view are the in-
sertions of the biceps tendon (28), the pectoantebrachialis tendon (12), and the brachialis
muscle (29). The origin of the pronator teres (44) in relation to the palmaris longus (40)
is visualized. Key: 12, pectoantebrachialis; 28, biceps brachii; 29, brachialis; 30, brachio-
radialis; 39, flexor carpi ulnaris; 40, palmaris longus; 41, flexor digitorum profundus (ulnar
head) ; 42, flexor digitorum profundus (humeral head) ; 43, flexor carpi radialis; 44, prona-
tor teres; 45, flexor digitorum profundus (central head); H, humerus, medial epicondyle;
R, radius, styloid process; U, ulna.

32. Extensor carpi radialis brevis.

This muscle is a partner of the extensor carpi radialis longus and also con-
nects the humerus to the hand.

OricIN. In common with the extensor carpi radialis longus from the lateral
epicondylar ridge, this muscle represents the lower fibers of origin. The tendon
forming in midforearm parallels that of the longus and passes with this tendon
in the groove at the distal end of the radius.

INSERTION. Spanning the wrist joint the tendon inserts into the dorsum
at the base of metacarpal 3.

VoL. XXXIX] HOWARD: SEA OTTER FORELIMB 463

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Ficure 44. Volar view of right forearm. All flexor musculature has been reflected to
expose the radius and ulna with intervening interosseus membrane. The flexor digitorum
profundi musculature has been reflected distally where the tendons are entering the carpal
canal beneath the deep carpal ligament. Distally, the pronator quadratus muscle (46) is
shown spanning the two forearm bones. Note insertions of the biceps brachii (28),
brachialis (29), and pectoantebrachialis (12). Key: 12, pectoantebrachialis; 28, biceps bra-
chii; 29, brachialis; 30, brachioradialis; 39, flexor carpi ulnaris; 40, palmaris longus; 41,
flexor digitorum profundus (ulnar head); 42, flexor digitorum profundus (humeral head) ;
43, flexor carpi radialis; 44, pronator teres; 45, flexor digitorum profundus (central head) ;
46, pronator quadratus; A, elbow joint; B, deep carpal ligaments; C, interosseus membrane;
H, humerus, medial epicondyle; R, radius, head; U, ulna.

Action. Owing to its more central location and insertion, the brevis serves
only to dorsiflex the wrist.

33. Extensor digitorum communis.

This muscle, along with the extensor digitorum lateralis, represents the long
extensors of the digits except for the thumb. The musculature is in the forearm
and the tendons extend distally to the middle phalanges.

Oricin. The muscle arises in common with the extensor digitorum lateralis
from the lateral epicondylar ridge of the humerus down to and including the
lateral epicondyle. The extensor digitorum lateralis fibers are the more distal

464 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

in origin. At its origin, the muscle overlies the extensor carpi radialis brevis
and appears to share some fibers with this muscle. On the dorsum of the fore-
arm the muscle lies between the radial-carpal extensors and the extensor digi-
torum lateralis. At wrist level four separate tendons develop and pass through
a groove at the distal radius, following which they spread out and span to each
of the digits 2, 3, 4, and 5. Over the metacarpal area small tendinous slips may
develop which rejoin the same or adjacent tendon at the metacarpal-phalangeal
joint area.

INSERTION. At the metacarpal-phalangeal joint level each tendon expands
and joins into the extensor hood and the extensor aponeurosis mechanism, but
the central fibers continue down the dorsum of the proximal phalanx to insert
at the base of the middle phalanx. At the metacarpal-phalangeal joint level a
light fibrous attachment is noted to the sesamoid at this site. The sesamoid,
however, is more associated with the joint capsule than with the extensor tendon.

ActTIOoN. The primary action of the tendon is to extend the metacarpal-
phalangeal joint through the hood mechanism. However, with the metacarpal-
phalangeal joint stabilized so that hyperextension does not occur, the central
fibers then act to extend the proximal interphalangeal joint.

34. Extensor digitorum lateralis.

This partner of the extensor digitorum communis muscle provides an extra
long extensor tendon for the fourth and fifth digits.

Oricin. The muscle arises in common with the extensor digitorum com-
munis representing the more distal portion of this origin, the lateral humeral
epicondyle and epicondylar ridge. Separation from the communis occurs on
the ulnar side of this muscle and near the wrist two individual tendons develop
which pass through a separate compartment in a groove of the distal radius.
From here, the more radial of the two tendons passes beneath the communis
tendon of digit 5 to the midmetacarpal area to join with the communis tendon
of digit 4 at the extensor hood level. The more ulnar tendon parallels the com-
mon extensor tendon of digit 5 to join into the extensor hood of this digit on
the ulnar side.

INSERTION. Joining with the communis extensor tendon the insertion is
the same; namely, the base of the middle phalanx of digits 4 and 5.

AcTION. The action is the same as that of the communis tendon for the
fourth and fifth digits.

35. Extensor carpi ulnaris.

This strong muscle joins the humerus with the hand and serves to balance
the action of the radial carpal extensors.
OricIn. This muscle arises from the lateral epicondyle just radial to the

VoL. XXXIX] HOWARD: SEA OTTER FORELIMB 465

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Ficure 45. An enlarged view of the volar surface of the right forepaw. The volar pad
has been removed and the palmaris longus tendon (40) divided. Distally, the palmaris
longus fascial expansion (palmar fascia) has been reflected radialward to disclose the origin
of the sublimis muscle (48) for digit 5. Note the continuation of deep muscle fibers from
the underside of the palmaris longus joining with muscle fibers arising from the deep trans-
verse carpal ligament to form the sublimis muscle of this fifth digit. Key: 30, brachioradialis ;
39, flexor carpi ulnaris; 40, palmaris longus; 41, flexor digitorum profundus, tendons from
the ulnar head; 42, flexor digitorum profundus (humeral head); 43, flexor carpi radialis;
48, flexor digitorum sublimis for digit 5; 49, abductor digiti quinti; 50, opponens digiti
quinti; A, transverse carpal ligament; B, pisiform bone.

466 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

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Ficure 46. Enlarged view of volar surface of the right hand. The palmar fascia has
been completely removed except for the groups of distal fibers which reinforce the sublimi
tendons. The flexor tendon sheath has been resected a bit distally for digits 2, 3, 4, and 5
to show the broadening of the sublimis to completely cover the profundus tendon at this
site. Note again, there is no sublimis tendon for digit 1. The palmaris longus fibers for this
digit attach to the rim of the flexor tendon sheath. The transverse carpal ligament (A) has
been divided and reflected to show the origin of the sublimis of digits 2, 3, and 4. The sub-
limis of digit 5 has been reflected from its origin and displaced ulnaward. Also shown are
the profundi flexor tendons of all digits and how these tendons arise from the proximal
musculature. Key: 30, brachioradialis; 39, flexor carpi ulnaris; 41, flexor digitorum pro-
fundus (ulnar head); 42, flexor digitorum profundus (humeral head); 43, flexor carpi ra-
dialis; 47, flexor digitorum sublimis (radial) ; 48, flexor digitorum sublimis (ulnar); 49, ab-
ductor digiti quinti; 50, opponens digiti quinti; A, transverse carpal ligament; B, fibrous
flexor tendon sheath.

VoL. XXXIX] HOWARD: SEA OTTER FORELIMB 467

Ficure 47. Enlarged view of the volar aspect of the right hand. On digit 4 the fibrous
flexor tendon sheath (A) has been divided along the ulnar margin and reflected radialward
to show the contents of the flexor tunnel. The concentration of transverse fibers on the
flexor tendon sheath provides the pulley mechanism of the sheath, thus preventing bow-
stringing of the flexor tendons when activated. Note the division and rejoining of the sub-
limis tendon to allow the profundus to come through. The sublimis then continues distally
to insert in the base of the middle phalanx (B). The profundus flexor tendon continues in
this sheath to the distal interphalangeal-joint level, inserting at the base of the terminal
phalanx which is in a hyperextended position at rest. The deep transverse carpal ligament
(D) has been divided and reflected. This broad ligament has its radial attachment mainly
to the radial sesamoid and volar wrist joint capsule along the radial side of the navicular-
lunate bone. On the ulnar side, attachment is mainly along the base of the pisiform with
attachment also to the wrist joint capsule in the area of the triquetrum. Key: 41, flexor
digitorum profundus (ulnar head); 42, flexor digitorum profundus (humeral head); 47,
flexor digiti sublimis (radial) ; 48, flexor digiti sublimis (ulnar of the fifth digit) ; 49, abduc-
tor digiti quinti; A, transverse carpal ligament; B, fibrous tendon sheath of digit 4, reflected;
C, insertion of sublimis flexor tendon; D, insertion of profundus flexor tendon.

468 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

5 em

Ficure 48. Enlarged ulnar view of the middle and distal phalanges of digit 4 to show
the hyperextended position of the terminal phalanx and the insertion of the profundus
flexor tendon. Note the close approximation of the claw to the terminal phalanx. Key: A,
middle phalanx; B, insertion of profundus flexor tendon on distal phalanx; C, claw.

origin of the supinator muscle adjacent and distal to the extensor digitorum
lateralis muscle. The muscle passes along the ulnar border of the dorsum of
the forearm adjacent to the extensor digitorum lateralis. Near the wrist a
strong tendon is formed which broadens and passes over the distal ulna with an
interposed bursa between the tendon and the bone.

INSERTION. Spanning the carpal area, the tendon continues distally to the
ulnar side of the base of the fifth metacarpal where it inserts into a slight bony
protuberance at this site.

Action. The direct action of this muscle is to ulnaflex the wrist but, if
this motion is opposed, the action becomes one of dorsiflexion of the wrist joint.

36. Extensor pollicis et indicis.

This muscle provides long extensor action for the thumb and an assistance
to the communis extensor tendon of digit 2.

OricInN. The muscle arises entirely from the proximal two-thirds of the ulna,
beginning at the articular notch level and extending distally along the ulnar
crest. The origin is just posterior to that of the abductor pollicis longus muscle.
The muscle underlies the extensor digitorum communis muscle and the tendon
which forms passes through the same compartment at the distal radius as the
communis tendons. Just distal to the wrist, the tendon divides into two separate
tendons which continue radiusward. The more radial one goes directly to the
base of the thumb, the other underlies the communis tendon to reach the ulnar
side of digit 2.

INSERTION. The pollicis tendon inserts into the base of the proximal phalanx
of digit 1 after spreading out somewhat in a hoodlike manner at the metacarpal-
phalangeal joint level. The indicis tendon, which is approximately twice the
size of the communis tendon of this digit, joins into the extensor hood on the

VoL. XXXIX] HOWARD: SEA OTTER FORELIMB 469

Ficure 49. Enlarged ulnalateral view of ulna border of the right hand to show relation-
ship of muscles of tendons about the fifth metacarpal (H). Note insertion of the extensor
digiti lateralis (34) is mainly into the base of the proximal phalanx, the communis extensor,
forming the main extensor aponeurosis. Although in close proximity the base of the fifth
metacarpal does not articulate with the joint surface on the triquetrum but rather more
radial on the hamate. The pisiform bone has been reflected outward a bit for a cleared view
of the ulna carpal extensor and flexor tendons. Key: 33, extensor digitorum communis;
34, extensor digitorum lateralis; 35, extensor carpi ulnaris; 39, flexor carpi ulnaris; 49, ab-
’ ductor digiti quinti; 50, opponens digiti quinti; 51, flexor digiti quinti brevis; A, proximal
phalanx; B, insertion of extensor digitorum lateralis; C, sesamoid bone; D, collateral ligament
at the metacarpal phalangeal joint; E, extensor hood at the metacarpal phalangeal joint;
F, pisiform; G, triquetrum; H, metacarpal (fifth); U, ulna, distal end.

ulnar side of the communis tendon, then joins with the communis tendon for
the remainder of its course.

Action. The pollicis portion extends the metacarpal-phalangeal joint of
the thumb and acts as an adductor of the thumb as well. The indicis portion has
the identical action to that of the comunis tendon of digit 2.

37. Supinator.

This deep, somewhat flat forearm muscle joins the humerus to the radius.

OricIN. Heavy tendinous fibers arise from the small area on the distal
and most prominent part of the lateral humeral epicondyle just ulnaward of
the origin of the extensor carpi ulnaris muscle. From its origin the muscle passes
directly toward the radius and promptly begins its insertion.

INSERTION. Insertion is into the proximal three-fourths of the radius start-

470 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Ficure 50. Palmar view of the right hand to show specifically the origin and insertion
of the single lumbricale muscle in this specimen (53). In this plate, the entire flexor sheath
has been removed from digit 4 and the proximal one-half removed from digit 5. For digit 1,
the palmar fascia attached to the rim of the flexor sheath has been split and reflected (40).
Key: 40, palmaris longus; 41, flexor digiti profundus (ulnar head); 42, flexor digiti pro-
fundus (humeral head) ; 47, flexor digitorum sublimis (radial) ; 48, flexor digitorum sublimis
(ulnar) ; A, digit 5.

Vor, XXXIX] HOWARD: SEA OTTER FORELIMB 471

5am

Ficure 51. Enlarged palmar view of right hand. The long digital flexor tendons have
been reflected distally to show the intrinsic hand muscles grouped about the first and fifth
metacarpals. These muscles are termed respectively the thenar and hypothenar muscle
groups. The opponens digiti quinti (50) has been detached from its origin and reflected
ulnaward. Note absence of an opponens pollicis muscle. The flexor pollicis brevis of digit
1 (55) has two heads designated as radial and ulnar by their position. A heavy fibrous
capsule at the wrist joint forms the floor of the carpal tunnel. Key: 39, flexor carpi ulnaris;
41, flexor digitorum profundus (ulnar head) ; 42, flexor digitorum profundus (humeral head) ;
49, abductor digiti quinti; 50, opponens digiti quinti; 51, flexor digiti quinti brevis; 52, ad-
ductor digiti quinti; 53, lumbricali; 54, abductor pollicis brevis; 55, flexor pollicis brevis; 56,
adductor pollicis; 59, interossei; A, transverse carpal ligament—reflected; B, pisiform bone;
C, radial sesamoid bone; D, volar capsule of wrist joint; E, proximal phalanx of fifth digit.

472 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Ficure 52. An enlarged radial view of the right first digit to show the relationship of the
extensor, extrinsic, and intrinsic muscles. Both pollicis and indicis portions of the extensor
pollicis et indicis tendons are shown. The indicis part lies behind the pollicis tendon. Note
the more distant extension of the radial fibers of the pollicis tendon at the insertion at the
base of the proximal phalanx. Note also the location of elastic tissue (dorsal elastic liga-
ment) which keeps the distal phalanx hyperextended. On voluntary flexion of the distal
phalanx, the elastic tissue simply stretches. There is no extensor tendon as such inserting
on the distal phalanx at digit 1 nor on the distal phalanges of the remaining digits. Visible
in this view is the radial head of the flexor pollicis brevis (55). Key: 36, extensor pollicis
et indicis; 38, abductor pollicis longus; 42, flexor digitorum profundus (humeral head) ; 54,
abductor pollicis brevis; 55, flexor pollicis brevis (radial head); A, carpus (multangular) ;
B, proximal phalanx; C, elastic tissue (dorsal elastic ligament) ; D, sesamoid; R, radius, dis-
tal end.

ing on the posterior and lateral aspects at the bicipital tubercle level and termi-
nating in tendinous fibers at the point of maximum bow of the radius. Thus
strong mechanical advantage is provided for its action.

AcTIon. This muscle is a strong supinator of the forearm.

38. Abductor pollicis longus.

This deep extensor muscle joins the ulna to the thumb.

OricIn. This muscle arises from the lateral aspect of the ulna all the way
from the articular notch to the end of the bone. The origin is adjacent to and
on the radial side of the extensor pollicis et indicis origin. Some fleshy fibers
of origin also arise from the proximal interosseus membrane and adjacent op-
posing surface of the radius. The muscle fibers converge in a triangular con-
figuration to emerge from under the extensor digitorum communis in the distal
forearm, where they then pass over the radial carpal extensors toward the radial
styloid. The tendon now formed passes in the groove of the radius at the radial
styloid level.

VoL. XXXIX] HOWARD: SEA OTTER FORELIMB 473

Ficure 53. Enlarged view of the ulnar side of the right first digit to show the extrinisic
and intrinsic muscles on this side. The principal insertion of the extensor pollicis (36) is
into bone somewhat to the ulnar side of the midline space of the proximal phalanx. The radial
fibers, as noted in figure 52, insert somewhat more distally. Visible in this view is the ulnar
head of the flexor pollicis brevis (55). Key: 36, extensor pollicis et indicis (pollicis part) ;
42, flexor digitorum profundus (humeral head); 55, flexor pollicis brevis (ulnar head); 56,
adductor pollicis; A, metacarpal of digit 1; B, proximal phalanx of digit 1; C, elastic tissue
(dorsal elastic ligament).

INSERTION. Emerging from the distal compartment of the radius the tendon
splits into a lesser one-fourth which inserts volarly into the radial sesamoid,
and a greater three-fourths which inserts into a bony tubercle to the radial side
of the dorsum of the base of the first metacarpal.

AcTIon. By virtue of its insertion this muscle acts to abduct the thumb
metacarpal, and also to radiflex the wrist to some degree. Like other extensor
musculature it can assist in supination of the forearm.

39. Flexor carpi ulnaris.

This large strong muscle occupies the ulnar side of the volar surface of the
forearm and lies adjacent to the palmaris longus muscle. It extends from the el-
bow area to the hand.

Oricin. Arising by muscular tendinous fibers from the medial surface of
the lower part of the olecranon crest the origin just overlies the origin of the
ulnar head of the flexor profundus digitorum muscles with a few fibers seem-
ingly in common. Other but lesser fleshy fibers arise from the adjacent medial
aspéct of the olecranon and over the joint capsule in this area, and a few from
the humerus along the articular ridge of the medial condyle. The muscle is
elongated, fusiform in type, and in its course down the forearm it overlies all
but the proximal part of the flexor digitorum profundus (ulnar head). The

474 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

ye

Syeelad

Ficure 54. An enlarged radiolateral view of digit 5 to show insertion of extensor digi-
torum communis (33) at the base of the middle phalanx. The elastic tissue at distal joint
level has been reflected from its attachment to the middle phalanx, thus allowing the distal
joint to assume a more flexed position. Insertions of the intrinsic musculature on the radial
side of digit 5 are shown. Key: 33, extensor digitorum communis; 41, flexor digitorum
profundus (ulnar head); 50, opponens digiti quinti; 51, flexor digiti quinti brevis; 52, ad-
ductor digiti quinti; A, metacarpal of digit 5; B, proximal phalanx; C, elastic tissue (dorsal
elastic ligament).

tendon of insertion begins to appear about halfway down the muscle. The mus-
cle fibers continue as far distally as the pisiform bone.

INSERTION. At the carpal level the tendon inserts into and encompasses
the pisiform bone but continues to pass deeply into the palm to its final inser-
tion at the volar surface of the base of the fifth metacarpal bone.

Action. This muscle is a strong volar flexor of the wrist and gives ulna-
flexion of the wrist as well.

40. Palmaris longus.

This is the most prominent muscle on the volar surface of the forearm. It
is a large, broad, flat muscle which lies superficially adjacent to the flexor carpi
ulnaris and connects the humerus with the hand. A tendinous surface presents
on the distal two-thirds of the muscle.

OricIn. Arising as the outermost muscle from the distal three-fourths of

VoL. XXXIX] HOWARD: SEA OTTER FORELIMB 475

Siem

Ficure 55. An enlarged palmar (volar) view of the right hand with all muscles and ten-

dons removed except for the abductor (57) and adductor (58) digiti secundus and the three

interossei (59). The interossei constitute the deepest layer of muscles, as the abductor and

adductor digiti secundi lie a bit superficial to them. Key: 57, abductor digiti secundi; 58,

adductor digiti secundi; 59, interossei; A, metacarpal, digit 5; B, head of fourth metacarpal;
C, pisiform bone; D, transverse carpal ligament—cut and reflected.

the medial epicondyle, the origin overlies the distal one-half of the origin of the
pronator teres muscle. Tendinous fibers appear the width of the muscle at the
junction of the proximal and middle thirds, and continue distally as a broad
tendinous sheath to the wrist. The superficial fibers spread out over the palm
forming a dense palmar fascia. Connective tissue fibers from this fascia extend

476 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Ficure 56. A radiolateral view of digit 4 of the right hand to show the pattern of in-
sertion of all interossei muscles except the one to the radial side of digit 2. Note how the
flat interosseus tendon joins the extensor aponeurosis at the extensor hood level overlying
the metacarpal phalangeal joint. The anatomical relationship of the long extensor probably
allows the interossei to extend the proximal interphalangeal joint when the metacarpal
phalangeal joint is in hyperextension. It appears that the hood mechanism probably checks
the excursion of the long extensor thus limiting its action on the proximal interphalangeal
joint when in the hyperextended position. Key: 33, extensor digitorum communis; 44, flexor
digitorum profundus; 47, flexor digitorum sublimis; 59, interosseus; A, extensor hood; B,
extensor aponeurosis; C, elastic tissue (dorsal elastic ligament); D, metacarpal of digit 4.

into the pads of the palm and the digits. The dense palmar fascia shows three
major fibrous bands extending toward the bases of digits 2, 3, and 4, with lesser
concentrations of fibers passing toward the bases of digits 1 and 5. Fibrous
extensions of the palmar fascia also pass deeply into the palm forming septum-
like structures which attach to the fibrous flexor tendon sheaths on either side
of the metacarpal heads. The interval between the metacarpal heads becomes
a hiatus through which the volar digital nerves and vessels pass into the digits.
When the palmaris longus muscle is divided just above the wrist and the distal
portion reflected, the undersurface can be observed. Here it is noted that toward
the ulnar side at wrist level muscle fibers continue and separate from the main
more superficial tendinous portion. These muscle fibers are now joined by ad-
ditional muscle fibers arising from the deep transverse carpal ligament. This
combined muscle shortly terminates in a small discreet tendon which then
promptly splits into two halves which enter the fibrous flexor tendon sheath
of digit 5 as a sublimis tendon. At its point of entry, this small tendon is rein-
forced by main fibers from the palmar fascia. The palmar fascia proper, which
spreads over the palm with the concentration of fibers and bands towards the
digits 2, 3, and 4, shows these bands attaching to the fibrous flexor sheath and

VoL. XXXIX] HOWARD: SEA OTTER FORELIMB 477

\
= a
. CaaS
J = PIE, A
a) ee e os ? Me
“ deren seen

Hiem

Ficure 57. Skull, right lateral view. Key: 3, clavotrapezius (origin); 8, rhomboideus
capitis (origin); A, lamboidal crest; B, mastoid process.

entering the sheath to reinforce the respective sublimi tendons to these digits.
For the thumb, the lesser fibrous band simply attaches to the fibrous flexor
digital sheath as there is no sublimis tendon for this digit.

INSERTION. As noted from the description above, the palmaris longus mus-
cle has three functions; namely, cupping of the palm by tensing the palmar fas-
cia, volar flexion of the wrist, and reinforcing the flexor sublimi tendons which
flex the proximal interphalangeal joints of digits 2, 3, 4, and 5.

41. Flexor digitorum profundus (ulnar head).

This muscle represents part of the profundus flexor mechanism of the digits.
The remaining part is the flexor digitorum profundus, humeral head and central
head.

OricIn. Musculotendinous fibers arise from the medial olecranon ridge
beneath the origin of the flexor carpi ulnaris with additional fleshy fibers of
origin continuing distally from the medial surface of the ulna in its upper half
tapering along the ulna crest. Tendon fibers begin to appear in the lower half
of the muscle but muscle fibers continue as far distally as the wrist. Slightly
above wrist level, junction occurs with the humeral and central heads with five
resulting strong tendons which pass through the carpal tunnel under the deep
transverse carpal ligament and then pass one to each digit. It is to be noted that
the contribution from the ulnar head makes up the tendons for digits 4 and 5.

478 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

: 5lcimn

Ficure 58. Skull, volar view. Key: 4, cleidomastoideus (origin) ; A, mastoid process.

INSERTION. Flexor profundus tendons of digits 4 and 5 enter the fibrous
flexor tendon sheath of the metacarpal-head level passing through a split of the
flexor sublimis tendon in the proximal phalanx area, the tendon continues dis-
tally in its sheath to its insertion on the volar surface of the base of the termi-
nal or distal phalanx.

Action. The profundus tendon flexes the terminal finger joint and all proxi-
mal finger joints as well.

42. Flexor digitorum profundus (humeral head).

This is a strong centrally placed forearm muscle connecting the humerus
with the digits. It is part of the profundus flexor mechanism.

OriciIn. Tendinous and muscular fibers arise from the medial humeral
epicondyle under the distal one-half of the origin of the palmaris longus muscle
and just distal to the origin of the flexor carpi radialis muscle. In the forearm,

VoL. XXXIX] HOWARD: SEA OTTER FORELIMB 479

Seen

Ficure 59. Atlas, volar view. Key: 6, omotrachleon (origin); 9, rhomboideus pro-
fundus (origin); A, transverse process; B, anterior tubercle.

the muscle lies between the flexor digitorum profundus (ulnar head) and the
flexor carpi radialis. Insertional tendon begins to develop about halfway to
the wrist and, at wrist level, junction with the ulnar and central heads occurs
and the five resulting tendons pass through the carpal tunnel and on to the
digits. The humeral head contributes to the tendons of digits 1, 2, and 3.

INSERTION. The profundus tendons of digits 2 and 3 insert as described
for those of digits 4 and 5. For the thumb, a similar fibrous flexor tendon
sheath is present but there is no sublimis tendon and only two phalanges. In-
sertion is into the base of the terminal phalanx on its volar surface.

AcTION. The profundus tendon flexes the distal joint of the digits and all
proximal digital joints as well.

43. Flexor carpi radialis.

This strong muscle with tendinous fibers at each end connects the humerus
to the hand.

OrIGIN. The heavy tendinous fibers of origin arise from the medial humeral
epicondyle under the proximal one-half of the origin of the palmaris longus mus-
cle and adjacent to the origin of the humeral head of the flexor digitorum pro-
fundus muscle. The flexor carpi radialis is fusiform in shape and passes down
the volar aspect of the forearm under the palmaris longus muscle and adjacent
to the humeral head of the flexor digitorum profundus. At wrist level, a heavy

480 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

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Ficure 60. Right scapula, lateral surface. Key: 16, teres major (origin); 17, acro-
miodeltoideus (origin); 18, spinodeltoideus (origin); 19, dorsoepitrochlearis (origin); 24,
infraspinatus (origin); 26, supraspinatus (origin); 28, biceps brachii (origin); 1, spino-
trapezius (insertion); 2, acromiotrapezius (insertion); 6, omotrachleon (insertion); 7,
rhomboideus major (insertion); 8, rhomboideus capitis (insertion); 9, rhomboideus pro-
fundus (insertion) ; A, axillary border of scapula; B, vertebral border of scapula; C, supe-
rior border of scapula; D, superior angle of scapula; E, inferior angle of scapula; F, spine
of scapula; G, tuberosity of scapular spine; H, glenoid surface; I, acromion process.

tendon forms which ‘passes to the ulnar side of the radial sesamoid and dips
deeply into the palm.
INSERTION. This tendon inserts on the volar surface of the base of meta-

carpal 2.
ActTIon. Strong palmar flexion of the wrist results from a pull on this ten-

don.

44. Pronator teres.

This is a strong thick muscle passing from the medial side of the distal hu-
merus to the radius.

VoL. XXXIX] HOWARD: SEA OTTER FORELIMB 481

Bem

Ficure 61. Right scapula, medial surface. Key: 22, triceps brachii caput medialis
(origin); 27, subscapularis (origin); 28, biceps brachii (origin); 10, serratus magnus
(insertion) ; 11, levator anguli scapulae (insertion); A, glenoid; B, supraglenoid tubercle.

OrIcIN. This muscle is the most proximal of those arising from the medial
humeral epicondyle, but the distal one-half of the origin underlies the origin
of the flexor carpi radialis. Arising mostly by tendinous fibers, this muscle
diagonally crosses the upper forearm and spreads out as it approaches the ra-
dius.

INSERTION. Starting along the radial margin of the ridge of the radius,
the insertion continues distally the full length of the bone. Tendinous fibers
make up the insertion for the most part, with a heavy concentration of fibers
at the maximum bow of the radius where the insertion comes in close relation-
ship to the insertion of the supinator muscle. Tendinous fibers continue in lesser
concentration along the distal radius to the volar aspect of the radial styloid
where they lie adjacent to the insertion of the brachioradialis muscle.

Action. As an antagonist to supination, this muscle is a strong pronator
of the forearm.

45. Flexor digitorum profundus (central head).

The deepest of the flexor profundus group, this muscle is roughly Y-shaped
and arises in the upper forearm to join with the ulnar and humeral heads of the
flexor digitorum profundus to give flexor action to the digits.

OrIGIN. There are three areas of origin of this muscle. The ulnar portion
arises as fleshy fibers from the anteromedial surface of the upper two-thirds

482 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

C

; Cin =
Ficure 62. Right scapula, glenoid view. Key: 28, biceps brachii (origin); A, glenoid;
B, superior angle of scapula; C, inferior angle of scapula; D, acromion process.

of the ulna. The origin starts near the articular lip of the ulna and adjacent and
lateral to the tendons of insertion of the brachialis and pectoantebrachialis
muscles. The radial portion arises from the anteromedial surface of the upper
two-thirds of the radius, starting near the articular rim at the neck of the radius
and just lateral to the bicipital tubercle. The interosseus portion arises from
the volar surface of the interosseus membrane between the radius and the ulna,
at the upper margin of the membrane. The muscle thus formed is flat and under-
lies the other profundi musculature which it joins from the underside at the
level where the tendons begin to form. Distally, the flexor profundus muscula-
ture overlies the pronator quadratus muscle.

INSERTION. The muscle fibers join into the flexor digitorum profundus
(central and ulnar heads) which contributes to the tendons formed and there-
fore has a common insertion with them.

VoL. XXXTIX] HOWARD: SEA OTTER FORELIMB 483

Ficure 63. Right humerus, posterior view. Key: 21, triceps brachii caput lateralis
(origin); 29, brachialis (origin); 30, brachioradialis (origin); 31, extensor carpi radi-
alis longus (origin); 32, extensor carpi radialis brevis (origin); 33, extensor digitorum
communis (origin); 34, extensor digitorum lateralis (origin); 35, extensor carpi ulnaris
(origin) ; 37, supinator (origin); 5, clavobrachialis (insertion) ; 13, pectoralis major (inser-
tion); 17, acromiodeltoideus (insertion); 18, spinodeltoideus (insertion); 24, infraspinatus
(insertion) ; 26, supraspinatus (insertion); A, head of humerus; B, greater tuberosity; C,
pectoral ridge; D, deltoid ridge; E, lateral epicondyle; F, lateral epicondylar ridge.

484 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

46. Pronator quadratus.

This muscle is flat and thin with fibers running transversely. It lies in the
distal forearm connecting the radius and ulna.

OricIn. Fleshy and tendinous fibers arise from the distal one-third of the
medial surface of the ulna. From its origin, the muscle fibers pass directly
transversely to the radius maintaining full width of the muscle.

INSERTION. Fleshy and tendinous fibers insert into the distal one-fourth
of the volar surface of the radius adjacent to the lower part of the insertion of
the pronator teres muscle.

AcTION. Pronation of the forearm is accomplished by this muscle.

47 and 48. Flexor digitorum sublimis.

There are four small flexor sublimi tendons, one each for digits 2, 3, 4, and
5. Digit 1 is without a sublimis tendon. The flexor sublimis of digit 5 has been
previously described under the palmaris longus muscle.

OricIn. The flexor sublimis tendons for digits 2, 3, and 4, arise from the
volar surface of the flexor digitorum profundus complex just proximal to the
wrist. The sublimis for digit 4 comes from the ulnar head, those for digits 2
and 3 from the humeral head. Because of their origin and position in the hand,
they have been grouped as (47) flexor digitorum sublimis (radial) for digits
2 and 3, and (48) flexor digitorum sublimis (ulnaris) for digits 4 and 5.

INSERTION. The sublimis tendons of digits 2, 3, 4, and 5 enter the digital
fibrous flexor tendon sheaths superficially to the profundus flexor tendons. At
this point, each is strongly reinforced by palmar fascia fibers to form a wide
flat tendon overlying and completely covering the profundus tendon. After a
short distance within the sheaths, the sublimis tendons abruptly split longi-
tudinally and curve around on either side of the profundus to rejoin into a thin
but broad tendon which hugs the proximal phalanx forming the back wall of the
flexor sheaths. The sublimi tendons continue distally across the proximal inter-
phalangeal joint to a broad insertion on the volar lip of the proximal end of
the middle phalanx.

Action. The sublimi tendons act to flex the proximal interphalangeal joints
of digits 2, 3, 4, and 5. In view of their origin, those of digits 2, 3, and 4 must
function in relation to the profundus flexor tendons. For digit 5, the anatomy
would allow for some independent flexion of the proximal interphalangeal joint.
The broad spread of the sublimis tendons at metacarpal-phalangeal joint
level could also serve as a pulley mechanism for the profundi tendons.

THE INTRINSIC MUSCEES OF THE HAND

A group of small muscles within the hand proper have been termed intrinsic
hand muscles. In view of their size, these muscles are not strong but they serve

VoL. XXXIX] HOWARD: SEA OTTER FORELIMB 485

a most important function in balancing the action of the long forearm flexor
and extensor musculature. These muscles, also, can provide the fine skill motion
of the digits. This latter function can hardly be recognized in the sea otter as
the digits are in the state of syndactyly and the thumb is unopposable.

For descriptive purposes, these intrinsic muscles can be grouped as fol-
lows. Those about the first ray are called the thenar intrinsic muscles. Those
about the fifth ray are called the hypothenar intrinsic muscles. The remaining
small muscles consist of the interossei, one each for the second, third, and fourth
digits, and two accessory intrinsic muscles for digit 2.

Also, the lumbrical muscle is generally considered an intrinsic muscle, al-
though its origin is from the flexor digitorum profundus group of tendons.

THE HYyPpoTHENAR GROUP OF INTRINSIC HAND MUSCLES

49. Abductor digiti quinti.

This is a rather large coarse muscle forming the ulnar border of the hand.

OricIn. Fleshy fibers arise from a triangular area on the distal surface of
the pisiform bone. The muscle becomes spindle-shaped passing distally.

INSERTION. The insertion is through a sesamoid bone into the ulnar volar
aspect of the base of the proximal phalanx.

Action. This muscle flexes the metacarpal phalangeal joint of digit 5 and
would also serve to abduct the extended digit within its anatomical limits.

50. Opponens digiti quinti.

This is a superficial slender fusiform muscle of the hypothenar group.

OricIn. Fleshy and tendinous fibers arise from the ulnar side of the deep
carpal ligament to form the thin muscle which first overrides the long flexors
of the fifth digit, then passes more deeply into the palm on a slightly oblique
course to overlie the adductor and flexor digiti quinti. At the metacarpal phalan-
geal joint level a small thin tendon forms.

INSERTION. The insertion is into the volar aspect of the base of the proximal
phalanx of digit 5 just to the ulnar side of the tendinous insertion of the adduc-
tor digiti quinti.

Action. The position of this muscle would provide slight opposition action
to the fifth ray if such were possible. The muscle probably serves to assist in
independent flexion of the metacarpal-phalangeal joint.

51. Flexor digiti quinti brevis.

This muscle is small and deep in the palm, being in the general plane of the
interossei muscles of the adjacent digits.

OricIn. This muscle arises from the fibrous capsule of the wrist joint to
the ulnar side of the flexor carpi ulnaris tendon. A fusiform muscle is formed
which passes directly to the radial side of digit 5 where a small tendon forms.

486 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

INSERTION. This small tendon inserts into the base of the proximal phalanx
and on the radial side dorsal to the insertion of the adductor and opponens mus-
cles.

AcTIoN. Flexion of the metacarpal-phalangeal joint of the fifth digit is
accomplished by this muscle and also some degree of adduction.

52. Adductor digiti quinti.

This flat triangular muscle is the next most superficial of the hypothenar
group.

OricIn. Fleshy fibers arise over a broad area of the volar fibrous wrist-
joint capsule adjacent to and on the ulnar side of the origin of the adductor
pollicis and extend ulnaward to the margin of the flexor carpi ulnaris tendon.
The muscle from its origin triangulates distally and obliquely toward the base
of the proximal phalanx of the fifth digit.

INSERTION. A short tendon arises which inserts into the base of the proxi-
mal phalanx on the ulnar side just radial to the opponens digiti quinti and over-
lying the insertion of the flexor digiti quinti brevis.

AcTIoNn. The action of this muscle is to adduct the fifth digit and also to
assist in flexion of the metacarpal-phalangeal joint.

THE LUMBRICAL MUSCLE

53. Lumbricali.

This small intrinsic hand muscle differs from the other by virtue of its origin.

OriciIn. Arising from the opposed surfaces of the profundus flexor tendons
of digits 3 and 4 in the proximal palm, the fleshy fibers form a short slim muscle
which traverses the palm partly by a very thin tendon which passes to the radial
side of digit 4.

INSERTION. This small tendon inserts into the radial side of the proximal
phalanx of digit 4 just distal to the joint surface. The tendon fibers terminate
into the periosteum and bone rather than entering into the extensor aponeuro-
sis.

AcTION. Owing to its site of insertion, the muscle gives radiolateral motion
of the metacarpal-phalangeal joint and also flexion of this joint.

Notes. The number of lumbrical muscles is variable. In another specimen
three lumbrical muscles were noted.

THe THENAR INTRINSIC MUSCLES

54. Abductor pollicis brevis.

This is a small fusiform muscle on the radial side of digit 1.
Oricin. Fleshy fibers arise, half from the radial sesamoid and half from

VoL. XXXIX] HOWARD: SEA OTTER FORELIMB 487

the adjacent deep carpal ligament to form a small muscle which passes along
the radial side of the metacarpal adjacent to the radial portion of the flexor
pollicis brevis.

INSERTION. This small muscle terminates distally in a tendon which joins
with the tendon of the radial part of the flexor pollicis brevis to insert in the
radial volar side of the base of the proximal phalanx through a sesamoid bone.

Action. The principle action is flexion of the metacarpal-phalangeal joint
of digit 1. A minor degree of abduction is probably also obtained.

55. Flexor pollicis brevis.

This muscle, which divides near its insertion, is the deepest of the thenar
group.

OrIGIN. Fleshy fibers arise from the volar fibrous wrist joint capsule between
the origin of the adductor pollicis and the abductor pollicis brevis. The muscle
then passes distally overlying the proximal area of the metacarpal of digit 1.
The muscle then divides into radial and ulnar halves. The terminal tendon of
each half inserts into the base of the proximal phalanx, the radial half through
the sesamoid with a tendon of the short abductor, the ulnar half with the tendon
of the adductor pollicis.

AcTIon. Both halves serve as flexors of the metacarpal-phalangeal joint
of digit 1.

56. Adductor pollicis.

This muscle is the largest of the thenar group.

OriIGIN. Fleshy fibers arise from the volar fibrous capsule of the wrist joint
between the origin of the flexor pollicis brevis on the radial side and the adduc-
tor digiti quinti on the ulnar side. The origin of this muscle overlies the origin
of the deeper intrinsic muscles of the second digit. The muscle thus formed
courses obliquely radiusward across the palm to the ulnar side of the proximal
phalanx of digit 1.

INSERTION. The short flat tendon which forms overrides but joins with the
tendon of the ulnar part of the flexor pollicis brevis to insert on the ulnar volar
lateral aspect of the proximal phalanx of digit 1.

Action. This muscle will adduct and flex the metacarpal-phalangeal joint
of digit 1.

THE AccEssoRY INTRINSIC MUSCLES OF DIGIT 2

57. Abductor digiti secundi.

This muscle is a short, somewhat thick, muscle which overlies the interossei
musculature.
OrIcIN. Fleshy fibers arise from the volar surface of the base of metacarpal

488 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

1. The muscle then passes slightly obliquely across the thumb web to the radial
side of the base of the proximal phalanx of digit 2.

INSERTION. Tendinous fibers developing at this level have a dual insertion.
The more superficial fibers form a sheetlike aponeurosis which passes dorsally
into the extensor hood and dorsal extensor aponeurosis. The deeper fibers join
with the tendon at the radial part of the interosseus muscle for this digit and in-
sert through a sesamoid bone into the base of the radial side of the proximal
phalanx.

AcTIon. The more superficial fibers which join with the extensor aponeuro-
sis act to extend the proximal interphalangeal joint and to flex the metacarpal-
phalangeal joint. The deeper fibers serve to radialflex the metacarpal-phalan-
geal joint.

58. Adductor digiti secundi.

This slender fusiform muscle lies in the same plane with the abductor digiti
secundi and also overlies the deeper interossei muscles.

OricIn. Fleshy fibers arise from a small area of fibrous capsule at the base
of metacarpal 3 slightly proximal and to the ulnar side of the origin of the inter-
osseus of digit 2. The muscle then passes slightly obliquely and distally to the
ulnar side of digit 2 where a short tendon develops.

INSERTION. At the metacarpal-phalangeal joint level the tendon passes
close to the volar plate and sesamoid bone and volar to the tendon of the ulnar
half of the interosseus muscle of this digit. The insertion is into the base of the
proximal phalanx on the ulnar side.

AcTIoN. This muscle would adduct digit 2 at the metacarpal-phalangeal
joint level and also assist in flexion of this joint.

THE INTEROSSEI MUSCLES

59. Interossei.

These muscles, three in number, are the deepest intrinsic muscles of the
hand. They are short and thick and somewhat fusiform in shape. Each muscle
divides about halfway from its origin into radial and ulnar halves which then
pass to the respective sides of each digit at metacarpal-phalangeal joint level.

OricIN. All interossei muscles arise as fleshy fibers from the volar aspect
of the fibrous capsule at the base of the metacarpals. Interosseus 2 between
metacarpals 2 and 3, interosseus 3 at the base of metacarpal 3, and interosseus
4 at the base of metacarpal 4. The ulnar half of interosseus 4 is the largest of
the interossei divisions and shows a slight tendency toward further division in
its distal one-third.

INSERTION. With the exception of the radial half of interosseus 2, all inter-
ossei form flattened tendons passing close to the volar plate at metacarpal-

VoL. XXXIX] HOWARD: SEA OTTER FORELIMB 489

Siem

Ficure 64. Right humerus, lateral view. Key: 29, brachialis (origin); 31, exten-
sor carpi radialis longus (origin); 32, extensor carpi radialis brevis (origin); 33, exten-
sor digitorum communis (origin); 34, extensor digitorum lateralis (origin); 35, extensor
carpi ulnaris (origin); 37, supinator (origin); 40, palmaris longus (origin); 42, flexor
digitorum profundus (humeral head—origin); 43, flexor carpi radialis (origin); 44,
pronator teres (origin); 5, clavobrachialis (insertion); 13, pectoralis major (insertion) ;
14, pectoralis minor (insertion) ; 15, latissimus dorsi (insertion) ; 16, teres major (insertion) ;
26, supraspinatus (insertion) ; 27, subscapularis (insertion) ; A, head of humerus; B, greater
tuberosity; C, pectoral ridge; D, deltoid ridge; E, lateral epicondyle; F, lateral epicondylar
ridge; G, medial epicondyle; H, lesser tuberosity.

490 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Ficure 65. Right humerus, anterior view. Key: 22, triceps brachii caput medialis
(two of three sites of origin); 40, palmaris longus (origin); 42, flexor digitorum pro-
fundus (humeral head—origin); 43, flexor carpi radialis (origin); 44, pronator teres
(origin) ; 13, pectoralis major (insertion); 14, pectoralis minor (insertion); 15, latissimus
dorsi (insertion) ; 16, teres major (insertion); A, head of humerus; B, greater tuberosity;
C, pectoral ridge; D, medial epicondyle; E, lesser tuberosity; F, medial epicondylar ridge.

VoL. XXXIX] HOWARD: SEA OTTER FORELIMB 491

Ficure 66. Right humerus, medial view. Key: 22, triceps brachii caput medialis
(origin) ; 23, anconeus externus (origin); 29, brachialis (origin); 27, subscapularis (inser-
tion) ; A, head of humerus; B, lesser tuberosity; C, medial epicondyle; D, medial epicondy-
lar foramen; E, lateral epicondylar ridge.

492 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

\\

4

= Eh SGA

TLL EEE

C4
Deen

Ficure 67. Right ulna and radius, posterior view. Key: 45, flexor digitorum profun-
dus (humeral head—origin) ; 19, dorsoepitrochlearis (insertion) ; 20, triceps brachii caput
longus (insertion); 21, triceps brachii caput lateralis (insertion); 22, triceps brachii caput
medialis (insertion) ; 30, brachioradialis (insertion) ; 37, supinator (insertion); A, olecranon
process of ulna; B, semilunar notch; C, styloid process; D, radial styloid; E, biceps tubercle
of radius; F, head of radius.

VoL. XXXIX] HOWARD: SEA OTTER FORELIMB 493

A

12 | PY xe — B

Ficure 68. Right ulna and radius, lateral view. Key: 36, extensor pollicis et indicis
longus (origin); 38, abductor pollicis longus (origin); 12, pectoantebrachialis (into fas-
cia—insertion) ; 20, triceps brachii caput longus (insertion) ; 21, triceps brachii caput lateralis
(insertion) ; 22, triceps brachii caput medialis (insertion) ; 23, anconeus externus (insertion) ;
28, biceps brachii (insertion); 30, brachioradialis (insertion); 37, supinator (insertion) ;
44, pronator teres (insertion); A, olecranon process, ulna; B, semilunar notch; C, styloid
process, ulna; D, styloid process, radius; E, bicipital tuberosity; F, head of radius.

494 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Kae wm

Ficure 69. Right ulna and radius, anterior view. Key: 5, clavobrachialis (insertion) ;
12, pectoantebrachialis (insertion) ; 22, triceps brachii caput medialis (insertion) ; 23, anco-
neus externus (insertion); 28, biceps brachii (insertion); 29, brachialis (insertion); 30,
brachioradialis (insertion) ; 44, pronator teres (insertion); A, olecranon process, ulna; B,
semilunar notch; C, styloid process, ulna; D, styloid process, radius; E, bicipital tuberosity ;
F, head of radius.

VoL. XXXIX] HOWARD: SEA OTTER FORELIMB 495

SMmeenas

Ficure 70. Right ulna and radius, medial view. Key: 39, flexor carpi ulnaris (ori-
gin) ; 45, flexor digitorum profundus (central head—origin) ; 5, clavobrachialis (insertion) ;
12, pectoantebrachialis (insertion) ; 20, triceps brachii caput longus (insertion); 22, triceps
brachii caput medialis (insertion) ; 28, biceps brachii (insertion) ; 29, brachialis (insertion) ;
A, olecranon process, ulna; B, semilunar notch, ulna; C, articular surface for ulna; D, bi-
cipital tubercle, radius; E, radial head.

496 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Senn

Ficure 71. Right ulna and radius, articulated, anterior view. Key: 45, flexor digi-
torum profundus (central head—origin) ; 46, pronator quadratus (origin); 5, clavobrachi-
alis (insertion) ; 12, pectoantebrachialis (insertion) ; 28, biceps brachii (insertion) ; 29, brachi-
alis (insertion) ; 30, brachioradialis (insertion) ; 44, pronator teres (insertion); 46, pronator
quadratus (insertion); A, olecranon process, ulna; B, semilunar notch, ulna; C, styloid
process, ulna; D, interosseus membrane.

VoL. XXXIX] HOWARD: SEA OTTER FORELIMB 497

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Ficure 72. Right ulna and radius, articulated, posterior view. Key: 36, extensor
pollicis et indicis longus (origin); 38, abductor pollicis longus (origin); 37, supinator
(insertion) ; A, olecranon process, ulna; B, head of radius; C, styloid process, radius; D,
styloid process, ulna.

498 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

phalangeal joint level and then, as aponeurotic sheets, join into the extensor
hood and extensor aponeurosis of each digit forming the lateral band of the ex-
tensor aponeurosis. At the proximal interphalangeal joint level, these lateral
bands and extensor aponeuroses terminate into the central extensor tendon to
insert into the dorsum at the base of the middle phalanx. The extensor tendon
structures do not continue to the terminal phalanx as might be expected. The
tendon of the radial half of the interosseus 2 joins with the tendon of the ab-
ductor digiti secundi to insert through a sesamoid into the base of the proximal
phalanx as previously described.

AcTION. The interossei tendons joining into the extensor aponeurosis serve
to balance the action between the long extensor and flexors of these digits. Act-
ing alone, they would impart lateral motion to the extended digits or serve to
flex independently the metacarpal-phalangeal joints. With the metacarpal-
phalangeal joints in hyperextension, they would also act to extend the proximal
interphalangeal joints. In this way, the balancing effect prevents deformity
from action of the long extensors and flexor tendons alone.

MUSCLE ORIGINS AND INSERTIONS

In viewing the origin and insertion of the muscles as shown in the skeletal
figures, some confusion may arise owing to the bowing and torsion of the long
bones, particularly the humerus. The views as described of each plate are based
on the proximal ends of the bones. For example, a posterior view of the proximal
end of the humerus gives, as a result of torsion of this bone, a lateral view of
the distal end.

Each muscle retains its individual number and, therefore, to distinguish
between origin and insertion a circled number means the area of origin and an
uncircled number means the area of insertion.

The bones of the forefoot are not shown, as tendon insertions only are present
and these areas of insertion are shown quite adequately on the muscle plates.

SUMMARY

The gross muscular anatomy of the forelimb of the sea otter has been pre-
sented. For the hand the intrinsic and extrinsic musculature is sufficient to
provide for dexterity, but such is extremely limited by reason of the anatomical
restrictions imposed by the integument and soft parts.

The forefoot, obviously designed for terrestrial use, has become, by virtue
of bimanual use, a highly skilled appendage for other purposes.

ACKNOWLEDGMENTS

The State of California Deparment of Fish and Game was most cooperative
in providing the specimen for anatomical study. The California Academy of
Sciences provided the invaluable aid of a disarticulated sea otter skeleton.

VoL. XXXIX] HOWARD: SEA OTTER FORELIMB 499

The Anatomical Studies of the California River Otter, by Edna M. Fisher,
proved to be an excellent guide during the dissection, and her unpublished notes
on the sea otter anatomy, which were made available through the courtesy of
Mr. Fred Tarasoff, were also of value. Unpublished notes and sketches by Fred
Tarasoff were also available and of some assistance.

The dissection and storage of materials took place at the Hopkins Marine
Station of Stanford University in Pacific Grove, California, and the personnel
of this institution, particularly Dr. I. Abbott, gave encouragement and assistance
in every way possible for which I am deeply indebted.

BIBLIOGRAPHY

BarABASH-NIKIFoROV, I. I.
1962. The sea otter (Enhydra lutris L.). Biology and economic problems of breeding.
Translated from the Russian by the Israeli Program for Scientific Translations.
National Science Foundation and U.S. Department of the Interior, Washington,
DEC 227 pages:
FIisHER, Epna M.
1939. Habits of the southern sea otter. Journal of Mammalogy, vol. 20, no. 1, pp.
21-36.
1942. Osteology and myology of the California river otter. Stanford University Press,
Stanford, California, vi + 66 pages.
—. Notes and sketches on the myology of the southern sea otter. (Unpublished).
Courtesy of Fred Tarasoff.

GoLp_ER, F. A.
1925. Bering’s voyages. American Geographical Society, Research Series, no. 2, 290
pages.

GRINNELL, J., JosEpH S. Dixon, and JEAN M. LINSDALE
1937. Fur bearing mammals of California. University of California Press, Berkeley,
California, 777 pages.
Hart, E. R.
1926. Muscular anatomy of three mustelid mammals, Mephitis, Spilogale, and Martes.
University of California Publications in Zoology, vol. 30, no. 2, pp. 7-38.
VAT Kee I:
1963. Tool-using performances as indicators of behavioral adaptability. Current
Anthropology, vol. 4, December, pp. 479-494.
Harris, C. J.
1968. Otters: a study of the recent Lutrinae. Weidenfeld and Nicolson, London, 397
pages.
Howe tt, A. B.
1930. Aquatic mammals. Charles C. Thomas, Springfield, Illinois, and Baltimore,
Maryland, xii + 338 pages.
JoLuie, M.
1962. Chordate morphology. Reinhold, New York, 478 pages.
Jacosi, A.
1938. Der Seeotter. Monographien der Wildsaugetiere, Band VI, 93 pages. (Transla-
tion: Fisheries Research Board of Canada, no. 521.)

500 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Kenyon, K. W.

1969. The sea otter in the eastern Pacific Ocean. North American Fauna, no. 68, U.S.
Department of the Interior, Bureau of Sport Fisheries and Wildlife, Washing-
ton, 352 pp.

Murtg, O. J.

1940. Notes on the sea otter. Journal of Mammalogy, vol. 21, no. 2, pp. 119-131.
Orre RT:

1966. Vertebrate biology (second edition). W. B. Saunders Co., Philadelphia, xii ++

483 pages.
Romer, A. S.
1964. The vertebrate body. W. B. Saunders Co., Philadelphia and London, viii +
643 pages.

Soxotov, A. S., and J. I. SoxoLov
1970. Some specific features of the locomotor organs of the river otter and sea otter
associated with their mode of life. Moscovskoe Obshchestvo Ispytatelel Prirody,
Otdel Biologicheskii, vol. 75, no. 5, pp. 5-17. (Translated from the Russian,
Hopkins Marine Station, Pacific Grove, California, 1972.)
TayLor, W. P.
1914. The problem of aquatic adaptation in the Carnivora, as illustrated in the oste-
ology and evolution of the sea otter. University of California Publications, Bul-
letin of the Department of Geology, vol. 7, no. 25, pp. 465-495.
TarASoFF, F. J.
. Notes and sketches on the myology of the sea otter. (Unpublished.)
Tarasorr, F. J., A. BISAILLON, J. PirrArD, and A. P. WuHrIeT
1972. Locomotory patterns and external morphology of the river otter, sea otter, and
harp seal (Mammalia). Canadian Journal of Zoology, vol. 50, no. 7, pp. 915-
929.
VANDEVERE, J. E.
1969. Feeding behavior of the southern sea otter. Proceedings of the Sixth Annual
Conference on Biological Sonar and Diving Mammals, Stanford Research
Institute, pp. 87-94.

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PROCEEDINGS

OF THE

CALIFORNIA ACADEMY OF SCIENCES
FOURTH SERIES

Vol. XXXIX, No. 21, pp. 501-506; 3 figs. December 19, 1973

A NEW HAWAIIAN CHITON,
RHYSSOPLAX LINSLEYI
(MOLLUSCA: AMPHINEURA: CHITONIDAE)

By
Glenn E. Burghardt

Steinhart Aquarium, California Academy of Sciences
San Francisco, California 94118

It has always puzzled conchologists that the Hawaiian Islands, which are
so rich in marine mollusks, are poor in species of chitons. Only four Hawaiian
species are known: Acanthochitona viridis (Pease, 1871), A. armata (Pease,
1871), Ischnochiton petaloides (Gould, 1846), and the new species described
below.

This new species has been known for some years but has gone unnamed
until this time. The first specimens brought to the attention of the author were
collected by Mrs. Harold Gudnason at Puu o Hulu Beach, near Maili, Oahu,
Hawaii, in the intertidal waters at low tide (0.0) on February 26, 1965.

Rhyssoplax linsleyi Burghardt, new species.
(Figures 1-3.)

Dracnosts. The surface of the valves, while appearing smooth to the
naked eye, looks pitted when viewed under low magnification. Under high
magnification however, it is clear that the surface is indeed smooth, the ‘pitted’
appearance being caused by a color pattern composed of marks shaped like
small starbursts.

In addition to the pattern of ‘pit marks,’ the anterior valve has a series of
between 26 and 36 (holotype has 36) shallow grooves which are weak and
difficult to see unless viewed under side light. The interior of the valve has
a series of 8 slits on the outer edge of the insertion plate. This edge is also
finely pectinated, a feature typical of the family Chitonidae.

Intermediate valves have the lateral areas defined but not sharply raised.

[501]

502 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

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(Up

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eet Te

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Pr

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Ficure 1. Holotype of Rhyssoplax linsleyiz. Length 13.3 mm., width 8.4 mm., height 2.5
mm.; Puu o Hulu Beach, Oahu, Hawaii.

They possess the same radiating series of bifurcating grooves as does the
anterior valve. Generally, these grooves are three in number (as in the
holotype), but can be single on young specimens. The central areas of these
valves are cut longitudinally with from 10 to 15 (13 in holotype) sharp lines.
Both lateral and central areas have the ‘pitted’ color pattern. The dorsal
ridge is smooth and shiny with the valves beaked. The jugal area is also smooth
but shows the ‘pit marks.’ The outer margins of the insertion plates are
pectinated and have a single slit.

The posterior valve appears smooth behind the slightly anterior mucro
but also has the unique ‘pitted’ color pattern. This area has a series of from
23 to 32 (26 in holotype) shallow radial grooves similar to those of the

VoLt. XXXIX] BURGHARDT: HAWAIIAN CHITON 503

Ficure 2. Rhyssoplax linsleyi: (a) anterior valve, (b) median valve, and (c) posterior
valve. Shown are both the dorsal surface (left) and interior surface (right) of the valves.

anterior valve. The interior of this valve shows the same series of slits—here
numbering 12—and pectinations as does the anterior valve.

DescripTION. Rhyssoplax linsleyi is a small chiton, averaging 9.6 mm.
in length (mean obtained from 116 specimens). It is fairly broad, with an
average ratio of width to body length of 3:5. The valves are moderately
arched, the average height being in a ratio of 1:5 to the body length.

The color of the valves is quite variable, ranging from a clean grey-white
to a mottled black and white (holotype) or green and white. A few specimens
in our sample were solid in color, either red or white, but such coloration
appears to be rare. The color pattern occurs on both lateral and central areas.
The interior of the valves is white with a slight bluish tint.

The girdle is of medium width and is covered with the fairly large, smooth

504 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

1B: Maui

Ficure 3. Map of Hawaiian Islands showing the collecting sites of Rhyssoplax linsleyi
on the islands of Oahu, Maui, and Kauai: (1) Puu o Hulu (type locality), (2) Ala Moana,
(3) Wainanalo, (4) Kuhuku Point, (5) Honolua, and (6) Poipu Beach.

scales typical of Chitonidae. This girdle is cream with green bands except on
the solid colored specimens, where the girdle matches the valves in color.

No radular studies were made.

Type LocaLity. The type series was collected at Puu o Hulu Beach, near
Maili, Oahu, Hawaii, on 26 February 1965, by Mrs. Harold Gudnason.

TYPE MATERIAL. Holotype. An adult specimen preserved in alcohol is
deposited in the California Academy of Sciences Department of Invertebrate
Zoology Type Series, no. 567. Measurements are 13.3 mm. in length. 8.4 mm.
in width, 2.5 mm. in height.

Paratypes. Twenty-four paratypes collected with the holotype have been
deposited in the mollusk type collections of the following museums and private
collectors: Academy of Natural Sciences of Philadelphia, two specimens, ANSP
330098; Bernice P. Bishop Museum, two specimens, BPBM 206931; Cali-
fornia Academy of Sciences, two specimens, CAS Geology 53956 (rearticu-
lated), CASIZ type series no. 594; Los Angeles County Museum of Natural
History, two specimens, LACM 1620; Museum of Comparative Zoology, two
specimens, MCZ 272531; San Diego Natural History Museum, two specimens,
SDSNH 62724, 62725; National Museum of Natural History, two specimens,
USNM 735016; Glenn and Laura Burghardt, six specimens; Mrs. Harold
Gudnason, four specimens.

VoL. XXXIX] BURGHARDT: HAWAIIAN CHITON 505

Discussion. This chiton has been collected from all sides of Oahu (Maili,
Kahuku, Waimonalo, and Ala Moana). It has also been collected by Dr.
Antonio Ferreira at Honolua Bay on the northwestern side of Maui, and by
Dr. George Ramsay at Poipu Beach on the southern side of Kauai. It is found
from the intertidal zone to a depth of 15 feet but not as an emergent species,
as it has never been discovered completely exposed by the low tides. Generally,
specimens can be found by turning small rocks, exposing the chitons clinging
to the undersides.

Measurements of 116 specimens studied are as follows:

Oahu (82 specimens) Maui (34 specimens)

Range Mean Range Mean
Length 3.5-14.5 8.0 mm. 7.0-21.0 13.4 mm.
Width 2.8— 6.2 4.9 mm. 4.0-12.4 8.2 mm.
Height 0.7— 2.9 1.8 mm. —- -=

According to Smith (1966) and Hyman (1967), many chitons brood their
eggs and some even carry their young in the pallial cavity. Upon examining
the specimens for this paper, I found that several were carrying eggs. The
eggs were found to be slightly less than 0.2 mm. in diameter.

This new species belongs to the family Chitonidae Rafinesque, 1915, genus
Rhyssoplax Thiele, 1893, the type species being the North African Chiton
affinis Issel, 1869 (International Committee on Zoological Nomenclature, 1971,
p. 18). The approximately 30 species of Rhyssoplax have been recorded from
Africa, Japan, Australia, New Zealand, and now the distribution includes the
Hawaiian Islands. No species of Rhyssoplax occurs on the Pacific coast of
North America (Burghardt and Burghardt, 1969). Species in this group are
sometimes referred to the genus Anthochiton Thiele, 1893, but Iredale (1914,
p. 40) and Smith (1960, pp. 65-66) have referred Anthochiton to the synonymy
of Rhyssoplax Thiele, 1893, which has been placed on the Official List of
Generic Names in Zoology (International Committee on Zoological Nomen-
clature, 1971, p. 18).

Rhyssoplax linsleyi does not closely resemble any other described species.
Its unique ‘pitted’ pattern, discussed above, distinguishes this species from all
other Rhyssoplax known to date. Edmonson (1946, p. 118, fig. 546) mentions
an undescribed chiton from Kanoehe Bay which undoubtedly belongs to R.
linsleyi. In the Bishop Museum there are two lots of chitons which were
labelled as a new species by Dall (BPBM no. 895.8 and 895.8a), but not
published (personal communication, Mrs. H. Gudnason). These also represent
R. linsleyi.

This new species is named in honor of Mrs. Gudnason’s father, Earle G.
Linsley, Honorary Associate in Astronomy, Bernice P. Bishop Museum and
Planetarium (1957-1962).

506 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

The author wishes to express his thanks to Mr. Allyn G. Smith of the
California Academy of Sciences for his review and assistance, to Mrs. Harold
Gudnason for permission to work on her Oahu specimens, to Dr. Antonio
Ferreira for permission to work on his Maui specimens, and to Betsy Harrison
and Hilda Manzak for specimens used for this study; and a special thanks to
wife, Laura, whose immeasurable help has made this paper possible. The
figures were done by Roderick MacPherson.

LITERATURE CITED

BurGHARDT, GLENN, AND LAURA BURGHARDT

1969. Collector’s Guide to West Coast Chitons. Special Publication No. 4, San Francisco

Aquarium Society, 45 pp., 4 pls.
EDMONDSON, CHARLES H.

1946. Reef and shore fauna of Hawaii. Special Publication no, 22, Bernice P. Bishop

Museum, 381 pp., 223 figs.
HyMaANn, LIBBIE

1967. The Invertebrates. Vol. 6, Mollusca no. 1. Pp. 70-142, figs. 29-57. New York,
McGraw Hill.

INTERNATIONAL COMMITTEE ON ZOOLOGICAL NOMENCLATURE ]

1971. Opinion 951. Rhyssoplax Thiele, 1893 (Amphineura): designation of a type-
species under the plenary powers. Bulletin of Zoological Nomenclature, vol.
28, pts. 1 and 2, pp. 18-19.

TIREDALE, ToM

1914. The chiton fauna of the Kermadec Islands. Proceedings of the Malacological

Society of London, vol. 11, pp. 25-51, pls. I-II.
SmitH, ALLYN G.

1960. Amphineura. Pp. 41—76, figs. 33-45, in: R. C. Moore (Ed.), “Treatise on Inverte-
brate Paleontology,’ Part I, Mollusca 1. Lawrence, Kansas, Geological Society
of America and University of Kansas Press.

1966. The larval development of chitons (Amphineura). Proceedings of the California
Academy of Sciences, vol. 32, no. 15, pp. 433-446, 11 figs.

arino =RIAN viCal
Vii hii Di Up

LIBRAK"
1974
PROCEEDINGS
OF THE Woods Hole, Mass

CALIFORNIA ACADEMY -OF-SCTENCES

FOURTH SERIES

Vol. XXXIX, No. 22, pp. 507-516; 3 figs. June 27, 1974

NEZUMIA (KURONEZUMIA) BUBONIS, A NEW
SUBGENUS AND SPECIES OF GRENADIER
(MACROURIDAE: PISCES) FROM HAWAII AND
THE WESTERN NORTH ATLANTIC

By
Tomio Iwamoto

California Academy of Sciences
San Francisco, California 94118

INTRODUCTION

In the course of exploratory fishing by the R/V Oregon off the Caribbean
coast of Colombia in the spring of 1964, a large black grenadier was captured by
a trawl fished at a depth of 400 fathoms. The fish resembled Nezumia atlantica
(Parr, 1946) in its general body shape and dentition, but it differed widely in
several meristic and morphometric features and in having a velvetlike texture
to its squamation and a peculiarly enlarged scaly swelling anterior to the vent.
Additional specimens were subsequently collected both in the Caribbean Sea
and in the Gulf of Mexico by the R/V Pillsbury and the R/V Oregon II. The
surprising presence of this species off Hawaii was revealed upon examination
of the fishes collected by Paul J. Struhsaker of the National Marine Fisheries
Service. An undescribed, closely related species from the South China Sea was
examined in June, 1970, in the collections of the Fisheries Research Station,
Aberdeen, Hong Kong.

In squamation and in head shape the new species resembles certain species
of Malacocephalus, and the swelling before the vent appears to be an exaggera-
tion of the slight swelling before the vent in members of that genus; but the
dentition, the strong serrations on the second spinous dorsal ray, and several
other features are totally in disagreement. In such features as length of upper
jaw and number of pyloric caeca, the species approaches certain species of Ventrt-
fossa; but in other more important characters, such as the structure of the light

[507]

508 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

organ and the shape of the head, it differs widely from most members of that
genus.

The difficulty of allocating the species to a genus has led to further, more
extensive study of the generic problems concerning Malacocephalus Gunther,
1862; Ventrifossa Gilbert and Hubbs, 1920; and Nezumia Jordan, 1904 (Iwa-
moto, MS). Publication of that study is being delayed because of the desire to
incorporate much additional material. The description given here of the new
species of Nezumia is being published at this time to make the name available to
other workers. I have refrained from describing as new the species from the
South China Sea because permanent disposition of the single representative has
not been assured. The specimen now is at the Aberdeen Fisheries Research
Station, Hong Kong. The new species and its closest congener from the South
China Sea share certain characters (as indicated in the diagnosis below) so dif-
ferent from those in other members of the genus Nezumia that subgeneric status
is designated.

METHODS

Methods for making counts and measurements generally follow procedures
outlined by Hubbs and Lagler (1958) and further described for macrourids by
Gilbert and Hubbs (1916) and Iwamoto (1970). The rounded snout of the spe-
cies treated here and the lack of a prominent terminal scute make determination
of the anterior tip of the nasal rostrum difficult. For this reason, measurements
normally taken from the tip of the nasal rostrum (e.g., snout length, head length)
are instead taken from whichever median point on the snout is more anterior.
All rays of the pectoral fin are counted, including the short, usually thin, sliver-
like uppermost ray and the 1 to 3 small lowermost rays. All gillrakers, including
rudimentary ones, along the medial sides of the first and second gill arches are
counted.

Institutional abbreviations are as follows: BPBM—Bernice P. Bishop
Museum, Honolulu, Hawaii; CAS—California Academy of Sciences, San Fran-
cisco, California; FRSA—Fisheries Research Station, Aberdeen, Hong Kong;
UMML—University of Miami, Rosenstiel School of Marine and Atmospheric
Sciences, Miami, Florida; USNM—Division of Fishes, United States National
Museum of Natural History, Washington, D.C.

ACKNOWLEDGMENTS

I thank the persons listed below for their assistance during the preparation
of this paper. The following made specimens available for examination: Harvey
R. Bullis, National Marine Fisheries Service (NMFS), Miami, Florida; Edward
F. Klima, NMFS, Pascagoula, Mississippi; Paul J. Struhsaker, NMFS, Hono-
lulu, Hawaii; William Chan, Fisheries Research Station, Aberdeen, Hong Kong;
C. Richard Robins, Rosenstiel School of Marine and Atmospheric Sciences,
Miami, Florida. Lillian J. Dempster and William N. Eschmeyer critically re-
viewed the manuscript.

Vol. XXXIX] IWAMOTO: NEZUMIA BUBONIS 509

Nezumia Jordan im Jordan and Starks, 1904
Subgenus Kuronezumia Iwamoto, new subgenus

Dracnosis. Species of Nezumia with light organ peculiarly enlarged into a
bulbous, scaly, wartlike structure anterior to anus and between pelvic fin bases
(fig. 3); bulbous structure housing a large lens which partly encapsulates
luminescent gland that fronts anterior wall of rectum. Light organ closely asso-
ciated with pelvic girdle, the large lens essentially overriding part of girdle. An-
terior dermal window of light organ at bottom of deep fossa and covered by
bulbous lens-luminescent-gland mass. Snout almost entirely covered with small
uniform scales; only thin margin above upper lips naked. Scales on suborbital
region small, uniform, not forming a stout ridge. Body scales small, densely
covered with extremely fine erect spinules. Color dark brown to black. Second
spinous ray strongly serrated.

Remarks. The bulbous structure housing the light organ in species of the
subgenus Kuronezumia is apparently unique in the family, although the struc-
ture is vaguely approached by a swelling in the same region in members of
Malacocephalus. (It appears to be an anteroventral swelling of the scaled area
between the periproct and the anterior dermal window and a gross hypertrophy
of the condition in Malacocephalus, where the area is slightly raised and the
anterior dermal window sits in a relatively deep fossa, with the posterior wall of
the depression steep, but not overlapping.) The species of the new subgenus
have a darker overall coloration than almost every other member of Nezumia
and resemble Trachonurus in that regard. Members of the subgenus are prob-
ably the largest of the genus, with V. (Kuronezumia) bubonis attaining a length
of over 630 mm. The largest specimen of N. atlantica (another large species
of Nezumia) I have measured is 450 mm. in total length.

Etymotocy. The name is derived from the Japanese, kuroi, black or dark,
and nezumi, rat, and is to be treated as feminine in gender.

Nezumia bubonis Iwamoto, new species.
(Figures 1-3.)

Counts. First dorsal rays II, 10-12; pectoral fin rays 23-26; pelvic fin rays
11-12 (one specimen with 9 in one fin, another specimen with 13 in one fin).
Gillrakers on first arch 1-2 + 7-9 (usually 2-+ 8); rakers on second arch
1-2 + 7-9. Scales below origin of first dorsal fin 14 to about 20, below origin of
second dorsal fin 12-14; lateral line scales counted from the anteriormost
scale posteriorly over distance equal to predorsal length 47-61. Abdominal
vertebrae 13.

MorpHoMEtryY. Head length of 10 specimens examined 30 to 125 mm.;
total length 140+ to 630 mm. The following measurements are in percent of
head length: snout length 26-32; ventral length of snout 13—18; orbit diameter
23-31; interorbital width 23-26; postorbital length 44-51; distance orbit to

[Proc. 4TH Ser.

CALIFORNIA ACADEMY OF SCIENCES

510

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Vol. XXXIX]

IWAMOTO: NEZUMIA BUBONIS

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CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

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Ficure 3. Ventral view of abdominal area of Nezumia bubonis showing position of bul-
bous swelling of light organ (l.0.) before anus. A—origin of anal fin.

angle of preopercle 34-44; suborbital width 17—20; length upper jaw 35-44;
greatest body depth 88-109; distance from base of outer pelvic fin to origin of
anal fin 23-37 (usually 31-37); distance anus to anal origin 11—22; interspace
between first and second dorsal fins 19-34; height first dorsal fin 89-92 (2
specimens); length pectoral fin 49-66; length outer pelvic ray 40-64; length
barbel 20-29; length outer gill slit 16-21.

DescriPTION. Body deep, its greatest depth just anterior to first dorsal
fin; trunk short, distance isthmus to anal origin about one-half length of head;
body tapering rapidly behind trunk into long straplike tail. Head about one-
fifth total length, laterally compressed, sides almost flat with no prominent
projections or ridges; interorbital space slightly convex, proportionately wid-
est in largest specimens; suborbital area broad, flat, evenly and completely
scaled, without a coarsely scaled ridge. Mouth almost terminal; posterior end
of maxillary lying under anterior half of orbits. Snout broadly rounded in
profile, without a well-marked tip, generally not projecting beyond upper jaws.
Orbits elliptic, proportionately largest in smallest specimens, usually longer in
vertical than in horizontal diameter; upper margins not entering dorsal profile.
Gill membranes united just behind vertical through posterior margin of orbits,
attached to isthmus with a moderately narrow posterior free fold.

Fin positions best seen in illustration of 59-mm.-head-length paratype from
Hawaii (fig. 2). Severe dorsal flexure distorts normal positions of fins of holo-
type. First dorsal fin complete in only two paratypes from Hawaii, measuring

Vol. XXXIX] IWAMOTO: NEZUMIA BUBONIS 513

89 and 92 percent of head length; spinous second ray strongly serrated along
leading edge and slightly produced. Pelvic fins finely but densely scaled along
exposed bases; outer ray slightly prolonged beyond other rays.

Head and body generally evenly and completely scaled. Exposed posterior
and posteroventral margins of interoperculum scaled. Gular and _branchio-
stegal membranes generally naked, although a few small patches of deeply
embedded scales occasionally found at bases of branchiostegal rays; in one
specimen, a few scales found at extreme tips of fourth and fifth branchiostegal
rays. A narrow margin of scaleless skin along snout just above upper lip.

Scales on body and head very small and strongly adherent. Long, fine,
slender spinules densely cover scales on body; spinules on nape scales almost
erect, those on other parts of body reclined at about a 30-degree angle from
vertical. Scales on head tend to have somewhat more stubby and conical spi-
nules generally arranged in 3—7 radiating rows. No stout modified scales present
on snout or suborbital region.

Pyloric caeca of two specimens moderately long (about 35 percent head
length), unbranched, 35 and 39 in number. Gas glands of swim bladder two,
kidney-shaped in outline although almost round in cross section; retia two,
long and thin, about 16 mm. long in 113-mm.-head-length specimen.

Dentition consists of broad villiform bands of small teeth in both jaws with
a slightly enlarged, evenly spaced outer series in upper jaw.

General color swarthy to dark brown with ventral surfaces of head, gill
covers, and abdomen black to brownish black. Lips, gular and branchiostegal
membranes, barbel, margins of orbits, periproct, and scaleless ventral margin
of snout black. Buccal membranes light gray to dark gray. Gill cavity linings
generally blackish towards outer and upper margins and paler ventrally and
deeper within the cavity. Peritoneal membrane pallid with numerous black
specks. All fins black.

COMPARISONS AND RELATIONSHIPS. The closest relative of Nezumia bubo-
nis is an undescribed species from the South China Sea. Together they comprise
the subgenus Kuronezumia. Nezumia bubonis differs from the undescribed
species in having more pelvic fin rays (normally 11-12 compared with 8),
slightly fewer scales below the second dorsal fin (12-14 versus 16) and a nar-
rower interorbital space (23-26 percent of head length compared with 29
percent).

Nezumia darus (Gilbert and Hubbs, 1916) most closely approaches the
members of subgenus Kuronezumia by its somewhat similar squamation and
head physiognomy. The absence of ventral tubercular swellings, the more
pronounced terminal snout scute, and the fewer scale rows below the second
dorsal fin immediately separate V. darus from both N. bubonis and its South
China Sea counterpart. Nezumia burragei (Gilbert, 1905), N. hebetatus (Gil-
bert, 1905), and N. macronemus (Smith and Radcliffe im Radcliffe, 1912)

514 CALIFORNIA ACADEMY OF SCIENCES [PRoc. 4TH SER.

all fall in a loose group near NV. darus in their resemblance to the species of
subgenus Kuronezumia. But, in addition to the subgeneric distinctions, NV.
burragei differs in having a well developed terminal snout scute and a sub-
orbital ridge formed of two rows of stout scutelike scales, and most of the
ventral surfaces of the snout and suborbital area naked; NV. hebetatus differs in
having a more pointed snout, extensive naked areas on both the upper and lower
surfaces of the snout and on the ventral surface of the suborbital region, a
narrow suborbital ridge formed of two rows of stout scales, and a posterior
margin of the preoperculum that is inclined forward; and N. macronemus
differs in having a more pointed snout and lanceolate scale spinules.

DIsTRIBUTION. Off the Hawaiian Islands, in the Gulf of Mexico, and in
the Caribbean Sea. Capture depths ranged from 732 to 1062 meters.

Etymotocy. The name comes from the Greek, bubonis, a tumor, and is to
be treated as a noun in apposition.

ReMARKS. The peculiar distribution pattern of this species cannot be ade-
quately explained because of the paucity of material and our present general
lack of knowledge of dispersal means of macrourid fishes. Probably the species
is much more widely distributed and the lack of more locality records is simply
a reflection of sketchy collecting efforts.

SPECIMENS EXAMINED. Holotype: CAS no. 27872, 86 mm. head length,
480 mm. total length (including a small pseudocaudal), western Gulf of Mexico
off Barra San Antonio, Mexico, (24°49’N., 96°27’W.), in 500 fathoms (914
meters), by 40-ft. otter trawl, R/V Oregon station 4814, 12 March 1964, bottom
temperature 41°F (5°C).

Paratypes (9 specimens from 7 localities): USNM 210592 (2, 53-100 mm.
head length, 284-500 mm. total length), off Colombia, R/V Oregon station
4902 (9°02’N., 76°31.5’W.), in 400 fathoms (732 meters), by 65-ft. otter trawl,
28 May 1964; UMML uncatalogued (1, 111 mm. H.L., 550 mm. T.L.), off
Colombia, R/V Pillsbury station 388 (10°16’N., 76°03’W.), in 450-580 fathoms
(823-1061 meters), by 40-ft. otter trawl, 15 July 1966; CAS no. 27873 (1, 125
mm. H.L., 630 mm. T.L.), Gulf of Mexico, R/V Oregon IJ station 10955
(21°41’N., 96°55’W.), in 490 fathoms (896 meters), by 150-ft. otter trawl,
3 June 1970; USNM 210593 (1, 113 mm. H.L., 560 mm. T.L.), Gulf of Mexico,
R/V Oregon II station 11136 (24°27’N., 87°38’W.), in 500 fathoms (914
meters), by 71-ft. otter trawl, 9 August 1970; BPBM uncatalogued (1, 39.6
mm. H.L., 180 + mm. T.L.), Hawaiian Islands, R/V Townsend Cromwell cruise
35, station 24 (21°06.5’N., 156°13.5’W.), in 640-686 meters, by 41-ft. otter
trawl, 5 April 1968; CAS no. 27874 (2, 30-56 mm. H.L., 140 + —246 mm. T.L.),
Hawaiian Islands, R/V Townsend Cromwell cruise 40, station 86 (21°06.8’N.,
156°13.7’W.), in 631-705 meters, by 41-ft. otter trawl, 23 November 1968;
BPBM uncatalogued (1, 59 mm. H.L., 247 mm. T.L.), Hawaiian Islands, R/V

Vol. XXXIX] IWAMOTO: NEZUMIA BUBONIS 515

Townsend Cromwell cruise 40, station 87 (21°04.6’N., 156°10.6’W.), in 623--
667 meters, by 41-ft. otter trawl, 23 November 1968.

Nezumia species.

Counts. First dorsal fin rays II,10; pectoral fin rays 24; pelvic fin rays 8.
Gillrakers on first arch 2 + 8; on second arch 2+ 7. Scales below origin of
first dorsal fin 17-18; below origin of second dorsal fin 16; lateral line scales
counted from the anteriormost scale for a distance equal to the predorsal length
50.

MorpHoMEtRY. The following are in percent of head length: snout length 30;
orbit diameter 25; interorbital width 29; postorbital length 48; distance orbit
to angle of preopercle 41; suborbital width 18; length upper jaw 41; greatest
body depth 100; interspace between first and second dorsal fins 21; height of first
dorsal fin 93; length pectoral fin 63; length outer pelvic ray 55; length barbel
22; length outer gill slit 20.

Dracnosis. A species of Vezumia of the subgenus Kuronezumia with 8 pelvic
fin rays. Scales very small, finely spinulated, 16 rows below origin of second
dorsal fin. Pectoral fin rays 24. Interorbital width 29 percent of head length.

REMARKS. The species agrees with the description previously given of Nezu-
mia bubonis so closely that it would be redundant to repeat the same characters.
A comparison of the differences between the two species of the subgenus is given
in the description of NV. bubonis.

DISTRIBUTION. South China Sea.

SPECIMEN EXAMINED. FRSA uncatalogued, (50 mm. H.L.), from the South
China Sea.

LITERATURE CITED

GILBERT, CHARLES H.

1905. The deep-sea fishes of the Hawaiian Islands. Pp. 576-713, pls. 66-101, figs. 230-
276. In, David Starr Jordan and Barton Warren Evermann: The aquatic re-
sources of the Hawaiian Islands. Bulletin of the United States Fish Commis-
sion, vol. 23, part 2.

GILBERT, CHARLES H., AND CArt L. Hupps

1916. Report on the Japanese macrouroid fishes collected by the United States fisheries
steamer “Albatross” in 1906, with a synopsis of the genera. Proceedings of the
United States National Museum, vol. 51, no. 2149, pp. 135-214, pls. 8-11.

1920. Contributions to the biology of the Philippine Archipelago and adjacent region.
The macrouroid fishes of the Philippine Islands and the East Indies. United
States National Museum, Bulletin 100, vol. 1, part 7, pp. 369-588.

GUNTHER, ALBERT
1862. Catalogue of the fishes in the British Museum, vol. 4, pp. 1-534.
Husps, Cart L., anpD Kart F. LAGLER

1958. Fishes of the Great Lakes region. Revised edition. Cranbrook Institute of Sci-

ence, bulletin 26, pp. 1-213.

516 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Twamoto, Tomio
1970. The R/V Pillsbury deep-sea biological expedition to the Gulf of Guinea, 1964-65.
19. Macrourid fishes of the Gulf of Guinea. Studies in Tropical Oceanography,
Miami, no. 4, part 2, pp. 316-431, figs. 1-27.
(MS). Macrourid fishes of the tribe Malacocephalini (Macrouridae: Gadiformes).
Doctoral dissertation submitted to the University of Miami, June 1972.
JorpAN, Davin S., AnD Epwrn C. STARKS
1904. List of fishes dredged by the steamer Albatross off the coast of Japan in the
summer of 1900, with descriptions of new species and a review of the Japanese
Macrouridae [by Jordan and Gilbert]. Bulletin of the United States Fish
Commission, vol. 22 for 1902, pp. 577-630, pls. 1-8.
Parr, ALBERT E.
1946. The Macrouridae of the western North Atlantic and Central American seas.
Bulletin of the Bingham Oceanographic Collections, no. 10, part 1, pp. 1-99.
RADCLIFFE, LEWIS
1912. Scientific results of the Philippine cruise of the fisheries steamer ‘‘Albatross,”
1907-1910. No. 21. Descriptions of a new family, two new genera, and twenty-
nine new species of anacanthine fishes from the Philippine Islands and con-
tiguous waters. Proceedings of the United States National Museum, vol. 43,
0. 1924, pp. 105-140, pls. 22-31, figs. 1-11.

PROCEEDINGS 90050 pi tenical Jal

OF THE eli ARY

ES Ets

CALIFORNIA ACADEMY OF SCIENCES, ,

FOURTH SERIES
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/

Vol. XXXIX, No. 23, pp. 517-562; 6 figs.; 4 tables.“~~~~~~June*27, 1974

REVISED DIATOM STRATIGRAPHY OF THE
EXPERIMENTAL MOHOLE DRILLING,
GUADALUPE SITE

By

Hans-Joachim Schrader

Geologisch-Paldéontologisches Institut und Museum der Universitat Kiel,
23 Kiel, Olshausenstrafe 40/60, F. R. Germany

Apstract: Thirty-four diatom-bearing samples were obtained from cores recovered
at the Experimental Mohole Drilling (EMD), Guadalupe Site. A planktonic-diatom
stratigraphy is defined through correlation of ranges of key species of the EMD with
a continuously cored section off California (DSDP, leg 18, Site 173). The Pliocene-
Miocene boundary (defined as the base of the Gilbert Reversed Paleomagnetic
Epoch) occurs between samples EMD-8-10-44/45 cm. and EMD-8-10-100/101 cm.
The Middle Miocene to Late Miocene boundary (defined as occurring within Geo-
magnetic Epoch 11) occurs between samples EMD-8-15-100/101 cm. and EMD-8-
15-200/201 cm. The youngest investigated sediment (EMD-8-9-5/6 cm.) cor-
relates with North Pacific Diatom Zone 10 of Middle Pliocene age (upper part of
the Gilbert Magnetic Epoch); the oldest investigated sediment (EMD-7-2-0/17
cm.) correlates with North Pacific Diatom Zone 24, with NN 6 Standard Nanno-
plankton Zone, and with the Dictyocha octacantha Silicoflagellate Zone. The sedi-
mentation rate for the interval from 80 to 95 meters below the sea floor is about 7
meters/million years; it increases abruptly over the interval from 95 to 115 meters
below the sea floor to approximately 30 m./m.y. and decreases again to approxi-
mately 7 m./m.y. for the interval from 115 to 135 meters below the sea floor.
One new genus and ten new species of diatoms are described, and five new com-
binations of names are proposed.

List of new taxonomic entries and new combinations:

a. New genus.

Cussia

[517]

518 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

b. New species.

Biddulphia moholensis CAS 54409
Coscinodiscus moholensis CAS 54410
Cussia lancettula CAS 54411
Cussia moholensis CAS 54413
Nitzschia burcklia CAS 54415
Nitzschia kanayensis CAS 54417
Nitzschia moholensis CAS 54419
Nitzschia riedelia CAS 54420
Nitzschia seiboldia CAS 54422
Rouxia moholensis CAS 54424

c. New combinations.
Cussia mediopunctata
Cussia mediopunctata var. matraensis
Cussia paleacea
Cussia praepaleacea

Cussia tatsunokuchiensis

ACKNOWLEDGMENTS

This investigation was financially supported by the Deutsche Forschungs-
gemeinschaft and the United States National Science Foundation. I thank W. R.
Riedel and T. Walsh of Scripps Institution of Oceanography, La Jolla for pro-
viding samples from the EMD; L. H. Burckle of Lamont-Doherty Geological
Observatory, New York for providing samples from Lamont cores; E. Seibold of
Geologisches Institut, Kiel and J. Koizumi of Institut Geological Sciences, Toyo-
noka for discussing stratigraphic problems; and Frau Schmidtmann, Frau Pro-
kopek, and Frau Enke for technical assistance. Dr. P. Rodda read the manuscript
and made many helpful suggestions.

INTRODUCTION

This study is a revision of the planktonic diatoms contained in 34 samples
from cores recovered during the Experimental Mohole Drilling near Guadalupe
Island off Baja California (Riedel e¢ a/., 1961). Age correlative samples from
cores (Lamont-Doherty Geological Observatory) and from a drill hole of the
Deep Sea Drilling Project, leg 18, Site 173 near Cape Mendocino, California
were also used in this study (chart 1, table 1).

Previous zonations of the Experimental Mohole Drilling cores have been
made on Foraminifera (Parker, 1964; Bandy and Ingle, 1970), on calcareous
nannoplankton (Martini and Bramlette, 1963; Martini, 1971), on silicoflagellates
(Martini, 1972), and on diatoms (Kanaya, 1971).

After publication of the Neogene diatom zonation of the East-Equatorial
Pacific (Burckle, 1972) and the completion of a Neogene diatom zonation of the

Vor. XXXIX] SCHRADER: DIATOM STRATIGRAPHY 519

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Cuart 1. Index map showing localities of investigated cores. DSDP 173: Deep Sea
Drilling Project, leg 18, Site 173; EMD: Experimental Mohole Drilling, Guadalupe Site;
RC-65, 208, 224: Robert Conrad cores (Lamont Doherty Geological Observatory).

TABLE 1. List of cores used in this study, with references to published descriptions.
IRDSDP—Initial reports of the Deep Sea Drilling Project; LDGO—Lamont Doherty Geo-
logical Observatory; SIO—Scripps Institution of Oceanography.

Water

Core No. Inst. Latitude Longitude Depth (m.) Author
RC-12-65 LDGO 04° 39’N 144° 58’ W 4868 Burckle 1972
RC-11-208 LDGO 05° 21’N 139° 58’ W 4920 Burckle 1972
RC-11-224 LDGO O3in327'5 122° 05.5’ W 4319 Burckle 1972
EMD-6,8,7,10 SIO 28° 59’N 117° 30’ W 3566 Riedel et al. 1961

Parker 1964

Martini 1971

Kanaya 1971
DSDP-173 SIO-DSDP 39° 57.7’'N W2SS AT? WY 2927 IRDSDP 1973

520 CALIFORNIA ACADEMY OF SCIENCES

EMD CORE LOG

[Proc. 4TH SER.

——

6
i HOLE 10
HOE 7
interval cored
9 core
mM BS ae interval recovered

Cuart 2. Core log, Experimental Mohole Drilling, Guadalupe Site.

interval recovered.

Black intervals:

VoL. XXXIX]

TABLE 2.

SCHRADER: DIATOM STRATIGRAPHY

521

Sample data for cores from the Experimental Mohole Drilling, Guadalupe
Site. Sedimentological data from Riedel et al. (1961).

Hole Core Sampled interval in cm. Sedimentological observations (Riedel ef al., 1961)
6 2 30/ 31 greenish gray, clayey siliceous-calcareous ooze
8 9 5/6 pale olive, siliceous-calcareous 00ze

140/141 pale olive, calcareous-siliceous 00ze
193/194 pale olive, calcareous-siliceous ooze
240/241 pale olive, calcareous-siliceous 00ze
290/291 olive-gray, siliceous-calcareous ooze
8 10 44/ 45 dusky yellow-green, silty siliceous ooze
100/101 dusky yellow-green, siliceous ooze
200/201 dusky yellow-green, siliceous ooze
300/301 dusky yellow-green, siliceous ooze
400/401 dusky yellow-green, siliceous ooze
8 11 30/ 31 yellowish gray, siliceous ooze
90/ 91 grayish green, siliceous-calcareous 00ze
200/201 greenish gray, siliceous 00ze
300/301 greenish gray, siliceous ooze
390/391 light olive-gray, calcareous-siliceous 00ze
8 12 94/ 95 pale olive, calcareous-siliceous ooze
203/204 grayish yellow-green, siliceous-calcareous ooze
8 13 30/ 31 dark greenish-gray, siliceous ooze
100/101 dusky yellow green, siliceous ooze
200/201 dusky yellow green, siliceous ooze
262/265 dusky yellow green, siliceous ooze
8 14 10/ 11 dark greenish gray, siliceous ooze
100/101 dark greenish gray, siliceous 00ze
400/401 dark greenish gray, siliceous ooze
8 15 35/ 36 dusky yellow green, siliceous-calcareous ooze
93/ 95 pale olive, calcareous-siliceous ooze
100/101 light greenish gray, calcareous siliceous ooze
200/201 light greenish gray, calcareous siliceous ooze
300/301 light greenish gray, calcareous siliceous ooze
504/505 pale olive, calcareous-siliceous 00ze
il 1 5/0 yellowish gray, calcareous-siliceous 00ze
41/ 42 greenish gray, siliceous ooze
7 2 O/F17 greenish gray, siliceous ooze
10 1 30/ 43 greenish gray, calcareous ooze

North Pacific based principally on a continuously cored stratigraphic hole
California (Deep Sea Drilling Project, leg 18, Site 173, Schrader, 1973a) it was
possible to prepare a revised diatom stratigraphy of the EMD which differs from
that of Kanaya (1971).

off

[Proc. 4TH SER.

ACADEMY OF SCIENCES

CALIFORNIA

CORRELATION

NEOGENE

S4INOZ

WOLVIG

I141DVd

IWIYOLVNODA

Pseudoeunotia

doliolus
praebergonii

Rhizosolenia

jouseae

Nitzschia

Thalassiosira

convexa

miocenica
porteri

Nitzschia

Nitzschia
=

SIOVIS
AINIYVW
VINYOIIIVD

Coscinodiscus

yabei

REPETIAN

DELMONTIAN

MOHNIAN
LUISIAN

ANOZddN

INOZ 1OI0VY

usd g;

maa

e| =

Ag:

VENTURIAN

17

a)

—————_> =
“

sDjuad ‘¢

nado: ‘uadGy

'
4

snwiyjNuadayuD “©

\luosiayjad"> “UODI4D] “>

INOZ ONNVN |<

Fa

10

INOZ WVAO4

HDOd3

iia

- |

°

INIDOINId

INIDOIW

j1V1

JINIDOIW JIGGIW

HDOdj OVW
‘10d ‘OVW

I

‘DO1LS1I41d

VWVALVW ISSNVD

|
“”

Lydd 119

gg wo

wn

ae

mnie Le

VoL. XXXIX] SCHRADER: DIATOM STRATIGRAPHY 523

METHODS

Samples from Lamont-Doherty Geological Observatory were taken by L. H.
Burckle; samples of the EMD were taken by T. Walsh, and samples from DSDP
Site 173 were made by the author.

Treatment of the diatom material and slide preparation followed the stan-
dardized methods of Schrader (MS.). Slides were mounted with Aroclor 4465
and key species were marked with a diamond microscopical specimen marker.
All important species were photographed through a Leitz Orthoplan, Apo Oil
92 X, 1.4 n.A. and 10 X Periplan ocular. Holotypes, paratypes, and some other
duplicates are deposited at the Department of Geology, California Academy of
Sciences. Other illustrated specimens are in the author’s collection. Additional
samples have been deposited at the Bundesanstalt fiir Bodenforschung, Hannover
(L. Benda), and at the Friedrich Hustedt Arbeitsplatz fur Diatomeenkunde,
Bremerhaven (R. Simonsen).

EpocHs AND AGE BOUNDARIES

The Upper-Middle Miocene boundary is placed within Geomagnetic Epoch
11 following Burckle (1972). This boundary is shifted by Berggren (1972) to
the upper part of Geomagnetic Epoch 10 (see chart 3).

The Miocene-Pliocene boundary is placed at the top of Geomagnetic Epoch
5 (bottom of the Reversed Gilbert Magnetic Epoch) (Burckle, 1972).

Other boundaries are defined by the range of species in the Equatorial Pacific
(Burckle, 1972) and the North Pacific (Schrader, 1973a).

Most of the Equatorial and North Pacific Zones are defined by evolutionary
events and morphotypic ranges of species (for definition of terms see Riedel and
Sanfilippo, 1971).

STRATIGRAPHIC DISTRIBUTION OF DIATOMS

The distribution of important planktonic diatom taxa is indicated in table 3.
A great effort has been made to recognize and define reworked taxa. Ranges
are plotted by symbols representing abundance and type of occurrence (alloch-
thonous or autochthonous). Abundance and preservation was generally good

Cuart 3. Correlation of the North Pacific Diatom Zonation (Schrader, 1973a) with Cali-
fornian Marine Stages (Berggren, 1972; Bandy and Ingle, 1970), the Equatorial Pacific
Diatom Zonation (Burckle, 1972), the Radiolarian Zonation (Riedel and Sanfilippo, 1971;
Moore, 1971; after Berggren, 1972), the Standard Calcareous Nannoplankton Zonation
(Martini, 1971; Bramlette and Wilcoxon, 1967; Baumann and Roth, 1969; Martini and
Worsley, 1970), the Planktonic Foraminiferal Zones (Blow, 1969; after Berggren, 1972),
the Paleomagnetic Stratigraphy (Burckle, 1972; Abdel-Monem et al., 1971), and the Radio-
metric Time Scale (Cox, 1969; Abdel-Monem et al., 1971).

[Proc. 4TH Serr.

CALIFORNIA ACADEMY OF SCIENCES

524

=~ a
re) Xu
prt eee re) 4 u
Fy) y y 3 8
H) 4y u Lot /O0¢% 8
& q fe Soe Loz/ooe - -8 - Wa
P) u u tor/oolt = S 8
g u u 8
ie Gils ) ¥ y 8
euoz a#uey yeTZAeY v u u 8
Teqek snostpoutosog ca) u u LoL /oot 8
9 y y= - u EDS PACH a AON} Wa
fe) u Lh gtd Vi" ¥ 9¢/29e~- =e =e We
COVER) i | er EC a he Gh ho Lor/oolt - - 98 - Wa
euoz eSuey TeTy1eg fp l=. <meta = OL: as BS /0€ = El = 8. =sNe
Tiezrod eTyOSz4IN Side |= ec eK Uy 8
es |S So Ses Ss Se uu 8
OTS XS Ss Pe ey eye 8
LG | as ed ==] rt Pl 8
= OC RR. i ae er a sate og /00¢ = EE = swat
OYA Tal) ea a | MP Sic Sy ce Jere SS ti wi: Ga lacy fAoyss > (AI tsp heer
SSE Men ed ete Mie O beth) SoS HS eee = u Cts BEeOSe =D ULES ee Sao
>
BOTUsOOTM STYOSZ4AIN €l SS Se a || hn: (ta Cu - } : Veen = Sle Lov /00P =O =8 = ane
=e) ee Xe (io mie mee” ee = Mh Rh OS L0£/00£ - OL - 8 - WO
<a Pee cane * Ce Oe Gee med == me Dp ear ete > Ce WT EY: Pe - Nod te 1oz2/ooe - OL - 8 - WO
su0Z eFue}; 2 eg ee Os Se! ite eS OX eS C PARE tOL/oOL - OL - 8 - Wa
TeTpzIeY Bxeauo0s ESSE EN I ES NP Cs a: Lge le ae A uy = Rf lle an = (1A Aad Sol EL ee Mie
vITSOTSSETBUY, tL Xone Xae ch Wat tS AEX TSS S> BU Ty = ell t6e/06e -6- @ - NG
a... eee SS SoU Re see ei SU Rt Be8 ye x Rig See tyz2/ove -6- 8 - Wa
euoz esuey TeT41ed fh te a (Cie (Rape ee Ti | = =u = VEL/26b = 16 = 98) = Wer
CISINIE CAMO EET SN Moe eo S a loeyo eS hy SS RRS wh > s tyL/ovL - 6 - @ - Wa
Of jos S 7 oe eto too h tee ee dis uouiy GQ) fe 92 Go Site
(ae Sa | ee sos ey Joe = 1 = 0b = Wa
5 © Joe
Oh [Si SS |S SSS SIS RS you 4 [2n/ OCS Rc EEO a
4 - + ——__—_—_—_—_—_—_—_—-+-- ~~ T
o ie LS SI i] 9 a) & Q to Es bo peyTOmMer yx
pH B34 3 a a ole |d}e ¢
tors Biers aa re) uy) e|/@)/5 | o b quepunqe = y
eis Boe lls: eee eee | Oe Cee 2 ae ee eee
» op SA aes. saa - Se ee ae eo neler ll ei|| Same She uowmopj = 9
NS ¢ SB ¢ J 2 a/p]|ol}p «
5 5 » ee a EL Wea iS ci quenbezy =
pos ns 3 ein a sen (ee oO QQ} 4 |
ot a Boe im S (if) of c 5 B} p a erel = ¥
Py « me 3 Q & Pp no 8 a wo alr
Oo = fo} ° oe Lone o °o i= Hn B ° o oO o
Se im ct ene oh Bao He) op ven @ewdwu 8S FY aololoaloa|s¢ (over tee queseaid you
a os p p =) aw pe tI Ss o o * o ° » @ r=] i=! o le} o - n =>
Be Se ec eee Sees 2 kee eee. yee 2
D> - 0 Dds <r Ce te ion 0 D He) ~~ o vs o PF Bl ple Scot en! SWNOLVIC
Qo 4 < @ ry (Cie er ct epee On an IS ye wo lo] B © 5
fa Ct » H i=] ° » © FY @ © Pp a [ele ae)
BS Ses Oe ie a aaa egg 5 ae eats edntepenp
a4 om Zi CP sae 8 a 8
a. » = A 4 5 ® Fd uTTTTId seTOYoW
xo.9 aie o Fs o a na eyuemtsedxy
mb mH 2 a 3 5 ot 7
Pp “0 a as 4 °
w + r . q
- wv io re wu. °
ty @ Fr ‘e r) 3
J » < ®
e. 3 8 i
m 8 p. e
re Q 7)
° 5
a

‘(Sajnqsnaf Uayosg KUDU puv payrja) ajwaapom—py {poos—H :sp paywjnqn] st smoywrp {0 uoNDadasarg
‘dADA—YS UOMMOI—J /JUDpUNnqD—Y ‘SD paqynjNngn] St ajdwups yova ut SmoqwIp fo aauDpuNgD 1010
‘saivagsS Aapjo paysomai—y ‘(Saivads [0 aauasdnaz0 Sspm) JuDp
~ungo—yp <(apys sag suamiags 0G uDYy} asom) Uuommor—J S(apys sag Ssuamizads QF uvyy adom) quanbas{—y {(apys sad
suaunrads maf D Kjuo) aavsA—y ‘quasaad Jou — :suorjpiaasqqn Suimopjof ay. Suisn paqvjnqgny aav saizags moywig ‘aqis adnjq
“DPONY “Suapaq aoyop jojuamiadxy ayy fo sajdwps aso wosf pasaaorad saizags moywup ruojyung fo ysvy “¢ AIAV,

VoL. XXXIX] SCHRADER: DIATOM STRATIGRAPHY 525

| a cal
o °o eo oo
oc oe] 4d 6 ac
ao | =] HO a 0
on ad eo oan mbmN
So |an 8 @ v7
: o o
°o & He “A wo o tf 3 fo
(2L6L) @TMONNE uoTzeU0Z mozRTG 3 = i A S a 5 a, 5 os
OTJTOBeT Tetrozenby 9uzy yRIA uoTZeTeI1109 8 ons Ce Cae ae
“A Id ) a dd 0
oa agn Sd od Od
od ak od og ga
Qi | soo Qed @ -d dod
NP “4 > & NP NP ov
PH |@ES Pu PH aH
ae 'coog da re) od
Zz Ay iS om ZA Zz Ay on
I
|
! I
| I
| |
8t ‘eT daqqur ‘ugadvNHOS ‘uotzeU07Z W poly DBD |H
WORETT OTFTO’T yxION UYTIA UOTZLETSeIIOD San [a ia
sTWIOJTT AAS bn Oe) t fed os oe ee otf oe oe og Oe pes oe oe ot ee en nt oe oo
Toqzeqeraid a H)efe tb 8 spt ee bs Op 8 bb 8 et cee pe cele oe ci ire oe et ot ot el ee
Bye wed “ Mpiyea pm tem bt tt bt bee oe tp et ft bt tlk at pa
'
wotusootm i fiji t totem 8b tle oe alt oem mo ie tm tm lol
"IPA BYELSQSY a mip te bt tom tp em tk tem me ty et tee tel ttm me
Toqreq etuaToOsozTyYy Seem er I ROmmMOlmkOmmlm elo 1 s}t tft t
SPEQCUTTeYTIeFzIe0 * Hl in| (erect [io arm cabal [cle ciear ts | 2s emo fuente ci
as) STSUsUTTEYIOeS * ON PU |i CP ay fre) Ce Thee Ce et el eC Cy SOR Ie at cle ete On 0
L
= eyeysnyue u Cytpe bt bt b te em 8 melt km I moe tt tt tt treme mle
™~
stTierngue steuoydeuyy
SS BoOTULOSTTeO i
BrrTyoing 4“
~ etlepata ft
in ertprloqtes “
A TtpToyutereead “a
a) aevesnol a
eotrputtTAD eStyoszzIN
(eeec0emoyeTp uoU) eT [938 eIOIIEW
eptpuetds etrerpey
wnyTnosnuto *
MNOTULO;TTeO uUntmsapoyzty
SnuIsstottdmts fs
BSTMIOFTeuNS snostTptmay

snydiomfjTod *j*o snynetmay

xazl snostpomyuyW

rare

- = not present

R
F = frequent

C = Common
A = abundant
X = reworked

Experimental
Mohole Drilling
Guadalupe Site
DIATOMS

Continued.

TABLE 3.

526 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

2 | 3 © 3
os | ca 4d 6 1
acai © 2 es as
Sree pee : ae
oO lel eo Ho oa
. om HY | a & ow 3%
(ZL6L) ATHONNA uoTzeuoZ moze TT we) is ies ag as os
OTJTOeG Tetarozenbm ayy uRTA UOTPETeIIOD ot omg ! aa: on ae
od he ° od r) “dA os
fq 'aan | sae fd On
od aK owe 0 og ad
24 |sgoo | Qain @ sr od
ne a> %o a+ Re or
SUH ie sie ee PH oH
A so @ Ee) A oa
ZA ha Om | wm Zz Om
|
|
«| |
|
. ‘ ‘ |
QL ‘Fe T daqur ‘uYaaqVHHOS ‘uoTtzeUu0Z ole-lo - lw pom i
WORST IIFTOeI YZAON UZTA UOTZPETSIIOD Sle pial paeen?
I
= i= =I =
41TS celal i 8 By Pa lt et Ii O
satnotds eZ#uods iyi pommr trot om
woneareeest | oP/P OO oOVOO|OY
aouepungy <aiule qa cee cod <|< <
suesut snjtosooutzoy =e (-ile~| [oe om —ilood |<
+ i pe
snostpoutosoo snostpopedsei) OG Oa etl OD oo OO 0
wnewomeuuto “ ait tmemimimom st tie
mnzrodvapuod mNTzRIGOTIYL 1 a 1ouot these ab nfs es
PWISBTSUOT XTINZOTBBETeUL Sits ISS Ae ies
TTasyozesn A pivje e tote dob bk to
trdn1zzsa0 ra cairo 0 0 OD OD iyo
exaauooerid u SN ee 0) A Osea Pee
PAT}EU Ma mlil) me Om mle mm bb bli
y ‘oads iu oo 2d oO OO to Ooo
enbryue ia CaN ob OO TO Da Oy
ewotTryueoxa i mili} i me 1 ee old bt time
suetdtoep a mlijot og to tye bt tom
eX9AU0D BITSOTSSETEYL zk 0 tected i) 1 Ole 0
STMIOJTABTO u CH ie Ds uP ame '
BaptotyoszzyIU emsuotsseleuyy <p|<« Lous) (Sais)
eueasnol erpeufc >] ils > fh 00
striny picid ade py jon }on OG om |oe om
septauo,tdtp *js°o ri i rt rien
eoTurostyTeo erxnoy pala} i ae f ijtot
a ee
ro = hort eele re
5 m TAT oO|\nno
a Cem Laie s N
a Pod >>> = SS
o c & © ojoJnNoMmo cojooo
H a ie] oo x Lam) HOT Oo OJM AO
i als) eh 1s) --N mt N
to oo | gs oO
i=} a oe! H o 3 z tyrpeoe De Uy
4d PP Co 8) UH ‘oO: oO)
di a 4 S Hh & O « H MN] OV OlO Sata
eon eele re
roy Py r " ‘ 1 | ee! 4
c ial ® je TN ti
oR mh 1) 0g Fe), eh
| 3} wjo|a@ o Dnojonno
d © 4 = hd
H ot &| O
@ o ol] & tyeja Fiery et ed eC) OO) Oe 0 te ae Oyo. 0
a oc 8 <
1 ORs ial Higiag = SEED ES SS SSS SSE SHEE EEE =
a = Oo] A fa] a ARPA AA AAA AAA Am RAM mw Aime ws

VoL. XXXIX] SCHRADER: DIATOM STRATIGRAPHY

EMD MULTIPLE CORRELATION

on
bo
~

an) |S) || ap
FL —s a
eS) OC <
O O N >
= Zz <<
a = Zz ~
72, || || Z4 ry he
16 a
—=— |
il lV
50 DELMON-| DELMON
TIAN TIAN
and or
ee) ame
Hullets MOHNI - ah
o AN
a UPPER ‘|
as ee OHNIA
al Lye
100 u | 2/9
= E
Oo -: 2
O mil.years 15 x
11.440.6 0 S| 8 | MOHNI- 7
= * AN
Z 18 7
O Nee LOWER
2 | = MOHNIAN
QO 12.3:0.4 o 35 5
Fa es
a |S LUISIAN =
Wilts
150 —
24 ; |
8
9 LUISIAN
fal
5
M

CuHart 4. Multiple microplankton correlation of the Experimental Mohole Drilling,
Guadalupe Site, with North Pacific Diatom Zonation (Schrader, 1973a), Silicoflagellate
Zonation (Martini, 1972), Standard Calcareous Nannoplankton Zonation (Martini, 1971),
and Californian Marine Stages (Parker, 1964; Kanaya, 1971). Potassium-Argon dates
from Dymond (1966).

528 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

EMD

1970

NPDZONES
BANDY & INGLE

50

DELMONTIAN

MOHNIAN
100

11440,6 x

12,3+0,4

LUISIAN

150 S
15,0+7 xt tyq) RELIZIAN
=
ee |

Cuart 5. Correlation of the diatom zonation of the Experimental Mohole Drilling,
Guadalupe Site, with the North Pacific Diatom Zonation (Schrader, 1973a) and with
Californian Marine Foraminiferal Stages (Bandy and Ingle, 1970). Absolute time scale
for the North Pacific Diatom Zonation from Schrader (1973a) and chart 4; dates for the
Californian Marine Stages from Bandy and Ingle (1970). Potassium-Argon dates from
Dymond (1966) and Krueger (1964).

BEST2eeeee ll

Vor. XXXIX] SCHRADER: DIATOM STRATIGRAPHY 529

except for sample EMD-10-1-30/43 cm. Additional symbols indicate the occur-
rences of sponge spicules and silt-size mineral components. The abundance of
spicules and silt grain correlates directly with the amount of reworked shallow
material including older fossil diatoms (EMD-8-9-193/194 cm.; EMD-8-9-
240/241 cm.; EMD-8-9-290/291 cm.).

Fifteen biostratigraphic units of the North Pacific Diatom Zonation and
five biostratigraphic units of the East-Equatorial Diatom Zonation can be dif-
ferentiated on the basis of ranges of planktonic diatom species (table 3).

Only cold and temperate water species are present in this Middle Miocene
to Pliocene interval. Comparable cool-water biofacies are present at DSDP
leg 18, Site 173.

BIOSTRATIGRAPHIC ZONATION OF THE EMD CORE.

1.) North Pacific Diatom Zone 10. The unit includes samples EMD-6-2-
30/31, and EMD-8-9-5/6 to EMD-8-9-240/241 cm. This unit is characterized
by the presence of Coscinodiscus temperei, Denticula hyalina var. hustedti,
Lithodesmium minusculum, Nitzschia cylindrica, N. jouseae, N. praereinholdii,
Rhizosolenia barboi, Thalassiosira convexa, and T. nativa. The zone is correla-
tive with the lower part of the Nitzschia jouseae Partial Range Zone (Burckle,
1972). Youngest investigated material of EMD is of Early Pliocene age ( ~ 4.2
m.y.). The top of the North Pacific Diatom Zone 10 was not found and should
lie higher in the section. Sample EMD-6-2-30/31 is tentatively equivalent to
EMD-8-9-140/141 cm.

2.) North Pacific Diatom Zone 11. The unit includes samples from EMD-
8-9-290/291 to EMD-8-10-44/45 cm. It is characterized by the occurrence of
Thalassiosira usatchevii, T. convexa, Lithodesmium californicum, L. minus-
culum, Denticula hustedtii, and D. hyalina var. hustedtii. This unit is correlative
with the upper part of the Thalassiosira convexa Partial Range Zone of Burckle
(1972) and is of Lower Pliocene age ( ~ 4.5—5.5 m.y.).

3.) North Pacific Diatom Zones 12-14. These three zones could not be sub-
divided in the EMD core and subdivisions on table 3 are tentative. The unit
includes samples from EMD-8-10-100/101 to EMD-8-12-94/95 cm. It is charac-
terized by the occurrence of Coscinodiscus temperei, Denticula dimorpha (lower
part of the unit), D. lauta, Lithodesmium minusculum, L. californicum, Nitzschia
praereinholdii, Rhizosolenia barboi, R. miocenica (lower part of the unit), Rouxia
californica, Synedra jouseana (lower part of the unit), T/alassionema claviformis
(lower part of the unit), and 7. nativa. This unit is correlative with the interval
from the lower part of the Thalassiosira convexa Partial Range Zone to the
Nitzschia miocenica Partial Range Zone of Burckle (1972) and is of upper Late
Miocene age ( ~ 5.5 m.y. to 7.6 m.y., top of Geomagnetic Epoch 5 to base of
Geomagnetic Epoch 7).

530 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

EMD
SEDIMENTATION
RATE

=e vy DYMOND 1966
50 O THIS PAPER

@ KRUEGER 1964

100

150

2 4 6 8 10 12 14 1% 18 20 m.y.

Vor. XXXIX] SCHRADER: DIATOM STRATIGRAPHY 531

4.) North Pacific Diatom Zone 15. The unit includes samples from EMD-
8-12-203/204 to EMD-8-13-30/31 cm. It is characterized by the occurrence of
Cussia praepaleacea, Denticula dimorpha, D. hustedtii, D. lauta, Nitzschia prae-
reinholdu, N. burcklia, Rhizosolenia barboi, R. miocenica, R. praebarboi, Synedra
jouseana, Thalassionema claviformis, and T. species A. (Burckle, 1972). This
unit is correlative with the upper part of the Nitzschia fortert Partial Range Zone
of Burckle (1972) and is of upper Late Miocene age (upper part of Geomagnetic
Epoch 8; ~ 7.8 m.y.).

5.) North Pacific Diatom Zone 16. This unit was only present in sample
EMD-8-13-100/101 cm. It differs from North Pacific Diatom Zone 15 in the
lack of Nitzschia praereinholdii and of Cussia paleacea, and in the consistent
presence of Rhizosolenia praebarbot. This zone is correlative with the lower part
of the Nitzschia porteri Partial Range Zone of Burckle (1972) and is of middle
Late Miocene age ( ~ 8.2 m.y.—middle part of Geomagnetic Epoch 8).

6.) North Pacific Diatom Zones 17-19. These zones could not be divided
and subdivisions on table 3 are tentative. The unit includes samples EM D-8-13-
262/265 to EMD-8-15-100/101 cm. It is characterized by the occurrence of
Bruniopsis mirabilis, Cussia paleacea, C. praepaleacea, Coscinodiscus yabei,
Denticula dimorpha (upper part of the unit), D. kustedtii, D. lauta, D. punctata
var. hustedtii, Mediaria splendida, Nitzschia burcklia (upper part of the unit),
N. riedelia (lower part of the unit), Rhizosolenia miocenica, R. praebarboi,
Synedra jouseana, and Craspedodiscus coscinodiscus (lower part of the unit).
This unit is correlative with the Coscinodiscus yabei Partial Range Zone of
Burckle (1972), the base of which has not been defined. It ranges from the
middle part of Geomagnetic Epoch 8 to the middle part of Geomagnetic Epoch 11
(~ 8.3 my. to 12 my.). The base of this unit represents the Late-Middle
Miocene boundary as defined by Burckle (1972), but note should be made of the
different boundary chosen by Berggren (1972) (chart 3).

7.) North Pacific Diatom Zone 20. The unit includes samples from EMD-
8-15-200/201 to EMD-8-15-300/301 cm. It is characterized by the occur-
rence of Actinocyclus cubitus, Cussia paleacea, C. praepaleacea, Coscinodiscus
vetustissimus var. javanicus, C. antiquum, C. yabei, Denticula hustedti, D. lauta,
D. nicobarica, D. punctata var. hustedtii, Mediaria splendida, Macrora stella,
Nitzschia species 2 (Schrader), N. riedelia, Rhizosolenia praebarboi, Synedra
jouseana, and Craspedodiscus coscinodiscus. This unit has not been correlated
to paleomagnetic stratigraphy, but it does correlate with NN 6 of the Standard
Calcareous Nannoplankton Zonation (Martini, 1971), and with the Corbisema
triacantha Zone of the Silicoflagellate Zonation (Martini, 1972). This Zone is of

<

CuHarT 6. Sedimentation rate of the core samples from the Experimental Mohole Drilling,
Guadalupe Site, as determined by Potassium-Argon dates (Dymond, 1966) and by diatom
zonation. Note change of sedimentation rate at 95 meters below the sea floor.

532 CALIFORNIA ACADEMY OF SCIENCES [Proc. 47TH SER.

upper Middle Miocene age. Dymond (1966) determined the absolute age of
sample EMD-8-15-86 cm. as 12.3 + 0.4 m.y. according to the Potassium-Argon
dating method. This age correlates well with the top of Zone 20 of the North
Pacific Diatom Zonation which has an approximate age of 12 my.

8.) North Pacific Diatom Zones 21-23. These zones could not be divided
precisely and the subdivisions on table 3 are tentative. The unit includes samples
from EMD-8-15-504/505 cm. to EMD-7-2-41/42 cm. It is characterized by the
occurrence of Actinocyclus ellipticus var. moronensis, Coscinodiscus lewisianus,
Cussia paleacea, C. praepaleacea, Coscinodiscus yabei, Denticula hustedtu, D.
lauta, D. nicobarica, Mediaria splendida, Nitzschia riedelia, N. species 2
(Schrader), and Rhizosolenia praebarboi. The unit has not been correlated to
paleomagnetic stratigraphy, but it is correlative with NN 6 of the Standard Nan-
noplankton Zonation (Martini, 1971), and is of Middle Miocene age.

9.) North Pacific Diatom Zone 24. This unit was found at EMD-7-2-0/17
cm. and is distinguished from the upper unit by the absence of Bruniopsis mirab-
ilis, Denticula hustedtii, Mediaria splendida, Nitzschia riedelia, and other
species. The unit has not been correlated to paleomagnetic stratigraphy, but it
is correlative with NN 6 of the Standard Nannoplankton Zonation (Martini,
1971) and with the Dictyocha octacantha Silicoflagellate Zone of Martini
(1972). Itis of Middle Miocene age.

SPECIES FOUND IN EXPERIMENTAL MOHOLE DRILLING SAMPLES

Genera are arranged alphabetically in the list, and species are arranged alpha-
betically within each genus.

Species and varieties have been identified following descriptions of Hustedt
(1930, 1959), Sheshukova-Poretzkaya (1967), Schrader (1973a), and Hanna
(1926, 1930, 1932).

Descriptions and discussions are given for species not adequately treated
in the literature.

Species which could not be positively assigned to a known species are num-
bered (if only one or two individuals were found) or described as new and
illustrated.

Holotypes and paratypes will be deposited in the California Academy of
Sciences diatom collection.

Genus Actinocyclus Ehrenberg
Actinocyclus cubitus Hanna and Grant.

Description. Hanna and Grant, 1926, p. 118, pl. 11, fig. 3.

Actinocyclus ehrenbergi Ralfs.

DEScRIPTION. Hustedt, 1930, pp. 525-532, numerous figures.
Remarks. No subdivisions of this species have been made.

VoL. XXXIX] SCHRADER: DIATOM STRATIGRAPHY 533

Taste 4. List of samples and corresponding Department of Geology catalog numbers,
California Academy of Sciences, San Francisco. A split of the cleaned fraction of each sample
is stored at the California Academy of Sciences.

Sample Number CAS Number
EMD-6-2-30/31 54427
EMD-10-1-30/43 54428
EMD-8-9-5/6 54429
EMD-8-9-140/141 54430
EMD-8-9-193/194 54431
EMD-8-9-240/241 54432
EMD-8-9-290/291 54433
EMD-8-10-44/45 54434
EMD-8-10-100/101 54435
EMD-8-10-200/201 54436
EMD-8-10-300/301 54437
EMD-8-10-400/401 54438
EMD-8-11-30/31 54439
EMD-8-11-90/91 54440
EMD-8-11-200/201 54441
EMD-8-11-300/301 54442
EMD-8-11-390/391 54443
EMD-8-12-94/95 54444
EMD-8-12-203/204 54445
EMD-8-13-30/31 54446
EMD-8-13-100/101 54447
EMD-8-13-262/265 54448
EMD-8-14-10/11 54449
EMD-8-14-100/101 54450
EMD-8-14-400/401 54451
EMD-8-15-35/36 54452
EMD-8-15-93/95 54453
EMD-8-15-100/101 54454
EMD-8-15-200/201 54455
EMD-8-15-300/301 54456
EMD-8-15-504/505 54457
EMD-7-1-5/6 54458
EMD-7-1-41/42 54459
EMD-7-2-0/17 54460

Actinocyclus ellipticus Grunow.
DEscrRIPTION. Hustedt, 1930, p. 533, fig. 303.

Actinocyclus ellipticus var. moronensis (Deby) Kolbe.
DEscrIPTION. Kolbe, 1954, p. 21, pl. 3, figs. 29-30.

534 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Actinocyclus ingens Rattray.

DESCRIPTION. Kanaya, 1971, p. 554, numerous figures.

Actinocyclus okhotensis Jouse.

DESCRIPTION. Koizumi, 1968, p. 208, pl. 32, figs. 7-10; Donahue, 1970,
p. 135, pl. 2, figs. 2-5.

Genus Asterolampra Ehrenberg
Asterolampra grevillei (Wallich) Greville.
DESCRIPTION. Hustedt, 1930, pp. 489-491, fig. 274.

Asterolampra marylandica Ehrenberg.
DEscriPTION. Hustedt, 1930, pp. 485-487, fig. 271.

Genus Biddulphia Gray
Biddulphia moholensis Schrader, new species.

DescriPTION. Valve lanceolate-rhombic with produced bluntly rounded
apices, 60 » long, 16» in the middle part, 4 » wide at the apices. Valve with
radially arranged rows of punctae, 15 in 10 »; punctae in quincunx. Middle part
slightly lifted from the valve surface with three spines and without any central
area. Valves with stronger solitary spines on both sides of the median apical line
near the valve mantle and with two to three solitary spines at both apices near
the valve mantle.

Discussion. No similar species was found mentioned in the literature. This
species was placed within the genus Biddulphia because of the shape of the valve
and the arrangement of the spines.

Hototype. Figure 4: 3-4, from EMD-8-9-290/291 cm. California Academy
of Sciences, Department of Geology no. 54409 (diatom collection).

DERIVATION OF NAME. This species is named ‘moholensis’ in memory of the
Experimental Mohole Drilling.

OccuRRENCE. Found only in the Neogene section off Baja California, in a
sample in North Pacific Diatom Zone 11.

Genus Bruniopsis (Tempere) Karsten

Bruniopsis mirabilis (Brun) Karsten.

DEscrIPTION. Kolbe, 1954, p. 24, pl. 4, fig. 44; Kanaya, 1971, p. 555.

Genus Cladogramma Ehrenberg
Cladogramma dubium Lohmann.

DescripTIon. Lohmann, 1948, p. 168, pl. 9, fig. 5; Sheshukova-Poretzkaya,
1967, p. 192, pl. 24, fig. 6, pl. 29, fig. 4.

VoL. XXXIX] SCHRADER: DIATOM STRATIGRAPHY 535

Genus Coscinodiscus Ehrenberg
Coscinodiscus antiquus (Grunow) Rattray.
DESCRIPTION. Grunow, 1884, p. 84, pl. 4 (D), fig. 24.

Coscinodiscus moholensis Schrader, new species.

DEscriPTION. Valve circular, 38 » in diameter, surface with a tangential
plication, concave on one side, convex on the other. Central area absent. Areolae
on the concave plication are round, on the convex plication they are polygonal,
radially arranged in a complete meshwork over the valve surface. Areolae 6—7
in 10», of about the same size all over the valve. Areolae forming fascicles.
Margin clearly defined with a secondary plication forming triangulate uplifted
areas 5 » wide. On top of each of these areas is one elongated areola. Margin
well defined and separated from the valve structure by a small hyaline area.
Margin 1-2 » wide, chambered, radially striated.

Discussion. This species differs from all other species of the Coscinodiscus
plicatus group in the formation of the triangulate uplifted marginal areas. No
reference to such a form was found in the literature.

DERIVATION OF NAME. Dedicated to the memory of the Experimental Mohole
Drilling.

HoLotyPe. Figure 3: 3-5 from EMD-8-13-100/101 cm. California Acad-
emy of Sciences, Department of Geology no. 54410 (diatom collection).

OccURRENCE. Found in the Neogene Section off Baja California, from a
sample in North Pacific Diatom Zone 16.

Coscinodiscus endoi Kanaya.
(Figure 3: 11-12.)

DESCRIPTION. Kanaya, 1959, pp. 76-77, pl. 3, figs. 8-11; Koizumi, 1968,
pazii, pl. 32, figs. 21-22.

Coscinodiscus lewisianus Greville.
DEscripTIon. Kanaya, 1971, p. 555, pl. 40.5, figs. 4-6.

Coscinodiscus marginatus Ehrenberg.

DESCRIPTION. Hustedt, 1930, pp. 416-418, fig. 223.
Discussion. No attempt has been made here to subdivide C. marginatus
and C. marginatus var. fossilis Jouse.

Coscinodiscus nodulifer A. Schmidt.
DEscripTION. Hustedt, 1930, pp. 426-427, fig. 229.

Coscinodiscus plicatus Grunow.

DEscripTION. Grunow, 1884, p. 86, pl. 3, figs. 10, 27; Kolbe, 1954, pp.
34-35 (no illustration) ; Schrader, 1973a, p. 703, pl. 6, fig. 23.

536 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

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Ficure 1. Magnification 1000 xX, Leitz-Orthoplan, Apo Oil 92/1.40 n.A. 1. EMD-8-
10-100/101 cm. Cussia moholensis Schrader, new species; 2. EMD-8-10-100/101 cm.
C. moholensis, California Academy of Sciences, Department of Geology no. 54414; 3. EMD-
8-9-290/291 cm., C. moholensis (type); 4. EMD-8-9-290/291 cm., C. moholensis, California
Academy of Sciences, Department of Geology no. 54413; 5. EMD-8-15-100/101 cm.,
C. lancettula Schrader, new species, California Academy of Sciences, Department of Geology
no. 54112; 6. EMD-8-11-300/301 cm., C. tatsunokuchiensis (Koizumi), new combination;
7. EMD-8-11-300/301 cm., C. tatsunokuchiensis; 8. EMD-7-1-5/6 cm., C. praepaleacea
(Schrader), new combination; 9. EMD-8-15-100/101 cm., C. lancettula (type) ; 10. EMD-8-

VoL. XXXIX] SCHRADER: DIATOM STRATIGRAPHY 537

Coscinodiscus symbolophorus Grunow.
(Figure 1: 30.)

DeEscrIPTION. Hustedt, 1930, pp. 396-398, fig. 208 (as Coscinodiscus stel-
laris var. symbolophora (Grunow) Jorgensen); Schrader, 1973a, p. 703, pl. 22,
figs. 8-9.

Coscinodiscus temperei Brun.

(Figure 2: 29-33.)
DeEscriPTION. Kanaya, 1959, p. 84, pl. 4, fig. 8.
Discussion. See Schrader, 1973a, p. 704.

Coscinodiscus vetustissimus Pantocsek.
DEscrIPTION. Hustedt, 1930, p. 412, fig. 220.

Coscinodiscus vetustissimus var. javanicus Reinhold.

DESCRIPTION. Reinhold, 1937, p. 102, pl. 8, figs. 7-8; Kanaya, 1971, p. 555,
pl4orzZ: fig. 1.

Coscinodiscus yabei Kanaya.
(Figure 3: 6-8.)

DEscrRIPTION. Kanaya, 1959, pp. 86-87, pl. 5, figs. 5-9; Schrader, 1973a,
p. 704, pl. 6, figs. 1-6.

Coscinodiscus species 1 Schrader.

DESCRIPTION. Valve circular, 24 in diameter, surface with a tangential
plication, concave on one side, convex on the other. Central area absent. Areolae
round, arranged in radial rows forming a complete network over the entire valve.
Areolae 8 in 10» and of the same size all over the valve surface. Secondary
spiral structure of areolae well developed, interrupted by the axis of the plica-

<

15-100/101 cm., C. lancettula, California Academy of Sciences, Department of Geology no.
54411; 11. EMD-8-15-93/94 cm., C. paleacea (Grunow), new combination; 12. EMD-8-15-
93/94 cm., C. paleacea; 13. EMD-8-15-504/505 cm., C. paleacea; 14. EMD-8-15-504/505
cm., C. paleacea; 15. EMD-8-9-240/241 cm., Denticula dimorpha Schrader; 16. EMD-8-
15-504/505 cm., D. lauta Bailey; 17. EMD-8-9-516 cm., D. lauta; 18. EMD-8-10-100/101
cm., Cussia species 1 Schrader; 19. EMD-8-10-100/101 cm., Cussia species 1 Schrader;
20. EMD-8-9-140/141 cm., Denticula hustedtii Simonsen and Kanaya; 21. EMD-8-9-
140/141 cm., D. hustedtii; 22. EMD-8-15-93/94 cm., D. punctata var. hustedtii Schrader;
23. EMD-8-15-504/505 cm., D. punctata var. hustedtii; 24. EMD-8-9-140/141 cm., D.
punctata var. hustedtii; 25. EMD-7-2-0/17 cm., D. hyalina Schrader; 26. EMD-7-2-0/17
cm., D. nicobarica Grunow; 27. EMD-7-1-5/6 cm., D. nicobarica; 28. EMD-8-15-300/301
cm., Triceratium antiquum Pantocsek; 29. EMD-8-15-35/36 cm., T. cinnamomeum Greville;
30. EMD-8-15-200/201 cm., Coscinodiscus symbolophorus Grunow; 31. EMD-8-11-30/31
cm., Thalassiosira oestrupii (Ostenfeld) Proshkina-Lavrenko.

538 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

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717 18 19 20 “ae

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21

Ficure 2. Magnification 1000 x, Leitz-Orthoplan, Apo Oil 92/140 n.A. 1. EMD-
8-11-30/31 cm., Thalassiosira oestruppi (Ostenfeld) Proshkina-Lavrenko; 2. EMD-8-11-
30/31 cm., T. oestruppi; 3. RC-12-65-1915/1917.5 cm., T. species A. Burckle; 4. EMD-8-
13-30/31 cm., T. species A. Burckle; 5. EMD-8-9-140/141 cm., T. convexa Mukhina;
6. EMD-8-9-240/241 cm., T. convexa Mukhina; 7. RC-12-65-1105/1107.5 cm., T. prae-
convexa Burckle; 8. RC-12-65-1105/1107.5 cm., T. praeconvexa; 9. EMD-8-10-44/45 cm.,
T. praeconvexa; 10. EMD-8-10-44/45 cm., T. praeconvexa; 11. EMD-8-9-240/241 cm.,
T. nativa Sheshukova-Poretzkaya; 12. EMD-8-9-140/141 cm., T. nativa; 13. EMD-8-9-

Vor. XXXIX] SCHRADER: DIATOM STRATIGRAPHY 539

tion. Margin clearly defined, not separated from the meshwork, 2—2.5 » wide,
radially striated.

Discussion. This species differs from C. yabei Kanaya in the round areolae,
and in the striated valve margin. It is close to Coscinodiscus flexuosus Brun but
differs in the nonseparated valve margin, and the nonchambered margin (see
Schrader, 1973a, p. 702). Only one specimen was observed.

FIGURED SPECIMEN. Figure 4: 13-14 from EMD-8-9-290/291 cm., a sample
from North Pacific Diatom Zone 11.

Coscinodiscus species 2 Schrader.

DescriPTION. Valve circular, 15-16 » in diameter, surface with a tangential
plication, concave on one side, convex on the other. Central area absent. Areolae
round on the concave plication, polygonal on the convex plication, arranged
in radial rows, forming a complete network over the entire valve. Areolae de-
creasing in size slightly towards the margin, approximately 11 in 10. Sec-
ondary spiral structure of the areolae well developed. Margin clearly defined,
not separated from the meshwork, 2 p. wide, radially striated, 16 in 10 », margin
not chambered.

Discussion. This species is close to the above mentioned species but differs
in the finer structure and the more finely striated margin. No similar species
have been found in the literature. Only one specimen was observed.

FIGURED SPECIMEN. Figure 2: 34-35 from EMD-8-9-193/194 cm., a sample
belonging to North Pacific Diatom Zone 10.

Coscinodiscus species 3 Schrader.

DeEscrIPTION. Valve circular, 21—22 » in diameter, surface with a tangential
plication, concave on one side, convex on the other. Central area absent. Areolae
round on the concave plication, polygonal on the convex plication; areolae de-
creasing in size slightly towards the margin, 10 in 10 », arranged in radial rows,
forming a complete network over the entire valve. Secondary radial rows present

<

140/141 cm., T. nativa; 14. EMD-8-9-193/194 cm., T. nativa; 15. EMD-8-9-240/241 cm.,
T. nativa; 16. EMD-8-9-5/6 cm., T. nativa; 17. EMD-8-11-300/301 cm., T. nativa;
18. EMD-8-9-193/194 cm., T. nativa; 19. EMD-8-9-193/194 cm., T. nativa; 20. EMD-
8-9-193/194 cm., T. nativa; 21. EMD-8-11-390/391 cm., T. nativa; 22. EMD-8-9-390/391
cm., 7. species 2 Schrader; 23. EMD-8-9-390/391 cm., T. species 2; 24. EMD-8-9-390/391
cm., 7. species 2; 25. EMD-8-9-5/6 cm., T. nativa Sheshukova-Poretzkaya; 26. EMD-8-9-
5/6 cm., T. nativa; 27. EMD-8-9-193/194 cm., T. species 2 Schrader; 28. EMD-8-15-
504/505 cm., 7. species 2; 29. EMD-8-9-5/6 cm., Coscinodiscus temperei Brun; 30. EMD-
8-9-5/6 cm., C. temperei; 31. EMD-8-9-5/6 cm., C. temperei; 32. EMD-8-10-44/45 cm.,
C. temperei; 33. EMD-8-10-44/45 cm., C. temperei; 34. EMD-8-9-193/194 cm., C. species
2 Schrader; 35. EMD-8-9-193/194 cm., C. species 2; 36. EMD-7-2-0/17 cm., C. species 3
Schrader; 37. EMD-7-2-0/17 cm., C. species 3; 38. EMD-7-2-0/17 cm., C. species 3;
39. EMD-8-9-290/291 cm., C. yabei Kanaya; 40. EMD-8-9-290/291 cm., C. yabez.

540 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Ficure 3. Magnification 1000 x, Leitz-Orthoplan, Apo Oil 92/1.40 n.A. 1. EMD-8-12-
94/95 cm., Thalassiosira usatchevii Jouse; 2. EMD-8-12-94/95 cm., T. usatchevii; 3. EMD-
8-13-100/101 cm., Coscinodiscus moholensis Schrader, new species; 4. EMD-8-13-100/101
cm., C. moholensis; 5. EMD-8-13-100/101 cm., C. moholensis; California Academy of
Sciences, Department of Geology no. 54410; 6. RC-12-65-2142.5/2145 cm., Coscinodiscus
yabet Kanaya; 7. RC-12-65-2142.5/2145 cm., C. yabei; 8. RC-12-65-2142.5/2145 cm.,
C. yabei; 9. EMD-7-1-5/6 cm., Liostephania species 1 Schrader; 10. EMD-7-1-5/6 cm.,
L. species 1; 11. EMD-8-9-140/141 cm., Coscinodiscus endoi Kanaya; 12. EMD-8-9-
140/141 cm., C. endoi; 13. EMD-8-15-504/505 cm., C. vetwustissimus Pantocsek.

Vor. XXXIX] SCHRADER: DIATOM STRATIGRAPHY 541

with an interstitial mesh. Secondary spiral structure of the areolae well devel-
oped. Margin clearly defined, not separated from the meshwork, radially striated
12 in 10». Margin chambered with two areolae between each chamber.

Discussion. This species is close to Coscinodiscus flexuosus Brun but dif-
fers in the absence of a hyaline area between the valve structure and the marginal
structure. Final taxonomic decision can only be made if more individuals are
found. Only one specimen was observed.

FIGURED SPECIMEN. Figure 2: 36-38 from EMD-7-2-0/17 cm., a sample
from North Pacific Diatom Zone 24.

Genus Craspedodiscus Ehrenberg
Craspedodiscus coscinodiscus Ehrenberg.

DescripTION. Kolbe, 1954, p. 36, pl. 1, fig. 4; Kanaya, 1971, p. 555, pl. 40.4,
figs. 1-3.

Genus Cussia Schrader, new genus

Coscinodiscus e.p. RATTRAY, 1890, p. 597, fig. 149.

Stoschia (?) e.p. GRUNOW in Van Heurck, 1883, pl. 128, fig. 6.
Coscinodiscus e.p. SCHRADER, 1973a. (C. paleaceus, C. praepaleaceus).
Rhaphoneis e.p. Hajos, 1968, p. 143, pl. XLI, figs. 16-27.

Rhaphoneis e.p. Koizumi, 1972, p. 349, pl. 42, figs. 3-4.

DescrIPTION. Cells solitary, free. Valves elliptical-lanceolate with a sym-
metrical and/or asymmetrical transapical axis, flat. Valve surface usually either
with round areolae or with transapical costae forming a pennate-like surface
ornamentation with a zigzag apical costa forming a pseudoraphe-like morpho-
logical band; in between the costae are hyaline intercostal membranes. Valve
center not furnished with a central area. Spines or apiculi absent. Valve margin
usually narrow with a coarse striation.

TYPE SPECIES. Stoschia (?) paleacea Grunow in Van Heurck, 1883, pl. 128,
fig. 6.

DisTRIBUTION. Widely distributed in Neogene marine sediments (Mediter-
ranean Sea and Pacific, Atlantic, and Indian oceans).

Discussion. Stoschia Janisch was not validly published by Janisch, but the
description was distributed to diatomists privately by the author (Janisch, C.:
The diatoms of the ‘‘Gazelle” Expedition, 17 plates with manuscript index, no
date). Rattray, 1890, p. 548, combined Stoschia with Coscinodiscus and com-
bined the species Stoschia admirabilis Janisch (circa 1888) with Coscinodiscus
reniformis Castracane (1886, p. 160, pl. 12, fig. 12). Grunow (1883 im Van
Heurck) established the species Stoschia paleacea; this species was newly com-
bined by Rattray (1890, p. 597, fig. 149) and named Coscinodiscus paleaceus
(Grunow) Rattray. As soon as the evolution of the genus under discussion was
clarified, the taxonomic position of some other species could be detected which
have been described under Rhaphoneis by Hajos (1968) and Koizumi (1972).

542 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

DERIVATION OF NAME. Dedicated to the scientific vessel Cuss J, which drilled
the Experimental Mohole Drilling.

Cussia lancettula Schrader, new species.

DEscriPTIon. Valves elliptical-lanceolate with acute rounded ends. 24-70 pu
long, 4-7 » wide. Transapical axis symmetrical. Valve surface flat with trans-
apical costae, 10-12 in 10 », slightly curved near the apices. Transapical costae
not divided in the middle part, divided and in decussate arrangement towards the
apices, connected in the middle by an apical zigzag line which simulates a pseudo-
raphe. Intercostal membranes homogenous even in oblique light. Margin not
separated from the valve structure.

Discussion. This species differs from Cussia praepaleacea in that the margin
is not separated, and the transapical structure is simple.

DERIVATION OF NAME. From the Latin word ‘lancea’ meaning ‘lance.’

Hototype. Figure 1: 9-10 from EMD-8-15-100/101 cm., a sample from
North Pacific Diatom Zone 19. California Academy of Sciences, Department of
Geology no. 54411 (diatom collection).

PARATYPE. Figure 1: 5. California Academy of Sciences, Department of
Geology no. 54412 (diatom collection).

Cussia mediopunctata (Hajos), new combination.
Raphoneis mediopunctata Hajos, 1968, p. 143, pl. XLI, figs. 16-27.

Discussion. The illustrations of Hajos verify this new combination. The
girdle view is most characteristic, as are the other mentioned morphological
features. Unfortunately the micrographs do not show clearly if as stated “in der
Langsmittellinie der Schalen zieht sich eine Punktreihe.” This species is close to
Cussia moholensis but differs in the presence of round areolae.

Cussia mediopunctata var. matraensis (Hajos), new combination.

Raphoneis mediopunctata var. matraensis Hajos, 1968, p. 144, pl. XLII, figs. 1-5.

Cussia moholensis Schrader, new species.

DEscrIPTION. Valves elliptical-lanceolate with acute apices, 76-85 » long,
9-10 » wide. Transapical axis symmetrical. Valve surface flat with marginal
transapical costae 7 in 10 » and extending 2-3 » into the valve surface. In be-
tween the marginal transapical costae are round areolae. Valve surface with
scattered, sometimes transapically orientated costae, which form a pseudoraphe-
like medium apical zigzag line. Intercostal membranes homogenous even in
oblique light.

Discussion. This species differs from Cussia lancettula in having scattered
medium costae and wider valves.

DERIVATION OF NAME. Dedicated to the memory of the Experimental Mohole
Drilling.

VoL. XXXIX] SCHRADER: DIATOM STRATIGRAPHY 543

Hototype. Figure 1: 3-4 from EMD-8-9-290/291 cm., a sample from North
Pacific Diatom Zone 11. California Academy of Sciences, Department of Geol-
ogy no. 54413 (diatom collection).

PARATYPE. Figure 1: 1-2. California Academy of Sciences, Department of
Geology no. 54414 (diatom collection).

Cussia paleacea (Grunow), new combination.
(Figure 1: 11-14.)

Stoschia paleacea GRUNOW in Van Heurck, 1883, pl. 128, fig. 6.
Coscinodiscus paleaceus (Grunow) RATTRAY, 1890, p. 597, fig. 149.
Coscinodiscus paleaceus (Grunow) Ratrray, in Schrader, 1973a, p. 703, pl. 3, figs. 10-12.

DEscripTION. Kolbe, 1954, p. 34, pl. 3, fig. 32.

Cussia praepaleacea (Schrader), new combination.
(Figure 1: 8.)

Coscinodiscus praepaleaceus SCHRADER, 1973a, p. 703, pl. 3, figs. 1-9.

DescripTION. Schrader, 1973a, p. 703, pl. 3, figs. 1-9.

Cussia tatsunokuchiensis (Koizumi), new combination.
(Figure 1: 6-7.)

Raphoneis tatsunokuchiensis KoizuMt, 1972, p. 349, pl. 42, figs. 3-4.

DEscriPTION. Koizumi, 1972, p. 349, pl. 42, figs. 3-4.

Cussia species 1 Schrader.

Discussion. Only two fragments were found with coarsely punctated trans-
apical lines, 7 in 10 », the punctae forming straight-apical lines. Middle structure
separated from the marginal line of punctae by a hyaline area parallel to the mar-
gin. No mention of similar species has been found in the literature.

FIGURED SPECIMEN. Figure 1: 18-19.

Genus Denticula Kiitzing

Denticula dimorpha Schrader.
(Figure 1: 15.)

DescripTION. Schrader, 1973a, p. 704, pl. 1, fig. 42.

Denticula hustedtii Simonsen and Kanaya.
(Figure 1: 20-21.)

DESCRIPTION. Simonsen and Kanaya, 1961, p. 501, pl. 1, figs. 19-25, pl. 2,
figs. 36-47.

544 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Ficure 4. Magnification 1000 x, Leitz-Orthoplan, Apo Oil 92/1.40 n.A. 1. EMD-8-9-
290/291 cm., Coscinodiscus plicatus Grun; 2. EMD-8-9-290/291 cm., C. plicatus; 3. EMD-
8-9-290/291 cm., Biddulphia moholensis Schrader, new species; 4. EMD-8-9-290/291 cm.,
B. moholensis, California Academy of Sciences, Department of Geology no. 54409; 5. EMD-
8-10-100/101 cm., Coscinodiscus yabei Kanaya; 6. EMD-8-10-100/101 cm., C. yabei;
7. EMD-8-10-100/101 cm., C. yabei; 8. EMD-8-10-100/101 cm., C. yabei; 9. EMD-
8-10-100/101 cm., C. yabei; 10. EMD-8-10-100/101 cm., C. yabei; 11. EMD-8-10-300/301
cm., C. plicatus Grunow; 12. EMD-8-10-300/301 cm., C. plicatus; 13. EMD-8-9-290/291
cm., C. species 1 Schrader; 14. EMD-8-9-290/291 cm., C. species 1.

VoL. XXXIX] SCHRADER: DIATOM STRATIGRAPHY 545

Denticula hyalina Schrader.
(Figure 1: 25.)

DEscrRIPTION. Schrader, 1973a, pp. 704-705, pl. 1, figs. 12-20, 22.

Denticula hyalina var. hustedtii Schrader.

DESCRIPTION. Schrader, 1973b, p. 418, pl. 1, fig. 9.

Denticula lauta Bailey.
(Figure 1: 16-17.)

DEscRIPTION. Simonsen and Kanaya, 1961, pp. 500-501, pl. 1, figs. 1-8.

Denticula nicobarica Grunow.
(Figure 1: 26-27.)

DESCRIPTION. Simonsen and Kanaya, 1961, p. 503, pl. 1, figs. 11-13.

Denticula punctata Schrader.
DescripTIon. Schrader, 1973a, p. 705, pl. 1, figs. 27-28, 25-26.

Denticula punctata var. hustedtii Schrader.
(Figure 1: 22-24.)

DescriPTION. Schrader, 1973a, p. 705, pl. 1, figs. 23-24.

Genus Ethmodiscus Castracane
Ethmodiscus rex (Rattray) Hendey.
DEscripTIon. Hendey, 1953, pp. 51-57, pl. 1, figs. 1-6, pl. 2, figs. 1-3.

Genus Hemiaulus Ehrenberg
Hemiaulus cf. H. polymorphus Grunow.

DESCRIPTION. Hustedt, 1930, pp. 880-881, figs. 525-526.
Discussion. The specimens which are tentatively identified here as Hemi-
aulus polymorphus differ from the original description in the lack of pseudoseptae.

Genus Hemidiscus Wallich
Hemidiscus cuneiformis Wallich.

DEscriPTION. Hustedt, 1930, pp. 904-907, fig. 542.

Hemidiscus simplicissimus Hanna and Grant.

DeEscripTIon. Hanna and Grant, 1926, p. 147, pl. 16, fig. 13; Schrader,
1973a, p. 706, pl. 24, figs. 12-13.

546 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Genus Liostephania Ehrenberg
Liostephania species 1 Schrader.

Discussion. Flat silicified disks belonging to this genus were found occa-
sionally with a moderately well preserved diatom association. Hanna and Brigger
(1970) pointed out that these disks are silicified remains of inner chambers of
either Asterolampra or Asteromphalus.

FIGURED SPECIMEN. Figure 3: 9-10.

Genus Lithodesmium Ehrenberg
Lithodesmium californicum Grunow.

DEscripTION. Schrader, 1973a, p. 706, pl. 12, figs. 11, 20.

Lithodesmium minusculum Grunow.

DescriPTIon. Schrader, 1973a, p. 706, pl. 12, fig. 7.

Genus Macrora Hanna (not a diatom genus, taxonomic position uncertain)
Macrora stella (Azpeitia) Hanna.

DEscrIPTION. Hanna, 1932, p. 196, pl. 12, fig. 7.

Genus Mediaria Sheshukova-Poretzkaya
Mediaria splendida Sheshukova-Poretzkaya.
DeEscripTION. Sheshukova-Poretzkaya, 1967, p. 306, pl. 47, fig. 14, pl. 48,
fig. 8.
Genus Melosira Agardh
Melosira sulcata (Ehrenberg) Kiitzing.
Description. Hustedt, 1930, pp. 276-278, figs. 118-120.

Genus Nitzschia Hassal
Nitzschia californica Schrader.
DESCRIPTION. Schrader, 1973a, p. 707, pl. 26, fig. 6; pl. 5; aieueiae
Nitzschia cylindrica Burckle.
(Figure 5: 23-30.)
DeEscripTIoNn. Burckle, 1972, pp. 239-240, pl. 2, figs. 1-6.

Nitzschia burcklia Schrader, new species.

DeEscrIPTION. Valve elliptical with slightly convex margins and broadly
rounded apices. 30-33 » long, 5 » wide. Transapical costae 11-12 in 10. In

Vor. XXXIX] SCHRADER: DIATOM STRATIGRAPHY 547

between the middle two costae one ‘secondary’ costa is inserted which is small
and derives at the central nodule. Intercostal membranes hyaline, even in oblique
light. Transapical costae and intercostal membranes straight in transapical
direction, reaching the well developed apical field. Apical field with 2—3 apical
costae. Keel well developed, keel punctae 11 in 10». Central nodule present,
raphe marginal.

Discussion. This species differs from the allied species N. riedelia, N.
rolandii, N. kanayensis in the presence of the ‘secondary transapical costae’ in
the middle of the valves.

DERIVATION OF NAME. Dedicated to Dr. Lloyd Burckle of Lamont-Doherty
Geological Observatory, New York.

HototyPe. Figure 6: 29 from EMD-8-9-240/241 cm., a sample from North
Pacific Diatom Zone 10. California Academy of Sciences, Department of Geology
no. 54415 (diatom collection).

PARATYPE. Figure 6: 30. California Academy of Sciences, Department of
Geology no. 54416 (diatom collection).

Nitzschia cf. N. heteropolica Schrader.
(Figure 5: 42.)
DescripTION. Schrader, 1973a, p. 707, pl. 26, figs. 1-2.

Discussion. Only one fragment was found in the EMD samples and thus
a positive identification could not be made.

Nitzschia invisa Schrader.
(Figure 6: 31.)

DescripTIoNn. Schrader, 1973a, p. 707, pl. 26, fig. 5.

Nitzschia jouseae Burckle.
DescripTIon. Burckle, 1972, p. 240, pl. 2, figs. 17-21.

Nitzschia kanayensis Schrader, new species.

DescripPTIoN. Valves elliptical with slightly convex margins, 18-25 » long,
4—4.5 » wide, apices broadly rounded. Transapical costae 13-16 in 10 p, straight
transapical in the middle part of the valve, slightly convex towards the apices.
Apical field well developed with 2—3 apical costae. Intercostal membranes hya-
line even in oblique light. Raphe marginal, keel indistinct; keel punctae hardly
visible, about 17 in 10 » (compare fig. 6: 28).

Discussion. This species is close to Nitzschia rolandii and N. riedelia but
differs in the finer structure and the apical fields.

DERIVATION OF NAME. Dedicated to Dr. Taro Kanaya.

HoLotypPe. Figure 6: 23 from EMD-8-15-200/201 cm., a sample from North

548 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Pacific Diatom Zone 20. California Academy of Sciences, Department of Geology
no. 54417 (diatom collection).

PARATYPES. Figure 6: 25, 28. California Academy of Sciences, Department of
Geology nos. 54418, 54426 (diatom collection).

OccuRRENCE. Found only in the Neogene section off Baja California and
at DSDP Site 173.

Nitzschia miocenica Burckle.
(Figure 5: 17-22.)

DescripTIon. Burckle, 1972, pp. 240-241, pl. 2, figs. 10-15.

Nitzschia moholensis Schrader, new species.

DEscrRIPTION. Valves elliptical with convex margins and slightly capitate
apices. Apices heteromorph. 26 » long, 5 » wide in the middle. Transapical costae
10 in 10 p, straight in transapical direction in the middle of the valves, slightly
curved towards the apices. Intercostal membranes with two rows of punctae (20
in 10 »), punctae in decussate arrangement 10 in 10 ». Raphe marginal, central
nodule absent, keel indistinct; keel punctae about 10 in 10 p.

Discussion. No similar species has been found mentioned in the literature.

DERIVATION OF NAME. Dedicated to the memory of the Experimental Mohole
Drilling.

HototyrPe. Figure 6: 10-11 from EMD-8-11-300/301 cm., a sample from
North Pacific Diatom Zone 14. California Academy of Sciences, Department of
Geology no. 54419 (diatom collection).

OccuRRENCE. Found only in the Neogene section of Baja California.

Nitzschia porteri Burckle non Frenguelli.
(Figure 5: 31-32.)

DeEscripTIon. Burckle, 1972, no description, pl. 2, fig. 16.

Discussion. Frenguelli, 1949, p. 116, pl. 1, fig. 33-34 described a species
with “elongato-ovalibus” valves and “striis transversis delicatis, 12-13 in 10 p.”
Burckle’s figured specimen has elliptical valves with parallel margins and distinct
transapical intercostal membranes with one row of punctae and transapical costae
which form a slight angle with the transapical direction. I have not yet changed
the name because of lack of the original Frenguelli material from Tiltil y Mejil-
lones.

Nitzschia praereinholdii Schrader.
(Figure 6: 1-9, 13-15.)
DEscripTION. Schrader, 1973a, p. 708, pl. 5, figs. 24-25, 20, 23, 26.

Nitzschia cf. N. praereinholdii Schrader.
(Figure 6: 16-19.)

VoLt. XXXIX] SCHRADER: DIATOM STRATIGRAPHY 549

DescripTIion. Schrader, 1973a, p. 708, pl. 5, figs. 20, 23-26.

Discussion. Specimens (from EMD-8-9-240/241 cm.) have been placed
doubtfully in NV. praereinholdii because they possess bent raphe bars near the
apices similar to Nitzschia marina Grunow.

Nitzschia riedelia Schrader, new species.

DESCRIPTION. Valves elliptical with slightly convex margins and broadly
rounded apices, 22-30 » long, 5—6 » wide in the middle of the valves. Transapical
costae straight in transapical direction in the middle part of the valve, curved
near the apices, 11-13 in 10» in the middle becoming closer near the apices,
approximately 15 in 10 ». Intercostal membranes hyaline even in oblique light.
Apical field not developed. Raphe marginal, central nodule absent, keel distinct,
about 15 keel-punctae in 10 p.

Discussion. This species is close to V. burcklia but differs in the absence
of apical fields and a central nodule. It is close to NV. rolandii but differs in
having finer transapical costae, no central nodule, and hyaline intercostal mem-
branes noticeable even in oblique light.

DERIVATION OF NAME. Dedicated to W. R. Riedel of Scripps Institution of
Oceanography, La Jolla.

Hototype. Figure 6: 21 from EMD-8-15-504/505 cm., a sample from North
Pacific Diatom Zone 21. California Academy of Sciences, Department of Geol-
ogy no. 54420 (diatom collection).

PaRATYPE. Figure 6: 20. California Academy of Sciences, Department of
Geology no. 54421 (diatom collection).

OcCURRENCE. Found in North Pacific sediments of Middle Miocene age.

Nitzschia rolandii Schrader.
DescriPTION. Schrader, 1973a, p. 708, pl. 5, fig. 31, pl. 26, figs. 3-5.

Nitzschia seiboldia Schrader, new species.

Description. Valves elliptical with slightly convex margins, and broadly
rounded apices. 28—45 » long, 5—6 » wide in the middle of the valves. Transapi-
cal costae straight. in transapical direction in the middle of the valve and be-
coming curved towards the apices, 12 in 10 ». Intercostal membranes punctated,
with two rows of fine punctae neighboured to the transapical costae. Punctae in
decussate arrangement, approximately 20-25 in 10 ». Raphe marginal, keel dis-
tinct with about 11 keel punctae in 10 », central nodule absent. Apices dimorph,
one apex with downwards curved transapical costae, the other with curved trans-
apical costae, apical field absent.

Discussion. This species is close to Nitzschia species 2 Schrader (Schrader,
1973a, p. 708, pl. 5, figs. 16-18); no other similar species has been found in the
literature. Specimens from the EMD generally are broken, and some are twisted

550 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

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Ficure 5. Magnification 1000 x, Leitz-Orthoplan, Apo Oil 92/1.40 n.A. 1. EMD-8-9-

193/194 cm., Thalassionema nitzschioides Grunow; 2. EMD-8-9-240/241 cm., T. nitzschi-
oides; 3. EMD-8-15-300/301 cm., Synedra jouseana Sheshukova-Poretzkaya; 4. EMD-8-15-

504/505 cm., Thalassionema nitzschioides Grunow; 5. EMD-8-15-504/505 cm., T. nitzschi-
oides; 6. EMD-8-15-504/505 cm., T. nitzschioides, aberrant specimens; 7. EMD-8-9-
240/241 cm., T. hirosakiensis (Kanaya) Schrader; 8. EMD-8-15-200/201 cm., Rowuxia
naviculoides Schrader; 9. EMD-8-9-240/241 cm., R. moholensis Schrader, new species;
10. EMD-8-11-30/31 cm., R. cf. R. naviculoides Schrader; 11. EMD-8-9-240/241 cm.,

Vor. XXXIX] SCHRADER: DIATOM STRATIGRAPHY 551

in apical direction (the author found numerous individuals in samples from San
Felipe CAS no. 39904 (Hanna in Hertlein, 1968) ).

DERIVATION OF NAME. Dedicated to Prof. E. Seibold, who encouraged the
author during many years with advice and help.

Hototyre. Figure 5: 40-41 from EMD-8-15-300/301 cm., a sample from
North Pacific Diatom Zone 21. California Academy of Sciences, Department of
Geology no. 54422 (diatom collection).

PaRATYPE. Figure 5: 45. California Academy of Sciences, Department of
Geology no. 54423 (diatom collection).

Nitzschia species.

The following Nitzschia species are treated informally and assigned numbers,
because they were found only once, or found only as fragments. The corrected
taxonomic assignments will require additional or better preserved specimens.

Nitzschia species 3 Schrader.

Discussion. Valve linear-elliptical, 15 » long, 4 » wide. 13 transapical costae
in 10. This species is close to Nitzschia porteri Burckle non Frenguelli, but
differs in the smaller size, in the heteropol apical fields, and in the finer struc-
ture of the intercostal membranes.

FIGURED SPECIMEN. Figure 5: 33-34.

=

R. moholensis Schrader, new species (type); 12. EMD-8-9-240/241 cm., R. moholensis,
California Academy of Sciences, Department of Geology no. 54424; 13. EMD-8-9-240/241
cm., R. species 1 Schrader; 14. EMD-8-9-193/194 cm., R. californica M. Peragallo;
15. EMD-8-9-193/194 cm., R. californica; 16. RC-12-65-1915/1917.5 cm., WNitzschia
porteri Burckle non Frenguelli; 17. RC-12-65-1105/1107.5 cm., N. miocenica Burckle;
18. RC-12-65-1105/1107.5 cm., N. miocenica; 19. RC-12-65-1105/1107.5 cm., N. miocenica;
20. RC-12-65-1105/1107.5 cm., N. miocenica; 21. RC-12-65-1105/1107.5 cm., N. miocenica;
22. RC-12-65-1105/1107.5 cm., N. miocenica (girdle view); 23. EMD-8-9-193/194 cm.,
N. cylindrica Burckle; 24. EMD-8-9-193/194 cm., N. cylindrica; 25. EMD-8-10-100/101
cm., NV. cylindrica; 26. EMD-8-10-100/101 cm., NV. cylindrica; 27. EMD-8-10-100/101 cm.,
Nitzschia cylindrica Burckle; 28. EMD-8-10-100/101 cm., N. cylindrica; 29. EMD-8-10-
100/101 cm., N. cylindrica; 30. EMD-8-10-100/101 cm., N. cylindrica; 31. EMD-8-10-
44/45 cm., N. porteri Burckle non Frenguelli; 32. EMD-8-10-44/45 cm., N. porteri;
33. EMD-8-10-44/45 cm., NV. species 3 Schrader; 34. EMD-8-10-44/45 cm., N. species 3;
35. EMD-8-15-100/101 cm., NV. species 7 Schrader; 36. EMD-8-15-100/101 cm., N. species
7; 37. EMD-8-15-200/201 cm., N. species 8 Schrader; 38. EMD-8-15-200/201 cm., N.
species 8; 39. EMD-8-15-200/201 cm., NW. species 8; 40. EMD-8-15-300/301 cm., WN.
setboldia Schrader, new species (type); 41. EMD-8-15-300/301 cm., NV. seiboldia (type),
California Academy of Sciences, Department of Geology no. 54422; 42. EMD-8-15-200/201
cm., Nitzschia~cf. N. heteropolica Schrader; 43. EMD-8-15-504/505 cm., N. species 4
Schrader; 44. EMD-8-15-504/505 cm., N. species 4; 45. EMD-8-15-300/301 cm., N.
seiboldia Schrader new species, California Academy of Sciences, Department of Geology
no. 54423.

552 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

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Ficure 6. Magnification 1000 x, Leitz-Orthoplan, Apo Oil 92/1.40 n.A. 1. EMD-8-
11-300/301 cm., Nitzschia praereinholdii Schrader; 2. EMD-8-11-300/301 cm., N. prae-
reinholdii; 3. EMD-8-11-300/301 cm., N. praeveinholdii; 4. EMD-8-11-30/31 cm., JN.
praereinholdi; 5. EMD-8-11-30/31 cm., N. praereinholdi; 6. EMD-8-9-193/194 cm.,
N. praereinholdii; 7. EMD-8-9-193/194 cm., N. praereinholdit; 8. EMD-8-9-193/194 cm.,
N. praereinholdii; 9. EMD-6-2-30/31 cm., N. praereinholdit; 10. EMD-8-11-300/301 cm.,
N. moholensis Schrader, new species (type); 11. EMD-8-11-300/301 cm., N. moholensis
(type), California Academy of Sciences, Department of Geology no. 54419; 12. EMD-8-
11-390/391 cm., NV. species 5 Schrader; 13. EMD-6-2-30/31 cm., N. praereinholdi Schrader;
14. EMD-6-2-30/31 cm., N. praereinholdii; 15. EMD-6-2-30/31 cm., N. praereinholdii;

VoL. XXXIX] SCHRADER: DIATOM STRATIGRAPHY 553

Nitzschia species 4 Schrader.

Discussion. Only fragments have been found. They display a central nod-
ulus, a marginal well defined keel with 9 keel punctae in 10 » and 17 transapical
costae in 10. Intercostal membranes with one row of punctae. No similar
species has been observed in the literature.

FIGURED SPECIMEN. Figure 5: 43-44.

Nitzschia species 5 Schrader.

Discussion. Only one fragment has been found. It has 9 transapical costae
in 10 », intercostal membranes with double rows of small punctae in decussate
arrangement, rows near the transapical costae. This species is close to Nitzschia
reinholdii Kanaya but differs from the latter in the lanceolate valve, the arrange-
ment of keel punctae, here 9 in 10 », and the finer punctated intercostal mem-
branes.

FIGURED SPECIMEN. Figure 6: 12.

Nitzschia species 6 Schrader.

Discussion. Valve elliptical with convex margins, 22 » long, 8 » wide, with
12 transapical costae in 10 », slightly convex near the apices. Intercostal mem-
branes hyaline even in oblique light, with about 14 keel punctae in 10 ». This
species is close to Nitzschia invisa Schrader but differs from it in the coarser
arrangement of transapical costae. It is also similar to Nitzschia porteri Fren-
guelli, but owing to the fact that I have not seen a NV. porteri from Tiltil the
correct taxonomic position could not be ascertained. Nitzschia porteri possesses
12—13 keel punctae in 10 » and 12-13 transapical striae in 10 p.

FIGURED SPECIMEN. Figure 6: 32.

Ps
16. EMD-8-9-240/241 cm., Nitzschia cf. N. praereinholdii; 17. EMD-8-9-240/241 cm.,
Nitzschia cf. N. praereinholdii; 18. EMD-8-9-240/241 cm., Nitzschia cf. N. praereinholdii;
19. EMD-8-9-240/241 cm., Nitsschia cf. N. praereinholdii; 20. EMD-7-1-41/42 cm.,
N. riedelia Schrader, new species, California Academy of Sciences, Department of Geology
no. 54421; 21. EMD-8-15-504/505 cm., NV. riedelia (type), California Academy of Sciences,
Department of Geology no. 54420; 22. EMD-8-15-504/505 cm., N. rolandii Schrader;
23. EMD-8-15-200/201 cm., N. kanayensis Schrader, new species (type), California
Academy of Sciences, Department of Geology no. 54417; 24. EMD-8-15-93/94 cm., N.
rolandi Schrader; 25. EMD-8-11-300/301 cm., N. kanayensis Schrader, new species,
California Academy of Sciences, Department of Geology no. 54418; 26. EMD-8-11-300/301
cm., N. rolandii Schrader; 27. EMD-8-10-44/45 cm., N. rolandiit; 28. EMD-8-9-240/241
cm., N. kanayensis Schrader, new species, California Academy of Sciences, Department of
Geology no. 54426; 29. EMD-8-9-240/241 cm., N. burcklia Schrader, new species (type),
California Academy of Sciences, Department of Geology no. 54415; 30. EMD-8-9-240/241
cm., N. burcklia, California Academy of Sciences, Department of Geology no. 54416;
31. EMD-8-11-300/301 cm., NV. invisa Schrader; 32. EMD-8-11-300/301 cm., N. species 6
Schrader.

554 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Nitzschia species 7 Schrader.

Discussion. Only one fragment of this small Nitzschia species has been
observed. It is 15 » long, 6 » wide, with 11-12 transapical costae in 10 », curved
near the apices; intercostal membranes have two rows of punctae with punctae
in decussate arrangement. No similar species has been observed in the literature.

FIGURED SPECIMEN. Figure 5: 35-36.

Nitzschia 8 Schrader.

Discussion. Only one specimen has been observed. It is 23 » long, 7 » wide,
with 9 transapical costae in 10 ,; intercostal membranes finely punctated.
Apices heteropol. Transapical costae curved near the apices. Keel distinct, with
9 keel punctae in 10 ». No similar species has been observed in the literature.

FIGURED SPECIMEN. Figure 5: 37-39.

Genus Rhaphoneis Ehrenberg

Rhaphoneis angularis Lohmann.
Description. Lohmann, 1938, pp. 92—93, pl. 22, figs. 6-8.

Rhaphoneis angustata Pantocsek.

DescripTIon. Sheshukova-Poretzkaya, 1967, pp. 241-242, pl. 41, fig. 8,
pl. 43, fig. 2.
Rhaphoneis margaritalimbata Mertz.

DEscrIPTION. Mertz, 1966, p. 27, pl. 6, figs. 1-3.

Rhaphoneis sacchalinensis Sheshukova-Poretzkaya.

Description. Sheshukova-Poretzkaya, 1967, p. 242, pl. 42, fig. 2.

Genus Rhizosolenia Ehrenberg
Rhizosolenia barboi Brun.
Description. Donahue, 1970, p. 136; Schrader, 1973a, p. 709, pl. 24, figs.
4-7,

Rhizosolenia hebetata forma hiemalis Gran.
DEscriPTION. Hustedt, 1930, pp. 590-592, fig. 337.

Rhizosolenia miocenica Schrader.
DescripTIon. Schrader, 1973a, p. 709, pl. 10, figs. 2-6, 9-11.

Rhizosolenia praealata Schrader.
DESCRIPTION. Schrader, 1973a, p. 709, pl. 10, fig. 13.

VoL. XXXIX] SCHRADER: DIATOM STRATIGRAPHY 555

Rhizosolenia praebarboi Schrader.
DescriPTIon. Schrader, 1973a, pp. 709-710, pl. 24, figs. 1-3.

Rhizosolenia styliformis Brightwell.
DescriPTIOoNn. Hustedt, 1930, pp. 584-588, figs. 333-335.

Genus Rouxia Brun and Heéribaud
Rouxia californica M. Peragallo.
DEscriPTIon. Hanna, 1930, pp. 186-188, pl. 14, figs. 6-7.

Rouxia moholensis Schrader, new species.

DEscrIPTION. Valves linear-oblong with parallel margins slightly constricted
in the middle. Apices broadly rounded, 35-42 » long, 6 » wide in the middle. The
two rudimentary raphe bars widely separated and situated near the apices.
Transapical striae 18 in 10 », transapical in the middle of the valve, becoming
radial towards the apices. Transapical striae formed by elliptical pores. No api-
cal lines are present. Axial area narrow, central area thin-elongated. Valves
isopol.

Discussion. This species differs from Rouxia diploneides in its finer struc-
ture and lack of apical lines.

DERIVATION OF NAME. Dedicated in memory of the Experimental Mohole
Drilling.

Hototyre. Figure 5: 11-12 from EMD-8-9-240/241 cm., a sample from
North Pacific Diatom Zone 10. California Academy of Sciences, Department of
Geology no. 54424 (diatom collection).

OccuRRENCE. Found in the Neogene Section off Baja California.

Rouxia naviculoides Schrader.
(Figure 5: 8.)
DescriIPTION. Schrader, 1973a, p. 710, pl. 3, figs. 27-32.

Rouxia cf. R. naviculoides Schrader.
(Figure 5: 10.)

Discusston. Specimens found are tentatively placed in R. naviculoides, but
they differ from others of that species in the butterflylike arrangement of the
central area.

Rouxia species 1 Schrader.

Discussion. Only one specimen of this species was found. It is close to R.
moholensis but differs in the coarser structure, and in the presence of one apical
line near the middle axis.

FIGURED SPECIMEN. Figure 5: 13.

556 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Genus Stephanopyxis Ehrenberg

Stephanopyxis turris (Greville and Arnotte) Ralfs.
DeEscripTIon. Hustedt, 1930, pp. 304-307, figs. 140-144.

Genus Synedra Ehrenberg

Synedra jouseana Sheshukova-Poretzkaya.
Description. Sheshukova-Poretzkaya, 1967, p. 245: Schrader, 1973a,
p: 710, pl. 23. figs... 21=23, 25, 38;

Genus Thalassionema Grunow

Thalassionema hirosakiensis (Kanaya) Schrader.
(Figure 5: 7.)

DESCRIPTION. Kanaya, 1959, pp. 104-106, pl. 9, figs. 11-15; Schrader,
1O73a. ip. UL, ple 23, tigs..31=33°:

Thalassionema claviformis Schrader.

DESCRIPTION. Schrader, 1973a, p. 711, pl. 23, figs. 11, 15.

Thalassionema nitzschioides Grunow.
(Figure 5: 1-2, 4-6.)
DESCRIPTION. Hustedt, 1959, pp. 244-246, fig. 725; Hasle, 1967, p. 111,
figs. 5, 27-34, 39-44.
Genus Thalassiosira Cleve
Thalassiosira convexa Mukhina.
(Figure 2: 5-6.)

Description. Donahue, 1970, pp. 136-137, pl. 3, figs. a-f.

Thalassiosira decipiens (Grunow) Joergensen.

DEscRIPTION. Hustedt, 1930, pp. 322-323, fig. 158.

Thalassiosira excentrica (Ehrenberg) Cleve.

Description. Sheshukova-Poretzkaya, 1967, pp. 141-142, pl. 14, fig. 4.

Thalassiosira antiqua (Grunow) Cleve.

DeEscriPpTION. Sheshukova-Poretzkaya, 1967, pp. 143-144, pl. 14, fig. 3.

Thalassiosira species A Burckle.
(Figure 2: 3-4.)

DescripTIONn. Burckle, 1972, p. 241, pl. 1, fig. 1.

VoL. XXXIX] SCHRADER: DIATOM STRATIGRAPHY 557

Thalassiosira nativa Sheshukova-Poretzkaya.
(Figure 2: 11-21, 25-26.)
DescripTIon. Sheshukova-Poretzkaya, 1967, p. 145, pl. 14, fig. 7.

Thalassiosira praeconvexa Burckle.
(Figure 2: 7-10.)
DeEscripTIon. Burckle, 1972, pp. 241-242, pl. 2, figs. 7-9.

Thalassiosira oestrupii (Ostenfeld) Proshkina-Lavrenko.
(Figure 1: 31; Figure 2: 1-2.)
DescripTion. Hustedt, 1930, p. 318, fig. 155 as Coscinosira oestruppi.

Thalassiosira usatchevii Jouse.
(Figure 3: 1-2.)
DeEscripTIoN. Sheshukova-Poretzkaya, 1967, p. 150, pl. 15, fig. 3.

Thalassiosira species 2 Schrader.

Discussion. This species is close to Thalassiosira nativa but differs in
lacking numerous scattered strutted tubuli in the center. No other similar species
were found mentioned in the literature.

FIGURED SPECIMEN. Figure 2: 22-24, 27-28.

Genus Thalassiothrix Cleve and Grunow

Thalassiothrix longissima Cleve and Grunow.
DescripTIon. Hasle, 1967, p. 114, fig. 20.

Genus Triceratium Ehrenberg
Triceratium antiquum Pantocsek.
(Figure 1: 28.)

DESCRIPTION. Pantocsek, 1886, p. 51, pl. 13, fig. 115; Azpeitia, 1911, p. 221,
plal2, tie. 2.

Triceratium cinnamomeum Greville.
(Figure 1: 29.)

DeEscriPTIon. Greville, 1863, p. 232, pl. 9, fig. 12.

Triceratium condecorum Brightwell.

DeEscriPTIon. Hanna, 1932, p. 221, pl. 17, figs. 1, 3.

SUMMARY AND CONCLUSIONS

(1) The North Pacific Diatom Zonation as established from cores obtained
during the DSDP leg 18, north of 35° North longitude. provided a basis

558

(4)

(6)

CALIFORNIA ACADEMY OF SCIENCES [PRroc. 4TH SER.

for establishing a diatom stratigraphy of core samples from the Experi-
mental Mohole Drilling site off Guadalupe Island, Mexico.

Paleoclimatic oscillations found at the EMD site are comparable to those
found at DSDP leg 18, Site 173; no warm period was detected comparable
to those found in Lamont Doherty Cores south of 10° longitude.

North Pacific Diatom Zones 20 to 24 are correlative with the lower part of
the Corbisema triacantha Silicoflagellate Zone and the upper part of the
Dictyocha octacantha Silicoflagellate Zone of Martini (1972), and with
NN 6 Standard Nannoplankton Zone of Martini (1971).

The base of the Late Miocene as defined by Burckle (1972), as occurring
within Geomagnetic Epoch 11 of the tropics, and as defined by Bolli (1957)
as the Globorotalia mayeri Zone below the Globorotalia menardii Zone, is
correlative with North Pacific Diatom Zone 19, and in the Experimental
Mohole Drilling it is between samples EMD-8-15-100/101 cm. and EMD-
8-15-200/201 cm. This boundary is correlative (chart 4) with the Corbr-
sema triacantha Silicoflagellate Zone (Martini, 1972), with the NN 6 Zone
of the Standard Nannoplankton Zone of Martini (1971), with the Luisian
stage of California (Parker, 1964; Kanaya, 1971; Bandy and Ingle, 1970).
Berggren (1972) placed the Middle to Late Miocene boundary within the
upper part of the N 15 Standard Foraminiferal Zone and indicated a radio-
metric age of 13.5 m.y. for the Luisian-Mohnian boundary, which correlates
with the Late to Middle Miocene boundary of Bandy and Ingle (1970).
The different correlations are compared in charts 3 and 4.

Dymond (1966) determined a Potassium-Argon date of 12.3 = 0.4 m.y.
for a glass sample from EMD-8-15-89 cm. This date agrees with the
correlation of the base of North Pacific Diatom Zone 19 with the paleo-
magnetic scale, and the correlation of the paleomagnetic scale with the
radiometric time scale (Berggren, 1972).

The Pliocene-Late Miocene boundary as defined by Burckle (1972) is at
the end of Geomagnetic Epoch 5 at about 5.5 million years B.P. In the
Experimental Mohole Drilling this boundary is between samples EMD-8-
10-44/45 cm. and EMD-8-10-100/101 cm., and it correlates with the
Dictyocha pseudofibula Silicoflagellate Zone of Martini (1972), with the
NN 10 Standard Nannoplankton Zone of Martini (1971), and with the
upper Mohnian Stage of Bandy and Ingle (1970).

The sedimentation rate for the Experimental Mohole Drilling Core in the
interval from 80—95 meters below the sea floor, is about 7 meters/million
years; for the interval from 95 to 115 meters it is approximately 30 m./m.y.,
and for the interval from 115 to 135 meters below the sea floor approxi-
mately it is again 7 m./m.y. The abrupt increase of sedimentation rate

VoL. XXXIX] SCHRADER: DIATOM STRATIGRAPHY 559

corresponds with the increase of silt sized minerals and sponge spicules. The
sedimentation rate determined by correlation of North Pacific Diatom Zones
to paleomagnetic stratigraphy and hence to the radiometric time scale,
differs from that obtained by using Dymond’s (1966) Potassium-Argon
dates (charts 4, 5).

(7) Published correlations of planktonic marine diatoms (Kanaya, 1971), of
planktonic foraminiferans (Parker, 1964; Bandy and Ingle, 1970) with the
Californian Marine Stages contain discrepancies which could not be resolved
in this paper because of a lack of stratotype diatom-bearing material from
California [the North Pacific Diatom Zones have not yet been correlated
to the Californian Marine Stages].

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INDEX TO VOLUME XXXIX

FOURTH SERIES

New names and principal references in boldface type.

italics.

Abe, Tokiharu, and William N. Eschmeyer,
A new species of the scorpionfish genus
Helicolenus from the north Pacific
Ocean, 47-53.

Abe, Tokiharu, see Eschmeyer, William N.

Acanthina grandis, 28

Acanthochitona

armata, 501
viridis, 501
Actaea
nana, 319
parvula, 315, 318, 329, 330
remota, 318
Acteocina infrequens, 28
Actinocyclus, 532
cubitus, 524, 531, 532
ehrenbergii, 524, 532
ellipticus, 524, 533
ellipticus moronensis 524, 532, 533
ingens, 526, 534
okhotensis, 524, 534

Agastache, 146, 147, 177

Agriopus, 307

Alaba supralirata, 28

Allips, 116, 117

concolor, 111, 116, 117, 117, 118
Allomycter, 389, 390, 393, 403, 405, 406
dissutus, 393

Alvania, 28
galapagensis, 28
halia, 28
hoodensis, 28
nemo, 28

Ambrosia, 146, 179
artemisiifolia, 145, 146

Amphineura:Chitonidae), A new Hawaiian
chiton, Rhyssoplax linsleyi (Mollusca:,
by Glenn E. Burghardt, 501-506.

Amphithalmus trosti, 28

Anachis, 28

tabogaensis, 28
Anatina (Raeta) undulata, 27

References to illustrations in

anatomy of the forelimb of the sea otter
(Enhydra lutris), Muscular, by L. D.
Howard, 411-500
Anodontia, 33
edentuloides, 33
(Lissosphaira), 33
(Lissosphaira) spherica, 33
philippiana, 33
spherica, 27, 30, 32, 33, 34
Anodota sphericula, 33
Anomia peruviana, 27
Anthochiton, 505
Apistus, 339
Aploactis, 373
breviceps, 373
(Apodes:Ophichthidae), Two new _ genera
and two new species of western Pacific
snake-eels, by John E. McCosker,
111-120.
Apogon evermanni, 130
Aprionodon, 391, 405
Arca
(Arcopsis) solida, 28
pacifica, 28
Archoplites interruptus, 77, 79, 80, 82, 83
Artemisia, 146
tridentata, 146, 173
Ashlock, Peter D., The Lygaeidae of the
Galapagos Islands (Hemiptera: Het-
eroptera), 87-103.
Aster, 146, 170
tanacetifolius, 146
Asterolampra, 534, 546
grevillei, 534
marylandica, 524, 534
Asteromphalus, 546
Atergatis elegans, 319
Atlantic, Nezumia (Kuronezumia) bubonis a
new subgenus and species of grenadier
(Macrouridae: Pisces) from Hawaii
and the western North, by Tomio
Iwamoto, 507-516.

[563]

564 CALIFORNIA ACADEMY OF SCIENCES

Atrina, 28
tuberculosa, 28

Baileya, 146
multiradiata, 146, 167
Balanus, 72
allium, 72
amphitrite, 67, 69
amphitrite amphitrite, 65, 66, 66, 67, 69
amphitrite hawaiiensis, 67
(Armatobalanus), 72
balanoides, 69
balanus, 69
crenatus, 69
eburneus, 67, 69, 70
improvisus, 65, 66, 67, 68, 69
Balcis, 28

berryi, 28
Baltra (South Seymour) Island, Galapagos
Islands, Pliocene fossils from, by
Leo G. Hertlein, 25-46.
Barbatia reeveana, 28
barnacle records (Cirripedia, Thoracica),

New, by Victor A. Zullo, Dea B.
Beach, and James T. Carlton, 65-74.

Bascanichthys, 116, 117

Beach, Dea B., see Zullo, Victor A.

Beryx spendens, 47

Biddulphia, 534

moholensis, 518, 534, 544

Bombonia, 135

brachyuran crabs of Easter Island, The,
by John S. Garth, 311-336.

Brigger, A. L., see Mandra, York T.

Walter C., and Marjorie V. C.
Falanruw, A new lizard of the genus
Emoia (Scincidae) from the Marianas
Islands, 105-110.

Bruniopsis, 534

mirabilis, 524, 531, 532, 534

Bufichthys, 341, 342, 343, 363, 364

erosa, 342, 343

grossa, 342

horrida, 342
Bulimulus calvus, 22

Brown,

Bulla punctulata, 28

Burghardt, Glenn E., A new Hawaiian
chiton, Rhyssoplax linsleyi (Mollusca:
Amphineura: Chitonidae), 501-506.

[Proc. 4TH SER.

California freshwater fishes, A key, based on
scales to the families of native, by
Richard W. Casteel, 75-86.

Calliscyllium, 257

Campogramma, 244, 246, 247

glaycos, 246, 246
liro, 246
vadigo, 246

Campogramma (Pisces: Carangidae), Com-
parative revision of Scomberoides,
Oligoplites, Parona, and Hypacanthus
with comments on the phylogenetic
position of, by William F. Smith-Vaniz
and Jon C. Staiger, 185-256.

Cancellaria, 28

ovata, 28

Cancer

convexus, 317
cymodoce, 321
dodone, 319
electra, 320
variegatus, 323
Cantharus, 28
janellii, 28
scissus, 28

Carangidae), Comparative revision of Scom-
beroides, Oligoplites, Parona, and
Hypacanthus with comments on the
phylogenetic position of Campo-
gramma (Pisces:, by William F. Smith-
Vaniz and Jon C. Staiger, 185-256.

Caranx, 191

(Gnathanodon) speciosus, 240

Carcharhinidae), Ctenacis and Gollum, two
new genera of sharks, (Selachii:, by
L. J. V. Compagno, 257-272.

(Carcharhiniformes: Triakidae), with a re-
definition of the family Triakidae and
a key to Triakid genera, Gogolia file-
woodi, a@ new genus and species of
shark from New Guinea, by L. J. V.
Compagno, 383-410.

Carcharhinus, 389, 391, 405

Cardium elenense, 28

Carlton, James T., see Zullo, Victor A.

Carpilius, 312

convexus, 315, 317, 329, 330

Carpilodes, 313

monticulosus, 318
rugatus, 318

VoL. XXXIX]

Casteel, Richard W., A key, based on scales,
to the families of native California
freshwater fishes, 75-86.

Casuarina, 110

Catostomus

latipinnis, 76
luxatus, 76
mniotiltus, 76
occidentalis humboldtianus, 76
occidentalis occidentalis, 76, 81
platyrhynchus, 76
rimiculus, 76
santaanae, 76
tahoensis, 76
Cavilucina densata, 33
Centronotus
argenteus, 221
glaycos, 244, 248
vadigo, 228, 229, 230

Cerambycidae), The genus Mecas LeConte
(Coleoptera:, by John A. Chemsak and
E. G. Linsley, 141-184.

Cerithiopsis, 28

curtata, 28
galapagensis, 28

Cerithium adustum, 28

Chaenogaleus, 392, 405

Chama, 28

Champson sechellensis, 303

Cheila equestris, 28

Chelonibia patula, 69

Chemical extraction techniques to free fossil
silicoflagellates from marine sedi-
mentary rocks, by York T. Mandra,
A. L. Brigger, and Highoohi Mandra,
273-284.

Chemsak, John A., and E. G. Linsley, The
genus Mecas LeConte (Coleoptera:
Cerambycidae), 141-184.

Cheroscorpaena, 339

Chione, 28

pertincta, 37
seymourensis, 37, 38
undatella, 28
Chiton affinis, 505
chiton, Rhyssoplax linsleyi (Mollusca: Am-

phineura: Chitonidae), A new
Hawaiian, by Glenn E. Burghardt,
501-506.

Chitonidae), A new Hawaiian chiton, Rhys-

INDEX

565
soplax linsleyi (Mollusca: Amphi-
neura:, by Glenn E. Burghardt, 501-
506.

Chlamys, 1, 2
amandi, 4

(Argopecten) circularis, 1, 28
benedicti, 4
(Chlamys) ineantata, 2, 3
(Chlamys) lowei, 1
hastata, 4
incantata, 2, 4
magnifica, 31
(Nodipecten) magnifica, 1, 28, 31, 32,
34, 36
(Nodipecten) nodosa, 31
Chlamys (Mollusca: Pelecypoda) from the
Galapagos Islands, Description of a
new species of, by Leo G. Hertlein,
1-6.
Chlorodiella, 312, 314, 320
barbata, 320
cytherea, 315, 320, 329
laevissima, 320
nigra, 320
Chlorodius cytherea, 320, 330
Choridactyloides, 339
Choridactylus, 308, 339
Chorinemus, 190, 191, 192, 229, 242, 248
altus, 223, 248
commersonianus, 200
delicatulus, 194, 248
exoletus, 194, 248
farkharrii, 248
guaribira, 225, 248
hainanensis, 199, 201, 248
lanceolatus, 221, 222, 249
leucopthalmus, 194, 249
maculosus, 249
mauritianus, 205, 207, 249
moadetta, 205, 206, 207, 249
occidentalis, 221, 242, 249
orientalis, 190, 205, 207, 249
palometa, 225, 249
saltans, 221, 249
sanctipetri, 205, 207, 208, 249
tala, 190, 199, 205, 250
tol, 190, 209, 210, 250
toloo, 199, 201, 250
Chorismopelor, 339
Chthamalus stellatus thompsoni, 67

566 CALIFORNIA ACADEMY OF SCIENCES

(Cirripedia, Thoracica), New barnacle rec-
ords, by Victor A. Zullo, Dea B.
Beach, and James T. Carlton, 65-74.

Cladogramma, 534

dubium, 524, 534

(Coleoptera: Cerambycidae), The genus
Mecas, by John A. Chemsak and E.G.
Linsley, 141-184.

Colubrina texensis, 163

Compagno, L. J. V., Ctenacis and Gollum,
two new genera of sharks (Selachii:
Carcharhinidae), 257-272.

Compagno, L. J. V., Gogolia filewoodi, a new
genus and species of shark from
New Guinea (Carcharhiniformes:
Triakidae), with a redefinition of the
family Triakidae and a key to
Triakid genera, 383-410.

Comparative revision of Scomberoides, Oli-
goplites, Parona, and MHypacanthus
with comments on the phylogenetic
position of Campogramma (Pisces:
Carangidae), by William F. Smith-
Vaniz and Jon C. Staiger, 185-256.

Conus, 28

fergusoni, 28
lucidus, 28

Corbicula manilensis, 68

Corbisema triacantha, 531, 558

Cordia, 93

Corythobatus, 339

Corythoichthys, 127, 135

flavofasciatus, 131

flavofasciatus conspicillatus, 131

flavofasciatus flavofasciatus, 131
Coscinodiscus, 535, 541

antiquus, 524, 531, 535

endoi, 524, 535, 540

flexuosus, 539, 541

lancettula, 518, 536, 542

lewisianus, 524, 532, 535

marginatus, 524, 535

marginatus fossilis, 535

moholensis, 518, 535, 540

nodulifer, 524

paleaceus, 541, 543

plicatus, 535, 544

praepaleaceus, 541, 543

reniformis, 541

species 1 Schrader, 537, 544

[Proc. 47TH Serr.

species 2 Schrader, 538, 539
species 3 Schrader, 538, 539
stellaris symbolophora, 537
symbolophorus, 524, 536, 537
temperei, 524, 529, 537, 538
vetustissimus, 524, 537, 540
vetustissimus javanicus, 524, 531, 537
yabei, 524, 525, 526, 531, 532, 537, 538,
539, 540, 544
crabs of Easter Island, The brachyuran, by
John S. Garth, 311-336.
Craspedodiscus, 54:1
coscinodiscus, 526, 531, 541
Crassostrea virginica, 65
Creusia, 71, 72
domingensis, 71, 71, 72
Ctena galapagana, 28
Ctenacis, 257, 258, 263, 264, 268, 269, 390,
405
felmanni, 257, 258, 260, 263, 264, 270
Ctenacis and Gollum, two new genera of
sharks (Selachii: Carcharhinidae), by
L. J. V. Compagno, 257-272.
Cuminga, 28
lamellosa, 28
Cussia, 517, 541
mediopunctata, 518, 542
mediopunctata matraensis, 518, 542
moholensis, 518, 536, 542
paleacea, 518, 524, 531, 532, 536, 543
praepaleacea, 518, 524, 531, 532, 536,
542, 543
species 1 Schrader, 536, 543
tatsunokuchiensis, 518, 536, 543
Cyclograpsus, 312, 325
longipes, 315, 325, 329, 330
Cyclopecten
(Cyclopecten) exquisitus, 1
(Cyclopecten) liriope, 1
(Cyclopecten) pernomus, 1
(Delectopecten) polyleptus, 1
(Delectopecten) zacae, 1
(Hyalopecten) neoceanicus, 1
Cylichna, 29
defuncta, 29
Cymatium lineatum, 29
Cymoninus, 89, 97
notabilis, 94, 97, 99
Cymus galapagensis, 95
Cyperus confertus, 97

Vor. XXXIX]

Cypraea nigropunctata, 29

Cyprinodon
macularius californiensis, 77, 80, 82
nevadensis calidae, 77
nevadensis nevadensis, 77
nevadensis shoshone, 77
salinus, 77

Cytharella camarina, 29

Dampierosa, 337, 339, 340, 365, 367, 369
daruma, 341, 351, 356, 367, 368, 369
Darwiniothamnus, 20
Darwinysius, 88, 89, 93, 94, 96
marginalis, 91, 95
wenmanensis, 92, 94, 94, 95, 96
Decterias, 339
Deleastes, 342, 343
daector, 342
Denticula, 543
dimorpha, 524, 529, 531, 536, 543
hustedtii, 524, 529, 531, 532, 536, 543
hyalina, 536, 545
hyalina hustedtii, 524, 529, 545
lauta, 524, 529, 531, 532, 536, 545
nicobarica, 524, 531, 532, 536, 545
punctata, 545
punctata hustedtii, 524, 531, 536, 545
Dentirostrum, 123, 124, 126, 127
jamssi, 121, 122, 123, 124, 125, 126, 127
Description of a new species of Chlamys

(Mollusca: Pelecypoda) from _ the
Galapagos Islands, by Leo G.
Hertlein, 1-6.

diatom stratigraphy of the experimental

Mohole Drilling, Guadalupe Site, Re-
vised, by Hans-Joachim Schrader,
517-562.
Dictyocha
octacantha, 517, 532, 558
pseudofibula, 558
Diodora alta, 29
Diplodonta
subquadrata, 35
subquadrata baltrana, 26, 28, 33, 34
Dirrhizodon, 392, 405
Doryrhamphus, 122, 123
melanopleura, 130
Dromia unidentata, 316
Dromida, 312, 314
unidentata, 315, 316, 329, 331

INDEX 567

unidentata hawaiiensis, 316
unidentata unidentata, 316
Dunckerocampus, 122, 123, 129
baldwini, 122, 123, 128, 128, 130
ben-tuviae, 122, 123
caulleryi, 123
caulleryi caulleryi, 122, 123
caulleryi chapmani, 122, 123
dactyliophorus, 122, 123, 130, 131, 132
multiannulatus, 122, 123
pessuliferus, 122, 123
Dylobolus, 141, 153
rotundicollis, 153, 157

Easter Island, The brachyuran crabs of, by
John S. Garth, 311-366.
Eleria, 190, 192
philippina, 190, 199, 201, 205, 249
Elytroleptus, 143, 144, 153
Emissola, 390
Emmydrichthys, 341, 343
vulcanus, 341, 359, 362
Emoia, 105, 107, 110
arnoensis, 105, 108, 109
atrocostata, 105, 108, 109, 110
boettgeri, 105, 109
boettgeri boettgeri, 108, 109
boettgeri orientalis, 108
caeruleocauda, 110
cyanura, 110
flavigularis, 105, 108, 109
nigra, 105, 108
slevini, 706, 107, 108, 109, 110
Emoia (Scincidae) from the Marianas Islands,
A new lizard of the genus, by Walter C.
Brown and Marjorie V. C. Falanruw,
105-110.
Encope
micropora, 27
micropora galapagana, 27
Engina pyrostoma, 29
Enhydra lutris, 411
Entelurus aequoreus, 130
Erato (Hespererato) marginata galapagensis,

29
Eridacnis, 257, 258, 263, 268, 269, 390,
391, 405

barbouri, 257
radcliffei, 257
sinuans, 257

568 CALIFORNIA ACADEMY OF SCIENCES

Erosa, 307, 337, 339, 340, 342, 343, 363, 365,
367, 369
daruma, 367
erosa, 341, 342, 351, 356, 364, 366, 367
fratrum, 364, 365
iridea, 364, 365
Eschmeyer, William N., Voshitsugu Hirosaki,
and Tokiharu Abe, Two new species
of the scorpionfish genus Rhinopias,
with comments on related genera and
species, 285-310.
Eschmeyer, William N., and K. V. Rama Rao,
Two new scorpionfishes (genus Scor-
paenodes) from the Indo-west Pacific,
with comments on Scorpaenodes
muciparus (Alcock), 55-64.
Eschmeyer, William N., and Kaza V. Rama
Rao, Two new stonefishes (Pisces:
Scorpaenidae) from the Indo-west
Pacific, with a synopsis of the sub-
family Synanceiinae, 337-382.
Eschmeyer, William N., see Abe, Tokiharu
Ethmodiscus, 545
rex, 525, 54.5

Etisodes
electra, 320, 330
sculptilis, 320

Etisus, 313
electra, 315, 320, 329
(Etisodes) electra, 320
rugosus, 320

Eucidaris thouarsii, 27

Eugaleus, 390

Eulimostraca, 29

galapagensis, 29
Euphorbia viminea, 95
Evips, 113
percinetus, 111, 112, 113, 174, 114, 115
extraction techniques to free fossil silico-
flagellates from marine sedimentary
rocks, Chemical, by York T. Mandra,
A. L. Brigger, and Highoohi Mandra,
273-284.
Eysenhardtia polystachya, 157, 159

Falanruw, Marjorie V. C., see
Walter C.
Fasciolaria, 67

Brown,

Fissurella, 29

[Proc. 47TH Serr.

macrotrema, 29
virescens, 29
Five new Indo-Pacific pipefishes, by Earl S.
Herald and John E. Randall, 121-140.
Flaveria linearis, 179
Florimetis cognata, 28
forelimb of the sea-otter (Enhydra lutris),
Muscular anatomy of the, by L. D.
Howard, 411-500.
Fundulus, 81
parvipinnis, 77
Furgaleus, 389, 391, 392, 393, 405
Fusinus dupetitthouarsii, 29

Gaillardia, 146, 179
pinnatifida, 146
pulchella, 146

Galapagos Islands, Description of a new
species of Chlamys (Mollusca: Pelec-
ypoda) from the, by Leo G. Hertlein,
1-6.

Galapagos Islands (Hemiptera: Heteroptera),
The Lygaeidae of the, by Peter D.
Ashlock, 87-103.

Galapagos Islands, Pliocene fossils from
Baltra (South Seymour) Island, by
Leo G. Hertlein, 25-46.

Galapagos, Three new land snails from Isla
Santa Cruz (Indefatigable Island), by
Allyn G. Smith, 7-24.

Galeocerdo, 391, 392, 405

Galeorhinus, 389, 390, 392, 393, 394, 398, 403,
405

Galeus, 390

recticonus, 394

Garth, John S., The brachyuran crabs of
Easter Island, 311-336.

Gastrocopta munita, 29, 31

genus Mecas LeConte (Coleoptera: Ceram-
bycidae), The, by John A. Chemsak
and E. G. Linsley, 141-184.

Geograpsus crinipes, 312, 315, 323, 329

Gila

bicolor bicolor, 76
bicolor obesa, 76
bicolor pectinifer, 76
crassicauda, 76
elegans, 77
mohavensis, 76
orcutti, 76

VoL. XXXIX]

Globorotalia
mayeri, 558
menardil, 558
Glycymeris maculata, 28
Gogolia, 383, 389, 390, 391, 392, 393, 394,
398, 405, 406
filewoodi, 383, 384, 386, 388, 394, 395,
396, 399, 400, 402, 403, 405
Gogolia filewoodi, a new genus and species
of shark from New Guinea (Carcha-
rhiniformes: Triakidae), with a
redefinition of the family Triakidae
and a key to Triakid genera, by
L. J. V. Compagno, 383-410.

Gollum, 257, 262, 264, 268, 269, 270,
390, 405
attenuatus, 257, 258, 261, 263, 264,
265, 270
Gollum, two new genera of sharks (Selachii;
Carcharhinidae), Ctenacis and, by
L. J. V. Compagno, 257-272.
Gordiichthys, 116
Granula, 29
minor, 29
polita, 29
Granulina margaritula, 29
Grapsilius maculatus, 321
Grapsillus cymodoce, 321
Grapsus
crinipes, 323
grapsus, 324
transversus, 324
grenadier (Macrouridae: Pisces) from
Hawait and the western North
Atlantic, Nezumia (Kuronezumia)

bubonis, a new subgenus and species of,
by Tomio Iwamoto, 507-516.
Guadalupe Site, Revised diatom stratigraphy
of the Experimental Mohole Drilling,
by Hans-Joachim Schrader, 517-562.
Guardiola, 146
tulocarpus, 146, 157
Gymnothorax, 130

North Atlantic,
Nezumia (Kuronezumia) bubonis, a
new subgenus and species of grenadier
(Macrouridae: Pisces) from, by Tomio
Iwamoto, 507-516.

Hawaii and the western

INDEX 569

Hawaiian chiton, Rhyssoplax linsleyi (Mol-
lusca: Amphineura: Chitonidae), A
new, by Glenn E. Burghardt, 501-506.
Helenium, 146
hoopesii, 146
microcephalum, 146, 167
tenuifolium, 146
Helianthus, 145, 146, 170, 179
annus, 146
tuberosus, 145, 146
Helicolenus, 47, 48, 51, 53
avius, 47, 49, 50, 51, 52, 53
barathri, 53
dactylopterus, 48, 51, 53
dactylopterus lahillei, 51
hilgendorfi, 51, 53
imperialis, 51
lengerichi, 53
maccullochi, 53
maculatus, 51
maderensis, 51
microphthalmus, 53
mouchezi, 51
papillosus, 53
percoides, 53
rufescens, 53
thelmae, 51
tristianensis, 51
uruguayensis, 51
Helicolenus from the north Pacific Ocean,
A new species of the scorpionfish genus,
by Tokiharu Abe and William N.
Eschmevyer, 47-53.
Hemiaulus, 545
polymorphus, 525, 545
Hemidiscus, 545
cuneiformis, 525, 545
simplicissimus, 525, 545
Hemigaleus, 392, 405
pingi, 392
Hemipristis, 392, 405
(Hemiptera: Heteroptera), The Lygaeidae of
the Galapagos Islands, by Peter D.
Ashlock, 87-103.
Hemitriakis, 257, 263, 389, 392, 393, 394, 398
403, 405
Heraeus, 89, 101
pacificus, 700, 101
Herald, Earl S., and John E. Randall, Five
new Indo-Pacific pipefishes, 121-140.

570 CALIFORNIA ACADEMY OF SCIENCES

Hertlein, Leo G., Description of a new species
of Chlamys (Mollusca: Pelecypoda)
from the Galapagos Islands, 1-6.

Hertlein, Leo G., Pliocene fossils from Baltra
(South Seymour) Island, Galapagos
Islands, 25-46.

Hesperoleucas

navarroensis, 76

parvipinnis, 76

symmetricus subditus, 76
symmetricus symmetricus, 76
symmetricus venustus, 76

Heterogaleus, 392

Heteroptera), The Lygaeidae of the Gala-
pagos Islands, (Hemiptera:, by Peter
D. Ashlock, 87-103.

Hexacreusia, 72

durhami, 71, 72, 73
Hipponix pilosus, 29
Hipposcorpaena, 285, 303, 304, 306
filamentosa, 304, 305

Hirosaki, Yoshitsugu, see Eschmeyer, William
N.

Howard, L. D., Muscular anatomy of the
forelimb of the sea otter (Enhydra
lutris), 411-500.

Hozukius, 48, 51

Hymeniacidon, 316

Hypacanthus, 228, 229, 230, 231, 241, 243,
244, 245, 247

amia, 229, 230, 231, 232, 238, 239, 241,
247

Hypacanthus, with comments on the phylo-
genetic position of Campogramma
(Pisces: Carangidae), Comparative
revision of Scomberoides, Oligoplites,
Parona, and, by William F. Smith-
Vaniz and Jon C. Staiger, 185-256.

Hypericum pratense, 93

Hypogaleus, 389, 392, 393, 394, 403, 405

Hypoprion, 391, 405

Hysterocarpus traskii, 77, 82, 83

Iago, 389, 392, 393, 398, 405

Ichthyocampus annulatus, 139

(Indefatigable Island), Galapagos, Three new
land snails from Isla Santa Cruz, by
Allyn G. Smith, 7-24.

Indo-Pacific pipefishes, Five new, by Earl S.
Herald and John E. Randall, 121-140.

[Proc. 4TH SER.

Indo-west Pacific, with a synopsis of the
subfamily Synanceiinae, Two new
stonefishes (Pisces: Scorpaenidae) from
the, by William N. Eschmeyer and
Kaza V. Rama Rao, 337-382.

Indo-west Pacific, with comments on
Scorpaenodes muciparus (Alcock),
Two new _ scorpionfishes (genus

Scorpaenodes) from the, by William N.
Eschmeyer and K. V. Rama Rao,
55-64.
Inimicus, 307, 308, 339
didactylum, 288
Ischnochiton petaloides, 501

Iselica, 29
kochi, 29
Isla Santa Cruz (Indefatigable Island),
Galapagos, Three new land snails

from, by Allyn G. Smith, 7-24.
Isogomphodon, 391, 405
Iwamoto, Tomio, Nezumia (Kuronezumia)
bubonis, a@ new subgenus and species
of grenadier (Macrouridae: Pisces)
from Hawai and the western North
Atlantic, 507-516.

Jaegeria hirta, 101

key, based on scales, to the families of native
California freshwater fishes, A, by
Richard W. Casteel, 75-86.

(Kuronezumia) bubonis, a new subgenus and
species of grenadier (Macrouridae:
Pisces) from Hawaii and the western
North Atlantic, Nezumia, by Tomio
Iwamoto, 507-516.

Lamiopsis, 392, 405

Lavinia
exilicauda exilicauda, 76
exilicauda harengus, 76

Leiaster leachii, 321, 322

Leptocharias, 390, 404, 405

Leptograpsus,
ansoni, 323
variegatus,

330, 331

Leptosynanceia, 337, 339, 340, 370, 375, 377
asteroblepa, 341, 351, 356, 372, 375, 376
greenmani, 375

312, 315, 323, 324.e¢2e

melanostigma, 370

Vor. XXXIX]

Lichia, 228

albacora, 222, 223, 248

glauca, 246

quiebra, 221, 249

tolooparah, 205, 207, 250
Linsley, E. G., see Chemsak, John A.
Liomera, 313

monticulosa, 318

rugata, 315, 318, 329, 330

rugatus, 318
Liostephania, 546

species 1 Schrader, 540, 54.6
Lithodesmium, 546

californicum, 525, 529, 546

minusculum, 525, 529, 54.6
Lophozozymus, 313

dodone, 315, 319, 329, 330
Loripinus (Pegophysema) spherica, 33
Loxodon, 391, 392, 405
Lucina spherica, 33
Lupa pubescens, 316
Lycus, 144

sallei, 144
Lygaeidae of the Galapagos Islands (Hemip-

tera: Heteroptera), The, by Peter D.
Ashlock, 87-103.

Lyropecten (Nodipecten) magnificus, 31
Lysodermus, 339

Macrora, 546
stella, 525, 531, 546

(Macrouridae: Pisces) from Hawaii and the
western North Atlantic, Nezumia
(Kuronezumia) bubonis, a new sub-
genus and species of grenadier, by
Tomio Iwamoto, 507-516.

Malacocephalus, 507, 508, 509

Malea ringens, 29

Mandra, Highoohi, see Mandra, York T.

Mandra, York T., A. L. Brigger, and
Highoohi Mandra, Chemical extraction
techniques to _ free _ fossil _ silico-

flagellates from marine sedimentary
rocks, 273-284.

Mangelia, 29

hancocki, 29

Marianas Islands, A new lizard of the genus
Emoia (Scincidae) from the, by
Walter C. Brown and Marjorie V. C.
Falanruw, 105-110.

INDEX 571

McCosker, John E., Two new genera and two
new species of western Pacific snake-
eels (Apodes: Ophichthidae), 111-120.
Mecas, 141, 142, 143, 144, 145, 146, 147, 148,
159, 161, 169, 174, 179, 180
albovitticollis, 179
ambigena, 143, 149, 173, 174, 177
bicallosa, 142, 143, 149, 172, 173, 178
brevicollis, 179
cana, 141, 142, 177, 178
cana cana, 149, 178
cana saturnina, 143, 145, 149, 179
cineracea, 142, 143, 148, 165, 166, 173,

178
cinerea, 141, 142, 143, 144, 149, 167,
168, 169

cirrosa, 143, 149, 171, 172

confusa, 142, 143, 145, 148, 163, 164,
165, 178

discovittata, 170

(Dylobolus), 148, 153

(Dylobolus) rotundicollis, 153, 154, 156,
158

femoralis, 141, 143, 144, 148, 165, 166

humeralis, 143, 149, 172

inornata, 142, 143, 145, 146, 163, 167, 179

laminata, 159, 161

laminata discoimpunctata, 161

laminata discopunctata, 161

laminata rufobasalis, 161

laticeps, 155, 157

laticeps mediopunctata, 155, 157

laticeps sutureflave, 155, 157

laticeps vitticollis, 155

marginella, 141, 143, 148, 162, 174

marmorata, 180

(Mecas), 148, 159

menthae, 143, 146, 149, 174, 175, 176,
177

mexicana, 155, 157

obereoides, 143, 144, 148, 159, 160, 161

(Pannychis), 148, 149

(Pannychis) sericea, 150, 151, 152

pergrata, 141, 143, 144, 149, 169, 170

pergrata semiruficollis, 170

pseudambigena, 173, 174

rotundicollis, 143, 144, 148, 153

rotundicollis mexicana, 155

rotundicollis ruficollis, 155

rubripes, 167, 169

on
~r
re)

rubripes callosoreducta, 167, 169
ruficollis, 153, 157
ruficollis mediomaculata, 153, 157
saturnina, 142, 145, 179
senescens, 142, 167, 169
sericea, 143, 144, 148
vitticollis, 155, 157
Mecas Leconte (Coleoptera: Cerambycidae),
The genus, by John A. Chemsak and
E.G. Linsley, 141-184.
Mediaria, 54.6
splendida, 525, 531, 532, 546
Megapitaria, 28
aurantiaca, 28
squalida, 28
Melosira, 54.6
sulcata, 546
Menippe parvulus, 318
Metrarga, 96
Miconia, 19, 20
Microdonophis, 113
Micrognathus, 137, 139
(Anarchopterus), 137
boothae, 139
brachyrhinus, 137
brevirostris, 139
brocki, 139
edmonsoni, 139
(Micrognathus), 137
(Minyichthys), 137, 138
(Minyichthys) brachyrhinus, 138
(Minyichthys) myersi, 137, 138, 138,
139
Microsyngnathus, 136
Minous, 308, 339
Mitra
gausapata, 29
lens, 29
Modiolus capax, 28
Modulus cerodes, 29
Mohole Drilling, Guadalupe Site, Revised
diatom stratigraphy of the Experi-
mental, by Hans-Joachim Schrader,
517-562.
(Mollusca: Amphineura: Chitonidae), A new
Hawaiian chiton, Rhyssoplax linsleyi,
by Glenn E. Burghardt, 501-506.
(Mollusca: Pelecypoda) from the Galapagos
Islands, Description of a new species
of Chlamys, by Leo G. Hertlein, 1-6.

CALIFORNIA ACADEMY OF SCIENCES

[Proc. 4TH SER.

Mugil cephalus, 77, 80, 83
Muscular anatomy of the forelimb of the
sea otter (Enhydra lutris), by L. D.
Howard, 411-500.
Mustela, 390
Mustelus, 257, 258, 263, 264, 269, 270, 389,
390, 394, 405
dorsalis, 258
felis, 393
henlei, 257, 258, 264
megalopterus, 258, 264
nigropunctatus, 258, 264, 392, 393
Mylopharodon conocephalus, 76, 79, 82

Naesiotus, 70, 11, 16, 21, 22
eavagnaroi, 12, 14, 16, 17
darwini, 70, 12
deroyi, 9, 10, 12, 13, 20
duncanus, 74, 17
jervisensis, 17
lycodus, 12, 19, 20
ochsneri, 12
rabidensis, 70, 12
sealesiana, 17, 18, 19, 20, 27, 21, 22
sculpturatus, 10, 12
wolfi, 22

Nassarius nodicenctus, 29

Natica, 40
darwinii, 40
dubia, 40
solida, 40

Negaprion, 392, 405

Negogaleus, 392

Neotriakis, 257
sinuans, 257

Neptunus
pubescens, 316
tomentosus, 316, 317

Nerita
asperata, 41
oligopleura, 29, 34, 41

Nesoclimacias. 96

Nesocryptias, 96

Nesomartis, 88

Neverita, 40
reclusiana, 40

New barnacle records, (Cirripedia, Thorac-

ica), by Victor A. Zullo, Dea B. Beach,
and James T. Carlton, 65-74.

VoL. XXXIX]

New Guinea (Carcharhiniformes: Triakidae) ,
with a redefinition of the family
Triakidae and a key to Triakid genera,
Gogolia filewoodi, a@ new genus and
species of shark from, by L. J. V.
Compagno, 383-410.

new Hawaiian chiton, Rhyssoplax linsleyi
(Mollusca: Amphineura: Chitonidae),
A, by Glenn E. Burghart, 501-506.

new lizard of the genus Emoia (Scincidae)

from the Marianas Islands, A, by

Walter C. Brown and Marjorie V. C.

Falanruw, 105-110.

species of the scorpionfish genus

Helicolenus from the north Pacific

Ocean, A, by Tokiharu Abe and

William N. Eschmeyer, 47-53.

Nezumia, 508, 509, 515

atlantica, 507, 509

burragei, 513, 514

darus, 513, 514

hebetatus, 513, 514

(Kuronezumia), 509, 513, 514, 515

(Kuronezumia) bubonis, 507, 509,
510, 511, 512, 513, 515

macronemus, 513, 514

new

Nezumia (Kuronezumia) bubonis, a@ new
subgenus and species of grenadier
(Macrouridae: Pisces) from Hawaii

and the western North Atlantic, by
Tomio Iwamoto, 507-516.
Ninus notabilis, 97
Nioche, 28, 34
zorritensis, 28, 34
Nitzschia, 54.6, 547, 551, 554
bureklia, 518, 525, 531, 546, 549, 552
californica, 525, 546
cylindrica, 525, 529, 546, 550
heteropolica, 547, 550
invisa, 547, 552, 553
jouseae, 524, 525, 526, 529, 547
kanayensis, 518, 547, 552
marina, 549
miocenica, 524, 525, 526, 529, 548, 550
moholensis, 518, 548, 552
porteri, 524, 525, 526, 531, 548, 550,
551, 553
praereinholdii, 525, 529, 531, 548, 549,
S52,
reinholdii, 553

INDEX

573

riedelia, 518, 525, 531, 532, 547, 549,
552
rolandii, 547, 549, 552
seiboldia, 518, 525, 549, 550
species A Schrader, 531
species 2 Schrader, 523, 549
species 3 Schrader, 550, 551
species 4 Schrader, 550, 553
species 5 Schrader, 553
species 6 Schrader, 552, 553
species 7 Schrader, 550, 554.
species 8 Schrader, 550, 554
Nodipecten magnificus, 31
Nofua, 342
Nysius, 88, 89, 91
baeckstroemi, 88
marginalis, 89, 94, 95
naso, 97
nubilis, 91
(Ortholomus), 91
(Ortholomus) naso, 89, 97
usitatus, 89, 90, 91, 94

Oberea bimaculata, 145
Odostomia
(Chrysallida) rinella, 29
(Menestho) aequisculpta, 29
(Miralda) galapagensis, 29
Oligoplites, 186, 188, 190, 191, 211, 212,
P13 214 905. 278, 218, 2194970122 1.
222. 228. D724 FI BIO, 34.2338 234,
236, 237, 238, 240, 241, 242, 244, 245,
247, 248, 249
africana, 246, 248
altus, 2135 2145 215 e2tgeee lO 220.5 2oe
223, 224, 225, 236, 247, 248, 249
inornatus, 215, 221, 222, 248
(Leptoligoplites), 213, 215
mundus, 213, 219, 223, 247, 249
palometa, 214, 215, 217, 218, 220, 225,
236, 247, 248, 249
rathbuni, 221, 249
refulgens, 213, 214, 215, 217, 219, 220,
233, 236, 247, 249
saliens, 214, 215, 217, 218, 218, 220, 223,
2329334 296.1241, 247-248, 249
saurus, 190, 213, 217, 220, 221, 222, 236,
247
saurus inornatus, 212, 214, 215, 219, 221,
222, 224, 233, 248

574

Saurus saurus, 214, 215, 218, 222, 232,
235, 237, 237, 238, 239, 242, 249
Oligoplites, Parona, and Hypacanthus, with

comments on the phylogenetic position
of Campogramma (Pisces: Carangi-
dae), Comparative revision of
Scomberoides, by William F. Smith-
Vaniz and Jon C. Staiger, 185-256.
Oliva, 29
Oncorhynchus
kisutch, 76
tshawytscha, 76
Oostethus, 122
Ophichthidae), Two new genera and two
new species of western Pacific snake-
eels (Apodes:, by John E. McCosker,
111-120.
Ophichthus, 113
Orcynus, 190
Orthaea, 98
insularis, 98
Orthodon, 83
microlepidotus, 76
Ortholomus, 88, 89, 91, 93
gibbus, 93
naso, 97
usingeri, 90, 91, 93, 94
Ostrea, 28
megodon, 28
palmula, 28

Pachybrachius, 89, 98
insularis, 98, 100
nesovinctus, 98, 99, 100, 101
pacificus, 101
vinctus, 101
Pachygrapsus, 324, 325
minutus, 324
planifrons, 324
transversus, 312, 315, 324, 330
Pacific Ocean, A new species of the scorpion-
fish genus Helicolenus from the north,
by Tokiharu Abe and William N.
Eschmeyer, 47-53.
Pacific snake-eels (Apodes: Ophichthidae),
Two new genera and two new species

of western, by John E. McCosker,
111-120.

Pacific, with a synopsis of the subfamily
Synanceiinae, Two new stonefishes

(Pisces: Scorpaenidae) from the Indo-

CALIFORNIA ACADEMY OF SCIENCES

[Proc. 4TH Serr.

west, by William N. Eschmeyer and
Kaza V. Rama Rao, 337-382.

Pacific, with comments on Scorpaenodes
muciparus (Alcock), Two new
scorpionfishes (genus Scorpaenodes)

from the Indo-west, by William N.
Eschmeyer and K. V. Rama Rao,
55-64.
Palaeoscomber, 190, 192
spinosus, 190, 211, 212, 212, 250
Pannychis, 141, 149
ducalis, 151, 153
sericeus, 151, 153
Paragaleus, 392, 405
Paraminous, 339
Parapterois, 307
Paratrachinotus, 243
Parona, 225) 226, 22/7; (241247524 seeoa ae
245, 247
sienata, 227, 22%, 232:2oo coarse
238, 239, 240, 241, 242, 247
Parona, and Hypacanthus with comments on
the phylogenetic position of Campo-
gramma, (Pisces:Carangidae), Com-
parative revision of Scomberoides,
Oligoplites, by William F. Smith-
Vaniz and Jon C. Staiger, 185-256.
Paropsis, 225
signatus, 225, 227
Pavona gigantea, 27
Pecten, 31
antillarum, 4
australis, 4
benedicti, 4
(Chlamys) coccymelus, 4
(Chlamys) nympha, 4
(Flabellipecten) sericeus, 1
(Lyropecten) magnificus, 31
madisonius, 4
magnificus, 31
(Oppenheimopecten) galapagensis, 1
(Pecten) slevini, 28
Pedipes angulatus, 29
Pegophysema, 33

Pelecypoda) from the Galapagos Islands,
Description of a new _ species of
Chlamys (Mollusca:, by Leo G.
Hertlein, 1-6.

Pelor, 339

Peloropsis, 287

VoL. XXXIX]

frondosus, 292, 295
xenops, 287, 292, 295
Persicula, 29
imbricata, 29
phrygia, 29
Phaenomonas, 116, 117
Phytoecia, 141
femoralis, 141, 147, 159, 165
pipefishes, Five new Indo-Pacific, by Earl S.
Herald and John E. Randall, 121-140.
(Pisces: Carangidae), Comparative revision
of Scomberoides, Oligoplites, Parona,
and Hypacanthus with comments on
the phylogenetic position of Campo-
gramma, by William F. Smith-Vaniz
and Jon C. Staiger, 185-256.
(Pisces:Scorpaenidae) from the Indo-west
Pacific, with a synopsis of the sub-
family Synanceiinae, Two new stone-
fishes, by William N. Eschmeyer and
Kaza V. Rama Rao, 337-382.
Plagusia, 312, 314
capensis, 328
chabrus, 328
dentipes, 312, 314, 315, 326, 328, 329,
331
depressa, 328
depressa depressa, 328
depressa immaculata, 329
depressa tuberculata, 329
gaimardi, 328
glabra, 328, 329
integripes, 315, 326, 327, 329
speciosa, 328, 329
tomentosa, 328
Pliocene fossils from Baltra (South
Seymour) Island, Galapagos Islands,
by Leo G. Hertlein, 25-46.
Pocillopora, 321, 322
damicornis, 321
danae, 321
diomedae, 321
Pogonichthys macrolepidotus, 76
Pogonophis, 113
Pogonoscorpius, 285, 287, 303, 306
sechellensis, 285, 287, 303, 304
seychellensis, 303
Polinices
dubius, 29, 38, 40
uber, 29

INDEX 575

Polycaulis uranoscopus, 373
Polycaulus, 339, 372
elongatus, 373
uranoscopus, 373
Polynices dubia, 40
Porites, 73
astreoides, 71, 72
californica, 72, 73
lobata, 73
Porthmeus, 228
argenteus, 228, 229, 230, 231
Portulaca, 91, 95, 96, 97
Portunus, 312
pubescens, 315, 316, 317, 329, 331
tomentosus, 317
Prionace, 391, 405
Pristipomoides microlepis, 57
Proscyllium, 257, 263, 268, 269, 390, 405
habereri, 257
venusta, 257
Prosopis, 179
Prosopium williamsoni, 76, 81, 81, 83
Protothaca
grata, 37
(Tropithaca), 28, 35, 37, 38
(Tropithaca) grata, 28
(Tropithaca) seymourensis, 28, 35, 38
Psammocora (Stephanaria), 27
Pseudoliomera
lata, 314, 319
remota, 312, 314, 315, 318, 319, 329
Pseudosynanceia, 337, 340, 369, 377
melanostigma, 341, 351, 356, 369, 370,
Sf
Pteroidichthys, 285, 287, 302, 303, 305, 306
amboinensis, 305, 306, 307
Pterois, 307
sphex, 307
Pteropelor, 285, 302, 303, 305
noronhai, 305, 306, 307
Ptychocheilus grandis, 76
Ptychognathus
easteranus, 312, 315, 325, 329
polleni, 325
Pyramidella
(Triptychus), 29
(Triptychus) olssoni, 29
(Voluspa), 29
Pyrene
castanea, 29

576 CALIFORNIA ACADEMY OF SCIENCES

fuscata, 29
haemastoma, 29
Pyrgoma, 72
Rama Rao, Kaza V., see
William N.
Randall, John E., see Herald, Earl S.
Raphiolepus, 190, 192
Revised diatom stratigraphy of the Experi-
mental Mohole Drilling, Guadalupe
Site, by Hans-Joachim Schrader,
517-562.
Rhaphoneis, 541, 554.
angularis, 525, 554
angustata, 525, 554
margaritalimbata, 525, 554
mediopunctata, 542
mediopunctata matraensis, 542
sacchalinensis, 525, 554.
tatsunokuchiensis, 543
Rhinichthys, 83
osculus klamathensis, 76
Rhinopias, 285, 287, 288, 289, 291, 295, 298,
301, 302, 303, 304, 306
aphanes, 285, 286, 288, 291
300, 301, 302, 307
argoliba, 285, 286, 288, 289, 290, 291,
292, 298, 302, 307
frondosa, 285, 286, 288, 291, 292, 295,
296, 296, 297, 298, 299, 300, 300, 302,
307
godfreyi, 285, 287, 303, 305, 306
xenops, 285, 286, 288, 290, 291, 292,
293, 294, 295, 296, 298, 302, 307
Rhinopias, with comments on related genera
and species, Two new species of the
genus, by William N.
Eschmeyer, Yoshitsugo Hirosaki, and
Tokiharu Abe, 285-310.
Rhizoprionodon, 392, 405
Rhizosolenia, 554
barboi, 525, 529, 531, 554.
hebetata, 525
hebetata hiemalis, 554.
miocenica, 525, 529, 531, 554.
praealata, 525, 554.
praebarboi, 525, 531, 532, 555
styliformis, 525, 555
Rhynchobatus, 394

Eschmeyer,

296, 298,

>]

scorpionfish

[Proc. 4TH Ser.

Rhyssoplax, 505
linsleyi, 501, 502, 503, 503, 504, 505
Rhyssoplax linsleyi (Mollusca: Amphineura:
Chitonidae), A new Hawaiian chiton,
by Glenn E. Burghardt, 501-506.
Richardsonius balteatus egregius, 77
Rissoina, 29
firmata, 29
signae, 29
Robinsonocoris, 94, 96
tingitoides, 88
Rouxia, 555
californica, 526, 529, 550,555
diploneides, 526, 555
moholensis, 518, 550, 555
naviculoides, 550, 555
species 1 Schrader, 555

Salmo
aguabonita aguabonita, 76
aguabonita whitei, 76
clarkii clarkii, 76
clarkii henshawi, 76
clarkii seleniris, 76
gairdnerii, 76
gairdnerii aquilarum, 76
gairdnerii gairdnerii, 76, 87
gairdnerii gilberti, 76
gairdnerii stonei, 76
Salvelinus malma parkei, 76
Salvia, 179
Saperda, 141, 142, 143, 147, 167, 173
bicallosa, 166, 172
cana, 141, 177, 178
cineracea, 166
cinerea, 142, 167
concolor, 142, 143
concolor unicolor, 143
inornata, 142, 143
pergrata, 141, 169
Scaeops sechellensis, 303
Scaevola, 110
Scalesia, 11, 12, 13, 16, 18, 19, 20, 22, 97
affinus, 97
gummifera, 97
helleri, 97
pedunculata, 11, 19
Schrader, Revised diatom
stratigraphy of the Experimental

Hans-Joachim,

Vor. XXXIX]

Mohole Drilling, Guadalupe site, 517-
562.

(Scincidae) from the Marianas Islands, A new
lizard of the genus Emoia, by Walter
C. Brown and Marjorie V. C.
Falanruw, 105-110.

Scoliodon, 391, 392, 404, 405

Scomber

aculeatus, 229, 230

amia, 228, 230
calcar, 218, 220, 223, 248
forsteri, 205, 207, 248
japonicus, 212

lysan, 205, 206, 249
maculatus, 205, 207, 249
madagascariensis, 194, 249
saliens, 218, 249

saurus, 213, 215, 221, 249

Scomberoides, 186, 187, 188, 190, 191, 192,
193, 195, 196, 197, 198, 199, 200, 201,
201, 202, 203, 204, 206, 212, 213, 215,
FAO. 231, 233, 234;.236).287, 238) 240,
241, 242, 243, 244, 245, 247, 248, 249,
250

commersonianus, 190, 191, 192, 194, 195,
196, 197, 198, 200, 200, 201, 202, 203,
204, 205, 206, 208, 232, 234, 236, 240
242, 247, 248, 249

formosanus, 191, 205, 207, 209, 210, 248

forsteri, 207

lysan, 190, 191, 193, 194, 195, 196, 197,
198, 200, 201, 202, 203, 204, 205, 206,
207, 208, 213, 233, 234, 236, 237, 240,
247, 248, 249, 250

moadetta, 191

noelii, 192, 249

orientalis, 191

oshimae, 205, 249

(Rhaphiolepis) , 190

saliens, 218

saltator, 192, 218, 249

sanctipetri, 191, 207

spinosus, 192, 211, 213, 241, 247, 250

tala, 191, 192, 194, 195, 196, 197, 198,
199, 200, 200, 201, 201, 202, 203, 204,
206, 209, 210, 233, 234, 236, 241, 247,
248, 249, 250

tol, 191, 194, 195, 196, 197, 198, 199, 200,
200, 201, 201, 202, 203, 204, 206, 209,
MONA 11 212,213, 232).233, 236, 237,

INDEX 577

238, 239, 247, 248, 250
toloo, 191, 201
tolooparah, 191, 206, 207, 247

Scomberoides, Oligoplites, Parona, and
Hypacanthus with comments on the
phylogenetic position of Campo-
gramma, (Pisces: Carangidae), Com-
parative revision of, by William F.
Smith-Vaniz and Jon C. Staiger,
185-256.

Scorpaena

alepidota, 352, 353
Bicapillata, 359
bicirrata, 359
Brachiata, 359
brachion, 359
frondosa, 287, 295
horrida, 341, 352, 353
monstrosa, 352

Scorpaenidae) from the Indo-west Pacific,
with a synopsis of the subfamily
Synanceiinae, Two new stonefishes
(Pisces:, by William N. Eschmeyer and
Kaza V. Rama Rao, 337-382.

Scorpaenodes, 55, 56, 58

investigatoris, 57, 58, 64
muciparus, 55, 56, 57, 58, 59
smithi, 57, 58, 59, 60, 62, 63,
steinitzi, 57

tribulosus, 55, 56, 58, 59
varipinnis, 57

Scorpaenodes from the Indo-west Pacific,
with comments on Scorpaenodes
muciparus (Alcock), Two new scor-
pionfishes (genus, by William WN.
Eschmeyer and K. V. Rama Rao,
55-64.

Scorpaenodes muciparus (Alcock), Two new
scorpionfishes (genus Scorpaenodes)
from the Indo-west Pacific, with
comments on, by William WN.
Eschmeyer and K. V. Rama Rao,
55-64.

scorpionfish genus from the
north Pacific Ocean, A new species of
the, by Tokiharu Abe and William N.
Eschmeyer, 47-53.

Helicolenus

scorpionfish genus Rhinopias, with comments
on related genera and species, Two new
species of the, by William WN.

578 CALIFORNIA ACADEMY OF SCIENCES

Eschmeyer, Yoshitsugo Hirosaki, and
Tokiharu Abe, 285-310.

scorpionfishes (genus Scorpaenodes) from the
Indo-west Pacific, with comments on
Scorpaenodes muciparus (Alcock),
Two new, by William N. Eschmeyer
and K. V. Rama Rao, 55-64.

Scylliogaleus, 389, 390, 393, 405

sea otter (Enhydra lutris), Muscular anatomy
of the forelimb of the, by L. D.
Howard, 411-500.

Sebastes, 48, 51

muciparus, 58
Sebastiscus, 48, 51, 53
marmoratus, 292, 295, 300

Sebastodes, 51

(Selachitz: Carcharhinidae), Ctenacis and
Gollum, two new genera of sharks, by
L. J. V. Compagno, 257-272.

Selar, 244

Seleastes daector, 359

Selene, 244

Seriola, 246, 247

shark from New Guinea (Carcharhiniformes:

Triakidae), with a redefinition of the

family Triakidae and a key to Triakid

genera, Gogolia filewoodi, a new

genus and species of, by L. J. V.

Compagno, 383-410.

(Selachii: Carcharhinidae), Ctenacis
and Gollum, two new genera of, by
L. J. V. Compagno, 257-272.
silicoflagellates from marine sedimentary

rocks, Chemical extraction techniques
to free fossil, by York T. Mandra,
A. L. Brigger, and Highoohi Mandra,
273-284.

Smith, Allyn, G., Three new land snails from
Isla Santa Cruz (Indefatigable Island),
Galapagos, 7-24.

Smith-Vaniz, William F., and Jon C. Staiger,
Comparative revision of Scomberoides,
Oligoplites, Parona, and Hypacanthus
with comments on the phylogenetic
position of Campogramma (Pisces:
Carangidae), 185-256.

snails, from Isla Santa Cruz (Indefatigable
Island), Galapagos, Three new land,
by Allyn G. Smith, 7-24.

sharks

[Proc. 4TH Srp.

snake-eels (Apodes: Ophichthidae) Two new
genera and two new species of western
Pacific, by John EE. McCosker,
111-120.
Solagmedens, 246
(South Seymour) Island, Galapagos Islands,
Pliocene fossils from Baltra, by Leo G.
Hertlein, 25-46.
Spondylus, 31
Stenostola, 141
gentilis, 170
pergrata, 170
saturnina, 141, 179
Stephanopyxis, 556
turris, 526, 556
stonefishes (Pisces: Scorpaenidae) from the
Indo-west Pacific, with a synopsis of
the subfamily Synanceiinae, Two new,
by William N. Eschmeyer and Kaza V.
Rama Rao, 337-382.
Stoschia, 541
admirabilis, 541
paleacea, 541, 543
Strombina gibberula, 29
Synanceia, 307, 337, 339, 340, 341, 342, 343,
357, 359, 363, 364, 365, 372, 375
alula, 341, 347, 348, 349, 350, 350, 351,
352, 350, SH7
asteroblepa, 375
bicapillata, 359
brachio, 341, 359
breviceps, 373
elongata, 373, 374
elongatus, 372, 373
erosa, 363, 364
grossa, 352
horrida, 337, 340, 341, 343, 347, 351,
352, 354, 355, 356, 357
nana, 341, 343, 344, 345, 346, 347, 351,
356, 363
(Nofua) platyrhynchus, 340
platyrhynchus, 340, 341, 342, 347, 351,
356, 357, 358
sanguinolenta, 359
thersites, 359, 361
trachynis, 352
verrucosa, 337, 339, 340, 341, 343, 347,
348, 351, 356, 357, 359, 360, 363
Synanceichthys, 339, 340, 341
verrucosus, 340, 359

VoL. XXXIX] INDEX

Synanceiinae, Two new stonefishes (Pisces:
Scorpaenidae) from the Indo-west
Pacific with a synopsis of the sub-
family, by William N. Eschmeyer and

579

excentrica, 526, 556

nativa, 526, 529, 538, 557
oestrupii, 526, 536, 538, 557
praeconvexa, 526, 538, 557

Kaza V. Rama Rao, 337-382.

Synanceja, 341, 342, 352

horrida, 352

(Nofua) platyrhynchus, 357

trachynis, 352

uranoscopa, 372

verrucosa, 359
Synanchia, 341, 342, 363, 364

erosa, 342, 363, 364
Synancia, 343

elongata, 373
Synancidium, 341, 343

erosum, 364

horrida, 343

horridum, 352
Synancya elongata, 373
Synancydium, 341
Synchiropus sechellensis, 303
Synedra, 556

jouseana, 526, 529, 531, 550, 556
Syngnathus, 133, 135, 136

balli, 135

banneri 135, 136, 136

caldwelli, 133, 134, 135

dactyliophorus, 130

maxweberi, 127

Taenianotis, 307
triacanthus, 307
Tagelus, 28
dombeii, 28
Tectarius galapagensis, 29
Tegula
aureotincta, 39
forbesi, 29, 36, 37, 38
rugosa, 39
snodgrassi, 29
Thalamita, 313, 315, 329, 330
Thalassionema, 556
claviformis, 526, 529, 531, 556
hirosakiensis, 550, 556
nitzschioides, 526, 550, 556
Thalassiosira, 556
antiqua, 526, 556
convexa, 524, 525, 526, 529, 538, 556
decipiens, 526, 556

species A (Burckle), 526, 531, 538, 556
species 2 Schrader, 538, 557
usatchevii, 526, 529, 540, 557
Thalassiothrix, 557
longissima, 526, 557
Thaleichthys pacificus, 76
Thoracica), New barnacle records, (Cirri-
pedia, by Victor A. Zullo, Dea B.
Beach, and James T. Carlton, 65-74.
Three new land snails from Isla Santa Cruz
(Indefatigable Island), Galapagos, by
Allyn G. Smith, 7-24.
Thynnus moluccensis, 205, 207, 249
Trachicephalus, 337, 339, 340, 342, 372, 373,
377
elongatus, 342, 373
uranoscopus, 341, 351, 356, 372, 373, 374,
375
Trachinotus, 191, 241, 243, 244, 245, 247
ovatus, 241
stilbe, 243
Trachonurus, 509
Trachycephalus, 372
uranoscopus, 373
Trachyurops, 244
Trapezia, 312, 314, 321, 322
areolata, 315, 320, 329, 330
cymodoce, 315, 321, 322, 329, 330
cymodoce ferruginea, 322
danae, 321
danai, 315, 321, 322, 329
digitalis, 322
ferruginea, 312, 315, 321, 322, 329, 330
ferruginea areolata, 320
hirtipes, 321
maculata, 315, 321
miniata, 322
reticulata, 320
wardi, 322
Triaenodon, 389, 390, 391, 392, 405
Triakidae), with a redefinition of the family
Triakidae and a key to Triakid genera,
Gogolia filewoodi, a new genus and
species of shark from New Guinea
(Carcharhiniformes:, by L. J. V.
Compagno, 383-410.

580 CALIFORNIA ACADEMY OF SCIENCES

Triakis, 257, 258, 263, 264, 269, 270, 389,
390, 392, 394, 405
acutipinna, 258, 264, 392
attenuata, 257, 258, 263, 264
barbouri, 257
fehlmanni, 257, 258, 263
habereri, 257
henlei, 257, 264
leucoperiptera, 257
maculata, 258, 264, 393
megaloptera, 392
scyllium, 257, 258, 264, 265, 392
semifasciata, 258, 262, 264, 393
venusta, 257
Triceratium, 557
antiquum, 536, 557
cinnamomeum, 526, 536, 557
condecorum, 526, 557
Trichophasia, 342, 372
Triphora, 29
galapagensis, 29
panamensis, 29
Trivia
pacifica, 30
radians, 30, 32 .
Turbo, 40
agonistes, 30, 36, 39
caboblanquensis, 40
vermiculosus, 30, 38, 40
Turbonilla (Chemitzia) houseri, 30
Turritella, 41
alturana, 42
broderipiana, 41, 42
broderipiana marmorata, 30, 38, 41, 42
(Broderiptella) , 42
gonostoma, 41
marmorata, 41, 42
Two new genera and two new species of
western Pacific snake-eels (Apodes:
Ophichthidae), by John E. McCosker,

[Proc. 4TH Serr.

Two new scorpionfishes (genus Scorpaenodes)
from the Indo-west Pacific, with com-
ments on Scorpaenodes muciparus
(Alcock), by William N. Eschmeyer
and K. V. Rama Rao, 55-64.

Two new species of the scorpionfish genus
Rhinopias, with comments on related
genera and species, by William N.
Eschmeyer, Yoshitsugo Hirosaki, and
Tokiharu Abe, 285-310.

Two new stonefishes (Pisces: Scorpaenidae)
from the Indo-west Pacific, with a
synopsis of the subfamily Synanceiinae,
by William N. Eschmeyer and Kaza V.
Rama Rao, 337-382.

Uranoblepus, 372

Uranoscopus
adhoesipinnis, 373
indicus, 373

Ventrifossa, 507, 508

Verbena, 93

Verbesina, 147
encelioides, 147

Vermicularia eburnea, 30

Volvarina taeniolata, 30

Xanthium, 145, 147, 179
spinosum, 147

Xyonysius, 89,97 .
californicus, 97
naso, 92, 94, 95, 97

Xyrauchen texanus, 76

Yozia, 140
Zullo, Victor A., Bea B. Beach, and James T.

Carlton, New barnacle records (Cirri-
pedia: Thoracica), 65-74.

Line 3 from bottom of table 1: for “Mugil cephalis” read “Musgil cephalus.”

111-120.
ERRATA
Page 33. Line 19 from top: for “Lissophaira” read “Lissosphaira.”
Page 77.
Page 263. Lines 23 and 24 should appear in reverse order.

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