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Prior to November 30, 1973, publications of the Natural History Museum have appeared under various titles — Leaflet
Series, Museum Graphic , Education Series , History Series, Science Series, Study Guides, Contributions in History, History
Bulletin, Science Bulletin, unnumbered catalogs of exhibitions, and other miscellaneous publications.

The publication program of the Natural History Museum continues to publish a series of reports that give accounts of
new discoveries in history and science as well as new advances in museum knowledge. The theme of the Natural History
Museum serials will emphasize basic research through the continuing active series:

Contributions in Science
Education Series
History Series
Science Series

In these series, the Contributions in Science are comprehensive papers of imperial octavo size whereas the Series are
papers of variable lengths of quarto or larger size. Issues in each serial are numbered separately and consecutively, and
report on the research and collections of the Natural History Museum, or of research activity of professional colleagues.
The Museum’s publications are distributed through mailing lists to libraries, universities, and similar institutions throughout

the world.

Acceptance of manuscripts will be determined by the significance of new information. Priority will be given to
manuscripts by staff members. All manuscripts must be recommended by the curator in charge of each discipline or by the
Editorial Board. Peer judgment will be accomplished through the various divisions of the Museum and by outside

reviewers.

All communications concerning submission of manuscripts should be directed to the Editor, Museum Publications; all
communications concerning exchange of publications should be sent to the Research Library; all communications
concerning purchase of publications should be sent to the Book Shop, Natural History Museum of Los Angeles County, 900

Exposition Boulevard, Los Angeles, California 90007.

Edward Qstermeyer
Editor

Printed in the United States of America by International Printing Services. Inc.

Number 301
March 16, 1979

CONTRIBUTIONS IN SCIENCE

NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY

A NEW FISH OF THE GENUS PHALLICHTHYS
(FAMILY POECILIIDAE) FROM COSTA RICA

By William A. Bussing

Published by the NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY • 900 EXPOSITION BOULEVARD • LOS ANGELES, CALIFORNIA 90007

The Science Bulletin and Contributions in Science of the Natural History Museum of Los
Angeles County were merged into a single imperial octavo serial, retaining the name Contribu-
tions in Science and beginning with Number 301.

This serial has been newly formatted for maximum use of typography and illustrations per page,
and sized for maximum use of paper. All photography has been produced utilizing a 200-line
screen for detail.

Since most institutions rebind Contributions in Science collectively into hardbound volumes,
and many researchers using them desire to photocopy material, two new features have been incor-
porated into the new format. A suggested citation line provides the abbreviated serial title, year,
number of publication, and total pagination, and appears in the lower left corner of each page; and
pages of Contributions in Science have been printed off-center to provide a two-inch gutter
between adjacent pages for easier accessibility in photocopying.

ISSN; 0459-8113

Suggested Citation: Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 301:1-8.

Contributions in Science are articles in the earth and life sciences, presenting results of original research in Natural
History. Science Bulletin (Numbers 1-30; 28 June 1965 to 10 April 1978) and Contributions in Science Numbers 1-300; 23
January 1957 to 16 June 1978) were merged into a single imperial octavo serial beginning with Number 301.

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 301:1-8.

A NEW FISH OF THE GENUS PHALLICHTHYS
(FAMILY POECILIIDAE) FROM COSTA RICA1

By William A. Bussing2

ABSTRACT: A new species, Phallichthys quadripunctatus, is described on the basis of 48 specimens from the
Baja Talamanca lowlands of Costa Rica. The new form is shown to be closely allied to another congener, P.
tico, of northern Costa Rica and southern Nicaragua. Both species are small, have sinistrally asymmetrical
gonopodia, and each bears a curved membranous swelling on the terminal element of gonopodial ray 4a. P
quadripunctatus was collected only in shallow streams and standing waters of the Rio Sixaola basin. Four
species now comprise the Central American genus Phallichthys.

Phallichthys and four other genera with much in common
morphologically and geographically, were placed in the tribe
Heterandriini by Rosen and Bailey (1963). Most members of the
tribe occur in Middle America, a few are found in western South
America, one in southwestern United States and one in Florida.
The species of Phallichthys are largely restricted to the Atlantic
versanf of Middle America (Fig. 1), although P. amates (Miller)
was apparently introduced by man to the Valle Central (Rio
Tarcoles headwaters) and was recently collected at four sites on
the Pacific slope in the Tempisque and Bebedero drainages of
northwestern Costa Rica (Bussing and Lopez 1977). Shortly
after Rosen and Bailey (1963) revised the family Poeciliidae,
Bussing (1963) described Phallichthys tico from northern Costa
Rica, and subsequently reported on Nicaraguan material of the
same species (1976). The present description of P. quadripunc-
tatus, also from the Atlantic slope, increases the number of Costa
Rican forms to three species. A fourth member of the genus, P.
fairweatheri, inhabits northern Guatemala and part of Beliz
(Rosen and Bailey 1959).

Phallichthys quadripunctatus new species

Figures 2, 3, 4

HOLOTYPE: LACM 36018- 1; a mature male 14.6 mm SL, collected
by William, Myrna and Eric Bussing in a small tributary of the Rio
Sixaola, 0.5 km NE of Chase on road between Puerto Viejo and Bratsi,
Limon Province, Costa Rica, on 4 October 1975.

PARA TYPES: LACM 36018-2; 9 males 12.8-15.5 mm SL and 7
females and immatures 12.5-20.7 mm SL, same data as the holotype.
USNM 216559; 3 males 11.7-15.0 mm SL and 2 females 14.6 and
15.9 mm SL, same data as the holotype. UCR 897-2; 10 males (2
cleared and stained) 10.3-15.4 mm SL and 8 females and immatures
8.8-21.0 mm SL. same data as the holotype. UCR 1017-1; 8 females
and immatures 7.2-32.7 mm SL, collected by William Bussing and
ichthyology class in a small tributary of the Rio Sixaola, 1.2 km SW of
Cruce Bribri on road between Puerto Viejo and Bratsi, Limon Province,
Costa Rica, on 18 September 1976.

MATERIAL AND METHODS: The type specimens are depos-
ited in the Natural History Museum of Los Angeles County
(LACM), the Museo de Zoologia, Universidad de Costa Rica

(UCR), and the National Museum of Natural History (USNM).

All body measurements refer to standard length (SL) in milli-
meters (mm), and were made according to the methods described
in Rosen and Bailey (1959). All proportional measurements in
the text are expressed as times in SL or times in head length.
Body proportions as thousandths of SL and meristic counts ap-
pear in Table 1. The nomenclature of the gonopodial system is
that used in Rosen and Bailey (1959).

DIAGNOSIS: This small Phallichthys is distinguished from its
congeners by the presence of typically four conspicuous spots on
each side of the body (Fig. 2), and an intermediate number of gill
rakers on the first gill arch (11-17, usually 13-16 on large
adults). Males of both the new species and Phallichthys tico dif-
fer from other Phallichthys in possessing a very long terminal
segment of gonopodial ray 4a (4 or 5 times longer than wide and
3 or 4 times longer than wide, respectively), which extends
beyond the other gonopodial elements, and bears a fleshy pad
dorsally and a membranous hook at a right angle to its axis (Fig.
3).

DESCRIPTION: Body moderately robust, laterally com-
pressed. Body depth greatest at pelvic-fin origin, and less in
mature males than in mature females; greatest body depth in SL
3. 2-3. 6 times in males and 2. 8-3. 4 times in females. Predorsal
profile slightly convex in both sexes; profile angular at dorsal-fin
origin. Postdorsal profile slightly concave. Ventral profile of
body rounded anteriorly; postanal profile straight. Caudal
peduncle depth in SL 5. 2-5. 7 times in males and 5. 5-6.0 times
in females.

'Review Committee for this Contribution:
Reeve M. Bailey
Robert J. Lavenberg
Donn E. Rosen

"Escuela de Biologia. Universidad de Costa Rica, Ciudad Universitaria,
Costa Rica; Research Associate in Ichthyology, Natural History Museum
of Los Angeles County; and Consejo Nacional de Investigaciones Cien-
tificas y Tecnologicas (CONIC1T) de Costa Rica.

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 301:1-8.

Bussing: Costa Rican Fish

Figure 1. Map of Central America showing known distributions of the species of Phallichthys. Range of P . fairweatheri and Guatemalan
distribution of P. amates from Rosen and Bailey (1963). Honduran distribution of P. amates after Martin ( 1972).

Head length 3. 1-3.7 times in SL; head width 4. 1-4.7 times in
SL. Eyes slightly below dorsal head profile; horizontal orbit
diameter 2.3- 3.3 times in head length. Least width of bony
interorbital 1.9- 2. 6 times in head length. Snout length always
shorter than orbit diameter, 2. 9-3. 7 times in head length. Post-
orbital distance 2.2- 2.5 times in head length.

Mouth opening dorsally; overall width 2. 1-2.9 times in head
length. Two rows of incurved teeth on each jaw. Teeth of outer
row of each jaw spatulate, that is, with expanded flattened tips,
apparently adapted to scraping surfaces. Teeth of inner series
sharp-pointed and conical, much shorter than teeth of outer

series. Teeth of both series gradually reduced in length toward
lateral edges of jaws; premaxillary teeth of outer series not pres-
ent at extreme edges of jaw.

Dorsal-fin origin at highest point on body; directly over anal-
fin origin in females, slightly posterior to origin of gonopodium
of males. Predorsal distance in SL 1.8- 1.9 times in males and
1.7- 1.8 times in females. Distance from dorsal-fin origin to
caudal-fin base in SL 2.0-2. 1 times in males and 2. 1-2.2 times
in females. Depressed length of dorsal fin in SL 3. 2-3. 6 times in
males and 3. 6-4.0 times in females. Dorsal-fin rays usually 9,
range 8(1) and 9 (21), frequency in parentheses.

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 301:1-8.

Bussing: Costa Rican Fish

TABLE 1

Meristics and proportions in thousandths of standard length of the holotypes and 10 male and 10 female
paratypes each of Phallichthys quadripunctatus and Phallichrhys tico.

Holotype

quadripunctatus

Paratypes

Holotype

P. tico

Paratypes

Dorsal rays

8-9

7-9

Anal rays

9-10

Pectoral rays

1 1

11-12

1 1

10-12

Pelvic rays

5-6

Caudal rays

12-14

Scales in lateral series

26-27

Total gill rakers

1 1-17

—

10-12

males

females

males

females

Standard length (mm)

14.6

12.0-15.2

17.3-32.7

17.4

13.7-17.6

17.7-32.9

Body, greatest depth

301

275-316

298-352

298

255-294

282-343

Caudal peduncle, least depth

185

175-191

167-182

183

154-180

158-173

Dorsal origin to snout tip

548

540-559

563-598

528

470-573

571-602

Anal origin to mand. symphysis

548

525-558

590-638

557

496-580

621-671

Dorsal origin to caudal base

493

483-507

464-48 1

465

422-483

431-458

Anal origin to caudal base

541

517-560

416-459

522

446-528

395-432

Head length

301

293-320

269-306

298

267-309

264-305

Head width

219

212-230

214-281

195

174-198

192-215

Snout length

87-99

92-99

83-105

88-107

Orbit length

1 16

113-133

83-116

109

96-111

75-101

Postorbital length of head

130

1 19-133

1 19-139

137

1 13-141

1 15-141

Interorbital, bony width

144

117-145

138-155

143

109-148

136-157

Mouth, over-all width

123

108-121

1 16-138

1 14

96-129

103-132

Dorsal fin, depressed length

288

275-309

249-280

264

244-27 1

224-257

Anal fin, depressed length

445

431-483

231-267

408

375-457

203-237

Caudal fin length

384

358-392

300-376

321

285-341

249-331

Pectoral fin length

267

263-280

229-262

224

193-228

181-235

Pelvic fin length

260

225-268

182-214

183

145-194

136-171

Anal-fin rays usually 10, range 9(1) and 10 (10). Preanal dis-
tance in SL 1.8- 1.9 in males and 1.6- 1.7 times in females.
Distance from anal-fin origin to caudal-fin base in SL 1 .8- 1.9 in
males and 2. 2-2. 4 times in females. Depressed length of anal fin
of females 3.8-4. 1 times in SL; depressed length of gonopodium
in males 2. 1-2.3 times in SL. Posterior border of anal fin in
females truncate.

Pectoral-fin rays usually 1 1, range 11 (21) and 12 (1). Tip of
fins extending posteriorly to origin of pelvic fins of females and
beyond origin of gonopodium of males; length of pectoral fin in
SL 3. 6-3. 8 times in males and 3. 8-4. 3 times in females.

Pelvic-fin rays usually 6, range 5(1) and 6 (21). Tip of fins in
both sexes extending posterior to origin of anal fin. Length of
pelvic tin in SL 3. 7-4. 4 times in males and 4. 7-5. 8 times in
females. Caudal-fin rays usually 13, range 12 (1), 13 (19) and 14
(2). Length of caudal fin in SL 2.6- 2.8 times in males and
2. 7-3. 3 times in females.

Total number of gill rakers increasing ontogenetically. Eleven
specimens of 12- 16 mm SL, including all males examined, with
11 ( 1 ), 12(1), 13 (5) and 14 (4) gill rakers on first arch. Eleven
larger specimens (17.0-32.7 mm SL), all females, with 15 (3),
16 (6) and 17 (2) gill rakers on first arch Scales in lateral series
usually 26, range 26 (20) and 27 (2). Vertebrae 29 on two
cleared and stained paratypes. Long, pointed urogenital (?)
papilla present on about 50 percent of large females; apparently
non-tubular, but serving to divide genital and urinary apertures.

GONOPODIAL SYSTEM: Gonopodium of mature males
sinistral, i.e. rays 3, 4 and 5 of anal fin forming a shallow
groove opening to left side of body. Ray 3 deep and compressed;
proximal segments sinistrally incurved to form ventral border of
trough; six subdistal segments of ray 3(1) forming long thin
spines; two minute terminal segments without spines (Fig. 3).
Three or four corresponding segments of ray 3(r) also produced
into spines.

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 301:1-8.

Bussing: Costa Rican Fish

Right and left halves of ray 4 symmetrical. Ray 4a slender,
tapering to tip; terminal segment much longer than subterminal
ones (4 or 5 times longer than wide). Terminal segment of ray 4a
forming tip of gonopodium; a fleshy swelling on dorsal border of
last segment from which hangs a pointed membranous knob.
Distal portion of ray 4p also slender; subterminal segments of
each half bearing long retrorse serrae; terminal two segments
without serrae.

Ray 5 symmetrical and composed of an anterior half which
forms bottom, and posterior half which forms sides, of a square-
bottomed groove on posterior surface of gonopodium. Both
halves of ray tapering at tip, terminal segments of ray 5a extend-
ing beyond end of ray 5p. Right and left halves of rays 6 to 10
symmetrical; rays 6, 7 and 8 constricted near middle of ray and
split into anterior and posterior elements distal ly; rays 9 and 10
split only into right and left halves.

Ligastyle of moderate length. Gonapophysis I curved antero-
ventrally, tip lying between lateral wings of primary gonac-
tinostal complex; a pair of long, slender uncini arising near center
of gonapophysis and curving posteroventrally (Fig. 4). Gona-
pophysis II projecting anteroventrally between preceding uncini,
and bearing a pair of posteroventrally projecting uncini. Gona-
pophysis III projecting anteroventrally, shorter and stouter than
others; a pair of short, pointed uncini arising near center of gona-
pophysis and directed posteriorly.

Gonactinost 1 free, pointed tip directed anterodorsally. Pri-
mary gonactinostal complex (gonactinosts 2, 3 and 4) with lateral
flanges along entire length of gonactinost 4; these lateral pro-
cesses flaring broadly and curving posteriorly on distal third of
primary complex. Gonactinost 5 lying between lateral processes
of primary complex. Gonactinosts 6 to 9 free and slender with
symmetrical or dextral lateral processes. Gonactinosts 6 and 7
slightly expanded laterally at tips, gonactinost 7 also with a dex-
tral lateral flange subterminally; gonactinosts 8 and 9 with tri-

FtGURE 2. Phallichthys quadripunctatus , new species from the
Rio Sixaola drainage, Costa Rica. Above, the holotype (LACM
36018- 1), a mature male, 14.6 mm SL. Below, a young female
paratype (UCR 897-2), 20.1 mm SL.

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 301:1-8.

Bussing: Costa Rican Fish

Figure 4. Gonopodial suspensorium of Phallichthys quadripunc-
tatus, new species (UCR 897-2), a male paratype, 14.9 mm SL.

angular dextral flanges on distal halves; gonactinost 10 short,
directed dextrally, and incorporated into gonactinost 9.

SENSORY CANALS: Sensory canals of head poorly devel-
oped. Supraorbital canal consisting of an open groove between
pores 3 and 4 (Gosline 1949) and a closed tube uniting pores 6
and 7; an indistinct groove between pores 2 and 3 and between
pores 5 and 6 visible on some specimens. Preopercular canal with
6 pores connected by closed tube. Preorbital canal with two pores
connected by closed tube; a shallow groove below lower pore on
some specimens.

COLORATION: In live material a tenuous light blue irides-
cence sometimes apparent on posterior margin of dorsal fin of
both sexes, and anal fin of females. Other color traits similar in
live and preserved examples. Generally four conspicuous subcir-
cular spots along horizontal axis of body (Fig. 2). Anterior spot
one scale row above fourth or fifth scales of lateral series;
slightly closer to dorsal-fin origin than posterior margin of eye.
Second spot one scale row above eleventh or twelfth scale of
lateral series, and below fourth dorsal-fin ray. Third spot cen-
tered on nineteenth or twentieth scale of lateral series; slightly
anterior to posterior margin of dorsal fin. Fourth spot over
twenty-fifth or twenty-sixth scale of lateral series; usually cen-
tered between dorsal and ventral margins of caudal peduncle or
slightly displaced ventrally. Individual variants (21 percent of
specimens collected) with three or five spots on one or both sides
of body. Three-spotted variants with first spot (4 cases) or second
spot (1 case) missing. Five-spotted variants with extra spot
between spots 2 and 3 (2 cases) or between spots 3 and 4 (8
cases); in these specimens adjacent spots slightly displaced to ac-
commodate extra spot. Holotype normal on left side (Fig. 2),
with five spots on right side. Four-spot pattern present as verti-

cally elongated blotches on newborn specimens.

Scale pockets edged in black, contrasting sharply with pale
yellow-gray ground color; belly and ventral half of head white.
Dorsal fin dark at base and posterior margin, clear between; large
melanophores on interradial membranes of proximal third of fin
and along distal one-fifth or one-sixth of fin. Anal fin with black
interradial membranes; a conspicuous spot at posterior base of
anal fin produced by melanophores on base of last two rays and
adjoining urosome. Gonopodium of male with small melano-
phores along rays 3 and 5; an intense black spot at base, and
along most of length of last three anal rays and extending onto
urosome. Pectoral, pelvic and caudal fins clear.

ETYMOLOGY: The specific name, quadripunctatus , refers to
the four distinctive spots on the body of this species.

ECOLOGY: The species was collected only in two shallow
rivulets and a roadside ditch. Collection UCR-897 was made in
a clear, shaded brook (0.5 m wide by 0.1 m deep, temperature
26°C, sand and gravel substrata), which also formed a stagnant,
turbid ditch (0. 1 m deep, temperatures 26° to 40°C, sand sub-
strata). The fish were collected in the strongly insolated ditch in
areas where bottom temperatures reached as high as 34°C. Only,
one other fish species was taken at this site, a species of Rivulus
from the running-water habitat.

Collection UCR- 1017 was made in shaded and unshaded por-
tions of a small stream (2.0 m wide by 0.2 to 0.5 m deep, tem-
perature 20°C, sand and rock substrate). At this station the new
species was not common, or perhaps just difficult to collect with
dip nets. Other species taken at this site were Phallichthys
amates, Alfaro cultratus and a species of Rivulus.

In an aquarium P. quadripunctatus is shy unless kept with
small, less timid species in a heavily planted tank. It readily
accepts small entomostracans (Ceriodaphnia spp.) and prepared
dry foods.

GEOGRAPHIC DISTRIBUTION: P. quadripunctatus is only
known from two tributaries of the Rio Sixaola in extreme eastern
Costa Rica (Fig. 1). It should also occur to the south in Pana-
manian tributaries of the Sixaola. The species has not been taken
at apparently suitable habitats in adjoining drainages to the north.
In their respective ranges, this form and Phallichthys tico are
sympatric, and often syntopic, with Phallichthys amates.

REMARKS

Phallichthys quadripunctatus presents the diagnostic character-
istics of the genus (Rosen and Bailey 1959; Bussing 1963). In
general physiognomy, size, and characteristics of the gonopodial
system, P . quadripunctatus and P. tico share a number of simi-
larities which distinguish them from their congeners: maximum
size (largest quadripunctatus , 32.7 mm SL and P. tico, 32.9 mm
SL vs. P. fairweatheri, at least 44 mm SL and P. amates, 67 mm
SL); terminal segment of gonopodial ray 4a several times longer
than wide (omitted from original description of P. tico ) and with
a membranous pad at the tip; right and left halves of gonopodial
ray 4 symmetrical; and a widely expanded primary gonactinost
complex.

Dr. Donn E. Rosen (in lilt.) has pointed out the similarity of
the fleshy pad at the tip of the gonopodium of P. fairweatheri
and P . quadripunctatus . A similar membranous swelling is pres-
ent in P. tico. Dr. Rosen suggests that if the pad is a derived
character for the group, then P . quadripunctatus and P . tico may
be the sister-group of P. fairweatheri, and these three the sister-

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 301:1-8.

Bussing: Costa Rican Fish

group of P. amates. The fact that P. fairweatheri , unlike its con-
geners, has dextral gonopodial asymmetry, may not be a strong
objection to this hypothesis. Rosen and Bailey (1959) noted that
a simple genetic change may determine the direction of gon-
opodial asymmetry and thus, would not necessarily signify a fun-
damental divergence between P . fairweatheri and its congeners.

ACKNOWLEDGMENTS

It is a pleasure to acknowledge the assistance of Myrna and
Eric Bussing, and members of the 1976 University of Costa Rica
ichthyology course who aided in the field work, and Reeve M.
Bailey, Robert J. Lavenberg and Donn E. Rosen who criticized
the manuscript. The work was supported by the Consejo Na-
tional de Investigaciones Cientificas y Tecnologicas (CONICIT)
de Costa Rica and the Universidad de Costa Rica.

RESUMEN

Se describe un nuevo poecilido, Phallichthys quadripunctatus,
de Baja Talamanca en la vertiente atlantica de Costa Rica. La
especie nueva es de parentesco muy cercano a otra especie
diminuta, Phallichthys tico, que habita el norte de Costa Rica y
el sur de Nicaragua. La nueva especie, tanto como P. tico, se
encuentran simpatricos con Phallichthys amates dentro de sus
respectivas distribuciones geograficas. Una cuarta especie habita
el norte de Guatemala y parte de Belice.

LITERATURE CITED

Bussing, W.A. 1963. A new poeciliid fish, Phallichthys tico,
from Costa Rica. Los Angeles Co. Mus. Contr. in Sci.,
77:1- 13.

1976. Geographic distribution of the San Juan ichthy-
ofauna of Central America with remarks on its origin and
ecology, hr. Thorson, T.R. (ed.). Investigations of the
ichthyofauna of Nicaraguan lakes. Univ. Nebraska Press,
pp. 157-175.

- AND M.I. LOpez S. 1977. Distribution y aspectos

ecologicos de los peces de las cuencas hidrograficas de
Arenal, Bebedero y Tempisque, Costa Rica. Rev. Biol.
Trop., 25(1). In press.

Gosline, W.A. 1949. The sensory canals of the head in some
cyprinodont fishes, with particular references to the genus
Fundulus. Occ. Papers Mus. Zool. Univ. Mich., 519:1-17.

Martin, M. 1972. A biogeographic analysis of the freshwater
fishes of Honduras. Doctoral dissertation. University of
Southern California. 598 pp.

Rosen, E.E., and R.M. Bailey. 1959. Middle- American poe-
ciliid fishes of the genera Carlhubbsia and Phallichthys,
with descriptions of two new species. Zoologica, 44:1-44.

. 1963. The poeciliid fishes (Cyprinodontiformes), their

structure, zoogeography, and systematics. Bull. Amer. Mus.
Nat. Hist., 126:1-176.

Accepted for publication April 21, 1977.

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 301:1-8.

L52X

Number 302
March 16, 197'9

DNTRIBUTIONS IN

NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY

f; ‘

A REVIEW OF THE SERRANID FISH GENUS ANTHIAS OF THE
HAWAIIAN ISLANDS, WITH DESCRIPTIONS OF TWO NEW

i;,!, nil, | I' j

Bv John E. Randall

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. , . ■ - ■

Number 302
March 16, 1979

CONTRIBUTIONS IN SCIENCE

NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY

A REVIEW OF THE SERRANID FISH GENUS ANTHIAS OF THE
HAWAIIAN ISLANDS, WITH DESCRIPTIONS OF TWO NEW

SPECIES

By John E. Randall

Published by the NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY • 900 EXPOSITION BOULEVARD • LOS ANGELES, CALIFORNIA 90007

CONTENTS

Abstract 3

Key to the Hawaiian Species of Anthias 3

Anthias (Mirolabrichthys) bicolor new species 4

Anthias (Pseudanthias) thompsoni Fowler 1923 7

Anthias ( Pseudanthias ) ventralis hawaiiensis new species and subspecies 9

Anthias (Pseudanthias) ventralis ventralis new subspecies 12

Literature Cited 13

ISSN: 0459-8113

Suggested Citation: Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 302:1-13.

Contributions in Science are articles in the earth and life sciences, presenting results of original research in Natural
History. Science Bulletin (Numbers 1-30; 28 June 1965 to 10 April 1978) and Contributions in Science (Numbers 1-300;
23 January 1957 to 16 June 1978) were merged into a single imperial octavo serial beginning with Number 301.

Contrib. Sci. Natur. Hist. Mus. Los Angeles Count}’. 1979. 302:1-13.

A REVIEW OF THE SERRANID FISH GENUS ANTHIAS
OF THE HAWAIIAN ISLANDS,

WITH DESCRIPTIONS OF TWO NEW SPECIES1

By John E. Randall2

Abstract: The serranid fish genus Anthias is represented in the Hawaiian Islands by three species: thomp-
soni (Fowler), which is distinctive in lacking prolonged dorsal spines, usually having 16 dorsal soft rays and 7
anal soft rays, and 50 to 58 lateral-line scales; bicolor, new species, with the second and third dorsal spines of
adults prolonged, usually with 17 dorsal soft rays, 7 anal soft rays, 57 to 64 lateral-line scales, and males with
a thickened and pointed upper lip; and ventralis, new species, with no prolonged dorsal spines, usually with 17
dorsal soft rays, 9 anal soft rays, and 39 to 46 lateral-line scales. A. ventralis is divided into two subspecies,
the Hawaiian A. v. hawaiiensis, and A. v. ventralis from other localities in Oceania.

In their Handbook of Hawaiian Fishes Gosline and Brock
(1960) listed ten species of serranid fishes, of which six belong in
the subfamily Anthiinae. Recent collections in the islands have
resulted in six additions to the subfamily, and adjustment is
needed of some of the anthiine names in the Handbook. In a
revision of Plectranthias Bleeker, Randall (in press) will describe
two new species of this genus from Hawaii, one of which was
reported as Pteranthias longimanus Weber by Gosline and
Brock. In addition he will record P. winniensis (Tyler) from the
islands. Pseudanthias kelloggi (Jordan and Evermann 1905),
which is only subspecifically distinct from azumanus (Jordan and
Richardson 1910) from Japan, also belongs in the genus
Plectranthias. A collection of ten postlarval specimens of Luzon-
ichthys Herre (BPBM 19926, 26-30.6 mm SL) taken by mid-
water trawl off Oahu in 1971 by Thomas A. Clarke appears to
represent an undescribed species. Description of this form is de-
layed in the hope of obtaining adults of the species. Katayama
(1975) described Caprodon unicolor from Midway. The other
species of this genus in Hawaii, which has been identified as C.
schlegelii (Gunther) (type locality, Japan), appears to be unde-
scribed. The species described as Caesioperca thompsoni by
Fowler (1923) should be classified in the genus Anthias, sub-
genus Pseudanthias . Two new species of Anthias, one in sub-
genus Pseudanthias and one in subgenus Mirolabrichthys, are
described herein. A paper on Anthias pleurotaenia (Bleeker), the
type species of Pseudanthias Bleeker 1873, and the distinction of
Anthias and Pseudanthias is in progress by P C. Heemstra and
Randall. The subgenus Mirolabrichthys Herre 1927 has been re-
vised by Randall and Lubbock (in press). Although both new
species of Anthias were first collected in the Hawaiian Islands,
they were later taken at other Indo-Pacific localities.

The holotypes and some paratypes of the new species and sub-
species are deposited in the Bernice P. Bishop Museum, Hono-
lulu (BPBM). Other paratypes have been sent to the Australian
Museum, Sydney (AM); British Museum (Natural History), Lon-
don [BM(NH)J; California Academy of Sciences, San Francisco
(CAS); Museum National d'Histoire Naturelle, Paris (MNHN);

Natural History Museum of Los Angeles County (LACM); and
National Museum of Natural History, Washington, D.C.
(USNM).

Data in parentheses in the descriptions below apply to the para-
types if different from the holotype. Tables 1 to 3 present the
meristic data of the three species of Anthias. Tables 4 to 6 consist
of the proportional measurements of the type specimens of the
new taxa. More measurements are given in these three tables than
are summarized in the text.

KEY TO THE HAWAIIAN SPECIES OF
ANTHIAS

la. Second and third dorsal spines prolonged in adults (espe-
cially in males); front of upper lip of males thickened and
moderately pointed; lateral-line scales 57 to 64; mandible

naked; subopercular margin smooth in adults

bicolor new species

lb. No dorsal spines prolonged; front of upper lip not thickened;

lateral-line scales 39 to 58; mandible scaled; lower suboper-
cular margin serrate 2

2a. Anal soft rays 7; dorsal soft rays usually 16; pectoral rays 20
to 22; lateral-line scales 50 to 58; margin of posterior half of

orbit rimmed with fleshy papillae thompsoni

2b. Anal soft rays 9; dorsal soft rays usually 17; pectoral rays 15
(rarely 16); lateral-line scales 39 to 46; margin of posterior
half of orbit smooth ventralis new species

‘Review Committee for this Contribution:

Phillip C. Heemstra
Robert J. Lavenberg
Roger Lubbock

2Bernice P. Bishop Museum, P. O. Box 6037, Honolulu, Hawaii 96818;
and Research Associate in Ichthyology, Natural History Museum of Los
Angeles County.

Contrib. Sci. Natur. Hist. Mus. Los Angeles County . 1979. 302:1-13.

Randall: Hawaiian Anthias

Anthias (Mirolabrichthys) bicolor

NEW SPECIES
Figures 1-3; Tables 1-4

HOLOTYPE: BPBM 10146, 99.2 mm SL, male, Hawaiian Islands,
Oahu, Waianae coast off Pokai Bay, small area of live coral with holes
and ledges, 33.5 m, quinaldine, J. E. Randall, P. M. Allen and E. S.
Helfman, 7 September 1970.

PARATYPES: BPBM 10141, 1(99.9 mm SL), Hawaiian Islands,
Oahu, off Pokai Bay, patch of coral rubble with some live coral, 27.5 m,
quinaldine, J. E. Randall, 31 August 1970; BPBM 10142, 4(76.5-99.2
mm SL), same data as preceding; CAS 35678, 1(94.5 mm SL), same
data as holotype; USNM 215284, 1(63.8 mm SL), same data as holotype;
BPBM 10145, 1(34.3 mm SL), Hawaiian Islands, Oahu, off Lahilahi
Point, cave in 27.5 m, quinaldine, J. E. Randall, 7 September 1970;
BPBM 19924, 2(26-26.9 mm SL), Hawaiian Islands off Oahu (21°
20-30' N; 158° 20-30' W), 200 m, 3.05-m Izaacs-Kidd midwater trawl,
R/V Ter it u, T. A. Clarke, 16 September 1970; BPBM 10173,
4(82.2-111 mm SL), Hawaiian Islands, Oahu, off Kahana Bay, W side
at edge of channel, cave in 26 m, rotenone, J. E. Randall, 27 September
1970; ANSP 134235, 1(83.4 mm SL), same data as preceding; LACM
35579-1, 1(81 mm SL), same data as preceding; MNHN 1974-48, 1(76
mm SL), same data as preceding; BPBM 13344, 3(27.5-57.5 mm SL),
Hawaiian Islands, Oahu, off Makapuu Point, collected from salvaged
vessel raised from 68 m, W. D. Madden, 14 October 1970; BPBM
15498, 1(29 mm SL), Hawaiian Islands, Oahu, off Makaha, ledge in
44.5 m, rotenone, J. E. Randall, ,L. Taylor, P. M. Allen and A. Na-
hacky, 16 June 1973; BPBM 15877, 4(45.6-63 mm SL), New Guinea,
Madang, Rausch Pass, S side, isolated coral block on sloping sand bot-
tom, 34 m, quinaldine, J. E. Randall, 18 August 1973; BPBM 19930,
1(82 mm SL), Mauritius, W coast, Medine, off La Fouche, 32 m, quin-
aldine, J. E. Randall, 19 November 1973; BPBM 17959, 1(66.1 mm
SL), Marshall Islands, Enewetak Atoll, Lagoon off Japtan (David) Island,
small low-profile patch reef on sand in 18 m, rotenone, J. E. Randall, P.
Lamberson and R. McNair, 13 December 1974; BPBM 18833, 1(72 mm
SL), Maidive Islands, Villingili Island, lagoon reef, cave in 35 m quinal-
dine, J. E. Randall, 18 March 1975; MNHN 1975-1139, 1(74.5 mm
SL), Hawaiian Islands, Oahu, off Makaha, ledge in 36.5 m, rotenone, J.
E. Randall, P. S. Lobel, J. Culp and J. L. Earle, 3 May 1975; BPBM
19916, 1(72.3 mm SL), Loyalty Islands, Mare Island, 55 m, P. Laboute
and Y. Magnier, 20 November 1975; AM 1.18721-001, 1(65 mm SL),
same data as preceding; BM(NH) 1976.3.15.1, 1(66.5 mm SL), same
data as preceding.

DIAGNOSIS: Dorsal rays X,16 to 18 (usually 17); anal rays

111. 7 or 8 (rarely 8); pectoral rays 19 to 21 (rarely 21); lateral-line
scales 57 to 64; gill rakers 11 or 12 + 26 to 28; depth of body
2.74 to 3.04 in SL; third dorsal spine prolonged in adult females,
and second and third spines prolonged in adult males; front of
upper lip of males thickened and moderately pointed; no papillae
along hind margin of orbit; margins of subopercle and interoper-
cle smooth in adults; pelvic fins long, reaching at least to anus
(generally to or beyond origin of anal fin); teeth on vomer large;
patch of teeth on palatines broad (about 10 irregular rows of teeth
at broadest place); no auxiliary scales on body; mandible naked;
dorsal and anal fins naked; upper half of body yellow-orange in
life, lower half lavender-pink; males with fleshy tips of elongate
second and third dorsal spines yellow.

DESCRIPTION: Dorsal fin X,17 (17 or 18, usually 17; two of
27 paratypes with XI, one of these with 16 soft rays); anal fin

131.8 (26 paratypes with 7, one with 8); pectoral rays 20 (19 or
20; one of 27 paratypes with 21 rays on one side), upper two
unbranched; pelvic fin 1,5; principal caudal rays 15 (upper and
lower unbranched); lateral-line scales 63 (60 on other side of
holotype) (57-64); scales above lateral line to origin of dorsal fin
9 (8-9); scales below lateral line to origin of anal fin 23
(21-23); circumpeduncular scales 33 (31-33); gill rakers 12 + 27
(11-12 + 26-28); pseudobranchial filaments 20 (16-20);
branchiostegal rays 7; vertebrae 26.

FIGURE 1. Anthias ( Mirolabrichthys ) bicolor, juvenile, 29 mm SL,
Oahu, Hawaii, BPBM 15498.

Body somewhat elongate, the depth 2.82 (2.74-2.96) in SL (a
little more elongate on non-Hawaiian specimens, the depth
2.84-3.04 in SL); body moderately compressed, the width 2.08
(1.97-2.36) in depth; head length 3.21 (2.81-3.23) in SL; snout
3.54 (3.36-4.27) in head, the front of upper lip of males thick-
ened, somewhat pointed, and freely movable dorso-ventrally; di-
ameter of orbit 4.34 (3.08-4.5) in head; posterior edge of orbit
without fleshy papillae; interorbital space convex, the bony
width 4.17 (3.86-4.4) in head; least depth of caudal peduncle
2.15 (2.15-2.84) in head.

Mouth oblique and moderately large, the maxilla reaching to
or posterior to a vertical at rear edge of pupil; mouth terminal
except on large males where it is slightly inferior due to hyper-
trophy of upper lip. Posterior end of maxilla rounded, its greatest
depth 1.2 in orbit of holotype; no supplemental maxillary bone.
Upper jaw with a band of villiform teeth which is broader an-
teriorly, the teeth at front enlarged; two large canine teeth an-
teriorly on each side of upper jaw (one laterally at front of jaw
which is perpendicular in jaw and not curved, and a larger slight-
ly curved one which lies nearly flat at posterior edge of the ante-
rior band of villiform teeth); an outer row of enlarged teeth ( 13 to
15 on holotype) on side of jaw, the more posterior ones inclined
forward and outward; lower jaw with a patch of villiform teeth
anteriorly, the front ones enlarged; two large curved canine teeth
on each side of lower jaw (one laterally at front of jaw which
points outward and the other about one-third back in jaw which
curves slightly posterior); a row of moderate- sized canines (14 to
17 on holotype) along jaw posterior to second large canine.
Vomer with unusually large teeth (11 on holotype); palatines with
a patch of villiform teeth in approximately 10 irregular rows at
broadest place. Tongue pointed; upper surface with scattered very
small papillae. Gill membranes free from isthmus. Gill rakers
slender and long (largest 1.17 in orbit of holotype), notably
longer than gill filaments (longest gill filament of first gill arch of
holotype contained 1.5 in longest raker).

Opercle with three flattened spines, the central one the largest
and most posterior, the upper one obtuse and indistinct, slightly
anterior to lower; two lower spines acute; distance between tips
of two lower spines about two-thirds the distance between central
and upper spines; margins of subopercle and interopercle smooth

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 302:1-13.

Randall: Hawaiian Anthias

(except some juveniles that may have a few small serrae on these
bones); lower margin of preopercle smooth; rounded comer and
upper edge serrate (25 on holotype); serrae increase in number
with size (12 in a 34-mm specimen, 15 in a 47-mm specimen, 22
in a 57.5-mm specimen, 23 to 30 in four specimens from 81 to
88 mm, and 36 in a 104-mm specimen). The 27.5 and 29-mm
paratypes have 9 serrae on the preopercle, the lower two at the
corner enlarged, especially the lowermost, which extends slightly
beyond posterior margin of operculum; in addition, there are two
spines on the interopercle (the one at upper edge large), one on
the subopercle, and for the smaller specimen, two on the lower
edge of the preopercle. These two specimens are similar morpho-
logically to the two post-larvae (BPBM 19924, 26-26.9 mm SL)
and clearly are transforming from the postlarval form. The post-
larvae have the same spination on the opercular bones, although
the two enlarged spines at the corner of the preopercle exhibit a
distinct serration.

Anterior nostril in a short membranous tube (higher dorso-
posteriorly) directly anterior to middle of eye about half the dis-
tance from edge of orbit to edge of groove separating upper lip
from snout; posterior nostril large, situated diagonally dorso-
posteriorly to anterior one, with little or no rim; greatest diameter
of opening about equal to distance between nostrils, 6.5 in orbit
of holotype.

Scales ctenoid; no auxiliary scales on body; head scaled except
mandible, throat and gill membranes, extreme front of snout and
a broad zone on side of snout anterior to center of eye, which
includes nostrils in its upper part. Small juveniles clearly show
five diagonal rows of scales on cheek between orbit and comer of
preopercle (plus a single scale just above upper posterior comer
of maxilla); in larger individuals small scales, which become
progressively larger, develop between the orbit and the larger
scales and near the margin of the preopercle; spinous portions of
dorsal and anal fins naked, the soft portions scaled basally (scales
extending farther out on interradial membranes than on rays);
caudal fin with small scales covering more than three-fourths dis-
tance to posterior margin; pectoral fins with small scales on basal
third or more of fin; pelvic fins with small scales on medial sur-
face.

Lateral line a smooth curve following contour of back; last
pored scale slightly anterior to end of hypural plate. Some pores
of cephalic lateralis system obscured by scales; prominent are: a
pore in front of anterior nostril, one between nostrils, one medial
to nostrils, two close together in interorbital space above front of
eye, a series near edge of orbit around posterior half of eye, six
on suborbital and preorbital region anterior to a vertical through
center of eye, and six in a mandibular series beginning at end of
lower margin of preopercle (last two on chin close together).

Origin of dorsal fin nearer a vertical at upper end of preopercu-
lar margin than upper end of gill opening; third dorsal spine pro-
longed in adult females, the length 4.85 to 5.82 in SL, more
prolonged in males, 2.42 (2.65-3.21) in SL; second dorsal spine
of females 7.51 to 8.28 in SL; second dorsal spine of males pro-
longed, 3.39 (2.63-2.93) in SL; tips of prolonged second and
third dorsal spines of males with a short fleshy cirrus; longest
dorsal soft ray 1.77 (1.79-2.02) in head; third anal spine 2.38
(2.12-2.37) in head; longest anal soft ray 1.23 (1.15- 1744) in
head; caudal fin lunate, the lobes filamentous, caudal concavity
3.22 (3.47-3.8) in SL; longest rays of pectoral fins (tenth or
eleventh) 3.77 (2.9-3.61) in SL; pelvic fins long, 2.49
(2.42-3.09) in SL, reaching to or beyond anus, and well beyond
spinous portion of anal fin in males and small juveniles.

Color of adults and large juveniles in alcohol pale with no dark
markings. Color of male holotype when fresh: upper half of body
salmon, lower half lavender-pink; an orange-yellow band faintly
edged in lavender running from lower part of eye to mid-pectoral
base, lavender line continuing anterior to lower part of eye as a
demarcation between orange-yellow color of most of snout and pale
pinkish ventral part of head; indistinct pale red streak from beneath
lower part of pectoral fin to above anus. Narrow red area basally at
front of dorsal fin; dorsal fin mainly yellow with indistinct blotches
of pale blue in middle, margin lavender, outer part of second and
third and nearly all of fourth interspinous membranes red; fleshy
cirri at tips of second and third spines bright yellow. Anal fin with
pale blue membranes, shading outwardly to light red, rays pink, and
margin pale bluish . Caudal fin with a broad zone of light red with a
slight wash of yellow in lobes, shading on central and posterior part
of fin to pale yellow with pink rays; pectoral fins pale pink. Pelvic
fins colored much like anal fin, the lateral edge with pale bluish
margin.

Color from field notes of a 99.9-mm female paratype (BPBM
101 04 ) when fresh: upper half of body orange- yellow, lower half
lavender pink (more lavender on abdomen and thorax); scales in
zone of integradation of color on mid-side with yellow centers and
pink edges; some irregular streaks of scales with yellow centers
extending a short distance into pink lower half of body; head above
lower edge of eye orange-yellow, below lavender with a wash of
orange, especially on maxilla and rest of head below and anterior
to eye; horizontal lavender line between orange-yellow and laven-
der colors on snout; tip of lower jaw lavender; orange band nearly as
broad as pupil and narrowly edged with lavender, running from
lower posterior portion of orbit to pectoral base. Dorsal fin primar-
ily pale red, irregularly yellow at base (yellow limited to first five
interspinous membranes); margin lavender, and an indistinct sub-
marginal zone of yellow; rays of fin rose red. Anal fin light
lavender-pink with some yellow areas, rays slightly darker, margin

TABLE 1

Fin-ray counts of Anthias

species

Dorsal Rays
XI 16 17

Anal Soft Rays
7 8 9 10

Pectoral Rays
16 17 18 19 20

21 22

bicolor

10 17

thompsoni

25 1

ventralis

hawaiiensis

19 1

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 302:1-13.

6 Randall: Hawaiian Anthias

Figures 2-4. Anthias from Oahu, Hawaii. 2, Upper, Anthias (Mirolabrichthys) bicolor, holotype, male, 99.2 mm SL, BPBM 10146. 3,
Middle, Anthias ( Mirolabrichthys ) bicolor, female, 83.5 mm SL, BPBM 10173. 4, Lower, Anthias (Pseudanthias) thompsoni, female,
126 mm SL, BPBM 11290.

lavender. Caudal lobes rose red; upper and lower edges narrowly
lavender, with traces of yellow submarginally; centro-posterior part
of fin light red-orange. Pectoral fins pale magenta, the membranes
pale. Pelvic fins pale yellowish, the rays light lavender.

Color from an ektrachrome transparency of 29-mm paratype
(BPBM 15498; Fig. 1): upper half of body yellow, lower half
lavender; dorsal fin pale yellow with violet margin. Pectoral fins
lavender; remaining fins pale with a lavender cast, upper and lower
margins of caudal fin, filamentous tip of pelvic fins, and tips of
longest anal rays violet. In preservative, this fish shows a faint
concentration of black pigment on the third interspinous membrane,
a smaller spot of the same density on the second membrane, and a
large but more diffuse one at the rear base of the spinous portion of
the fin, extending in about equal area onto the body. In the 27.5-mm
paratype these same dark areas are more evident. They are even
more conspicuous on the two postlarvae (26-26.9 mm SL); in
addition, the postlarvae have a blackish spot on the pelvic fins
between the spine and the third ray about one- third distance from the
base (only a faint remnant of this spot may be seen on the 27.5 and
29-mm specimens) and three less distinct blackish spots on caudal
peduncle (dorsally, ventrally and at mid-base of caudal fin).

REMARKS: Adult and juvenile specimens of A. bicolor have
been collected in the Hawaiian Islands, Marshall Islands, Loyalty
Islands (kindly supplied by Pierre Fourmanoir of ORSTOM, New
Caledonia), New Guinea, Maidive Islands and Mauritius in a
depth range of 18 to 68 m. However, I have observed the species
in 8 m in Hawaii, and aquarium fish collector Anthony Nahacky
has seen it in as little as 5 m off Makua, Oahu. This fish usually
is found in the vicinity of caves or ledges, but not infrequently it
is encountered on small patch reefs where shelter seems minimal
(even here an unexpected crevice generally will account for its
presence). Typically it occurs in aggregations, though these may
consist of only a few individuals. It feeds on zooplankton, and
may rise as much as 3 or 4 m above the bottom in feeding.
Although it may intermingle with feeding aggregations of A.
thompsoni, it tends to maintain itself apart from this abundant
Hawaiian species.

As in many other species of the subfamily Anthiinae, males
are consistently larger than the females, probably as a result of
sex reversal; also, they are notably less abundant than females.
Although the differences in color between the sexes are not as
marked as in some anthiines, the greatly prolonged second and

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 302:1-13.

Randall: Hawaiian Anthias 7

Figures 5-7. Species of the subgenus Anthias ( Pseudanthias ). 5, Upper, Anthias ventralis hawaiiensis, holotype, male, 76.9 mm SL,
Oahu, Hawaii; BPBM 10560. 6, Middle, Anthias ventralis hawaiiensis, female, 51.8 mm SL, Oahu, Hawaii; BPBM 10159. 7, Lower,
Anthias ventralis ventralis, male, 51.5 mm SL, Pitcairn Island; BPBM 16883.

third dorsal spines with a yellow cirrus at each tip clearly differ-
entiates the male.

Two postlarvae (BPBM 19924) were collected in 200 m with a
midwater trawl off leeward Oahu by Thomas A. Clarke of the
University of Hawaii who kindly made them available as para-
types. The tow was made at 2006-2222 hours. The net remained
open during descent and ascent, thus one cannot be certain of the
depth of capture; however, the lowering and raising of the net
required only 16 minutes of the 136-minute towing time.

The species appears to attain a larger size in the Hawaiian Is-
lands than elsewhere. Four males in Hawaii range from 99.2 to
1 1 1 mm SL. Six males from outside Hawaii measure 63 to 82
mm SL. This difference may be due, at least in part, to the cooler
water of the Hawaiian region. The largest non-Hawaiian speci-
men, however, is the 82-mm one from Mauritius, also a cool
subtropical locality.

Named bicolor in reference to the salmon pink of the upper
half and lavender pink of the lower half of the body. When
viewed underwater in its usual moderately deep habitat (hence
with shades of red subdued), the back appears more yellow and
the lower side more blue, thus enhancing the bicolored effect and

making this species easily distinguished from A. thompsoni. An-
ticipating the scientific name, some fish fanciers in Hawaii al-
ready have begun to use the common name “Bicolored Bass” for
this species.

Anthias (Pseudanthias) thompsoni Fowler 1923

Figure 4; Tables 1-3

Caesioperca thompsoni Fowler 1923. B. P. Bishop Mus., Occ. Pap.

8(7):379 (type locality, Honolulu).

DIAGNOSIS; Dorsal rays X,16 (one of 33 specimens with
17); anal rays III, 7; pectoral rays 20 or 21 (usually 21, one of 33
with 22); lateral-line scales 50 to 58; gill rakers 10 to 12 + 25 to
28; depth of body 2.7 to 3.3 in SL; no dorsal spines prolonged
(fifth spine usually the longest, but fourth to tenth spines nearly
equal); front of upper lip of males not thickened and not pointed;
posterior edge of orbit with a row of fleshy papillae; margins of
subopercle and interopercle smooth; pelvic fins short, usually not
reaching anus (just reaching anus in large males); caudal fin lu-

Contrib. Sci. Natur. Hist. Mus. Los Angeles County . 1979. 302:1-13.

Randall: Hawaiian Anthias

TABLE 2

Lateral-line scale counts of Anthias

species 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 61 62 63 64

bicolor 2 5 4 6 6 2 2 1

thompsoni 1 1 2687422

ventralis

ventralis 1 1 1 2 2 3 1

hawaiiensis 2 2 6 6 1 2 1

nate, the caudal concavity of large males up to 1.85 in SL; small
villiform teeth on vomer forming a broad V-shaped (nearly tri-
angular) patch; band of teeth on palatine narrow, consisting of
only two irregular rows; anxiliary scales present over entire body;
mandible scaled; dorsal and anal fins naked.

Color in alcohol pale with no dark markings. Color of an adult
female when fresh: body red-orange dorsally, with a yellow spot
basally on each scale; laterally the yellow has shifted to the edges
of the scales, and the red-orange color is more magenta; ventrally
the yellow fades to light lavender, and the scale centers to pale
heliotrope. A double band, yellow above and magenta below,
from lower edge of eye to lower opercle, then turning upward
along margin of opercle to level of upper base of pectoral fin; a
similar but faint double band extending onto snout from front of
eye; a narrow yellow rim on posterior half of eye. Dorsal fin red;
membranes mottled with yellow; margin of soft portion lavender.
Anal rays salmon; the membranes light yellow; margin of ante-
rior half of soft portion lavender (to tip of acute angle of margin).
Caudal fin yellow-orange; upper and lower edges red, shading
outwardly to lavender. Pectoral rays salmon, membranes trans-
parent; base of fin yellowish with faint lavender marking in ap-
proximate V-shape. Pelvic rays salmon, membranes light yellow.
Color of a large adult male is similar but the magenta streak on
the head is broader with a wide band of yellow on either side;
and the pectoral fin base more yellow with the lavender V more
evident; large area of yellow on the basal rayed part of fin, with a
faint broad outer margin of lavender; maxilla pale yellow, with a
broad pale magenta border; lavender upper and lower margins of
caudal fin and red submarginal lines are more pronounced, with a
faint posterior margin of lavender and red submarginal line in
central part of fin.

REMARKS: A. thompsoni is known only from the Hawaiian
Islands. It is common near rocky vertical discontinuities in the
bottom. Brock and Chamberlain (1968) commented on its abun-
dance in loose schools on near-shore escarpments from submarin-

ing observations off the Waianae coast of Oahu. Strasburg, Jones
and Iversen (1968) also observed it from a research submarine, in
the depth range of 55 to 110 m, as did Clarke (1972:311) who
noted that it was “seen frequently down to about 150 m, but
never deeper.”

Bishop Museum collections of this species have come from 26
to 157 m. It occurs in water at least as shallow as 10 m; however,
it is not common in the lesser depths.

The stomach contents of six specimens, 98 to 119 mm SL,
from three different lots, were examined. This material consisted
entirely of zooplankton, primarily copepods. Also, shrimp larvae
frequently were eaten. Adults feed up to several meters above the
bottom (exceptionally as much as 9 m).

On April 25, 1972 I speared a jack (Caranx melampygus) 410
mm fork length off Barber’s Point, Oahu that had three A.
thompsoni in its stomach measuring 75 to 85 mm SL.

Females are much more numerous than males. Males have
been observed in courtship; they elevate the dorsal fin during the
display, which appears to take on a deeper red color; also, there
is a red band along the back and a red border behind the opercu-
lum.

The holotype (BPBM 3376, 122 mm SL) is in the Bishop Mu-
seum and is in good condition. Corrections are in order for some
of the counts given by Fowler in the description. The fish has 21
(not 22) pectoral rays, 54 (not 58) lateral-line scales, and 28 (not
27) lower-limb gill rakers. Fowler described thompsoni in Caesi-
operca Castelnau, a genus otherwise known only from temperate
waters of Australia and New Zealand. A specimen of the type
species of the genus, C. rasor (Richardson), was examined. It
clearly is divergent from thompsoni, having a supplemental
maxillary bone (absent in thompsoni), 21 dorsal soft rays, 14
pectoral rays, and a shallowly forked caudal fin.

The largest specimen of A. thompsoni (BPBM 15443) mea-
sures 149 mm SL. The Natural History Museum of Los Angeles
County has specimens in two series(LACM 32242-1, 15:89-144

TABLE 3

Gill-raker counts of Anthias3

species

Upper Limb
8 9 10 11

Lower Limb
24 25

bicolor

thompsoni

ventralis

hawaiiensis

3Gill-raker counts were made on the first arch and include all rudiments; the raker at the angle is included in
the lower-limb count.

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 302:1-13.

Randall: Hawaiian Anthias

TABLE 4

Measurements of type specimens of Anthias (Mirolabrichthys) bicolor
expressed as a percentage of the standard length

HOLOTYPE PARATYPES

BPBM

10148

BPBM

15498

BPBM

13344

BPBM

10142

BPBM

10173

BPBM

10173

Standard length (mm)

99.2

29.0

47.0

67.6

88.3

104.3

Depth of body

35.4

33.8

35.5

36.5

34.5

35.7

Width of body

17.0

15.5

15.0

17.0

17.5

18.2

Head length

31.2

35.5

32.0

33.9

31.0

31.9

Snout length

8.8

8.6

7.5

8.7

8.2

9.5

Diameter of orbit

7.2

11.5

10.2

8.5

7.8

7.1

Bony interorbital width

7.5

8.3

7.7

7.8

8.1

Length of maxilla

16.4

15.6

15.1

15.8

15.9

16.9

Least depth of caudal peduncle

14.5

12.5

13.6

15.1

14.4

14.8

Length of caudal peduncle4

21.0

20.0

21.3

20.7

21.5

21.3

Predorsal length

26.9

29.9

26.2

27.2

25.6

26.5

Preanal length

62.2

62.4

63.5

65.2

62.7

64.2

Prepelvic length

33.3

36.2

33.7

35.3

33.3

35.4

Length of first dorsal spine

4.3

6.5

—

4.7

5.1

Length of second dorsal spine

29.5

13.3

12.1

13.3

12.3

38.0

Length of third dorsal spine

31.2

15.5

—

20.6

17.2

41.3

Length of tenth dorsal spine

10.5

14.5

14.3

13.9

11.4

11.9

Length of longest dorsal ray

17.6

17.5

17.6

17.9

17.0

17.8

Length of dorsal fin base

65.4

63.2

65.8

63.6

65.6

66.6

Length of first anal spine

7.0

8.9

8.5

7.1

6.9

8.0

Length of second anal spine

—

19.7

17.6

16.6

14.2

14.1

Length of third anal spine

13.1

15.2

15.1

14.6

13.1

13.7

Length of longest anal ray

25.2

31.0

25.0

23.4

24.7

26.9

Length of anal fin base

18.5

20.7

19.5

19.6

18.7

19.0

Length of pectoral fin

26.5

34.4

30.2

31.2

28.1

27.7

Length of pelvic spine

16.1

21.0

18.2

18.1

16.4

16.2

Length of pelvic fin

40.1

41.4

33.4

32.9

32.4

41.2

Length of caudal fin

44.4

52.4

51.7

44.7

40.3

47.5

Caudal concavity5

31.0

28.9

34.9

29.6

26.3

33.7

■•Measured diagonally from rear base of anal fin to mid-base of caudal fin.

5Measured horizontally between verticals at tips of shortest and longest caudal fin rays.

mm SL; LACM 32668-14, 17:83-148 mm SL), which are
nearly as large. These were taken by trawl off Haleiwa, Oahu in
120 m.

John E. Fitch of the Department of Fish and Game, State of
California, examined the otoliths from these LACM specimens.
He detected one presumed annular ring in the otoliths of four
specimens 83-95 mm, two in 14 specimens 93-112 mm, three
in three specimens 112-127 mm, four in four specimens 1 37—
140 mm, and five in a 145-mm specimen. He wrote me,
“. . . While they (the otoliths) are not the best in the world for
reflecting ages, I feel reasonably certain of my readings.”

Five postlarval specimens of A. thompsoni (BPBM 19925,
23.4 to 24.3 mm SL) were collected off leeward Oahu by
Thomas A. Clarke of the University of Hawaii from the R/V
Teritu in a 3.05-m midwater trawl at a depth of 225 m on 1 1 June
1971. The time of the tow was 0200- 0416 hours; the lowering
and raising of the net required 18 minutes of this time. These fish
have a very long serrate spine at the comer of the preopercle
(nearly as long as diameter of orbit), with 7 to 9 smaller spines
on upper limb of preopercle (adjacent spine somewhat enlarged
and serrate), and three spines on lower limb (middle spine some-
what enlarged and serrate); also, there is a spine on the subopercle
and 2 to 5 on the interopercle . There are no distinctive large
blackish spots as on the postlarvae of A. bicolor, but there is a

concentration of dark pigment basally on the third interspinous
membrane of the dorsal fin and lesser dark blotches may be
found on the lower part of the second membrane and basally on
the fourth to seventh spines. Also a small amount of dark pig-
ment may be seen on some specimens on the membranes behind
the tips of the second to seventh spines and near the base of the
pelvic fins centered on the first and second soft rays. As in many
postlarvae, there are some fine dots of melanin over the occipital
region of the head.

Anthias ( Pseudanthias ) ventralis hawaiiensis

NEW SPECIES AND SUBSPECIES
Figures 5-6; Tables 1-3, 5

HOLOTYPE: BPBM 10560, 76.9 mm SL, male, Hawaiian Islands,
Oahu, Moku Manu; off N side at base of drop-off, 49 m, rotenone, J. E.
Randall, W. J. Baldwin and A. Stark, 9 October 1968.

PARATYPES: BPBM 10561, 7(38.3-71.1 mm SL), same data as
holotype: ANSP 134236, 1(62.5 mm SL), same data as holotype, AM
1.18720-001, 1(49 mm SL), same data as holotype; BM(NH)
1976.3.15.2, 1(65.1 mm SL), same data as holotype; LACM 35578- 1,
1(69.4 mm SL), same data as holotype; BPBM 10159, 3(36.3-62.2
mm SL), Hawaiian Islands, Oahu, off Kahana Bay, NW side of channel
at entrance to bay, ledges and caves in 26 m, quinaldine, J. E. Randall
and P. M. Allen, 12 September 1970; MNHN 1974-17, 1(60.2 mm SL),
same data as preceding; BPBM 10172, 1(52.9 mm SL), same locality as

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 302:1-13.

Randall: Hawaiian Anthias

TABLE 5

Measurements of type specimens of Anthias (Pseudanthias) ventralis hawaiiensis
expressed as a percentage of the standard length

HOLOTYPE PARATYPES

BPBM

10560

BPBM

15058

BPBM

10159

BPBM

10159

BPBM

10159

BPBM

10561

Standard length (mm)

76.9

24.0

36.3

51.8

62.2

71.1

Depth of body

37.8

35.8

35.4

34.7

34.0

38.4

Width of body

18.6

17.4

15.8

16.4

15.6

18.4

Head length

29.7

34.5

33.6

31.4

30.5

30.0

Snout length

7.4

7.5

7.4

7.2

7.3

7.7

Diameter of orbit

7.8

11.6

11.0

9.9

8.8

8.3

Bony interorbital width

8.9

9.2

9.1

8.9

9.0

Length of maxilla

14.8

15.8

15.4

14.2

14.4

14.5

Least depth of caudal peduncle

15.0

14.1

15.1

14.9

14.5

15.1

Length of caudal peduncle

20.7

19.7

19.6

18.7

20.2

19.2

Predorsal length

32.4

32.1

33.1

32.6

32.7

32.8

Preanal length

60.2

63.3

62.0

63.1

61.7

62.5

Prepelvic length

33.2

35.3

33.1

33.7

32.1

33.7

Length of first dorsal spine

5.8

5.6

5.5

5.6

5.0

5.3

Length of longest dorsal spine

12.9

16.0

15.7

15.3

13.4

13.2

Length of longest dorsal ray

32.3

18.8

23.8

21.8

26.2

30.0

Length of dorsal fin base

62.6

54.5

61.1

59.9

60.0

60.8

Length of first anal spine

6.2

7.9

7.7

7.1

6.4

6.1

Length of second anal spine

12.9

16.3

17.2

15.2

12.9

12.7

Length of third anal spine

13.6

15.8

16.6

14.3

12.9

12.8

Length of longest anal ray

40.2

30.8

30.3

32.4

37.5

39.4

Length of anal fin base

23.4

22.9

22.3

21.6

Length of pectoral fin

30.9

33.3

32.5

31.4

30.2

32.1

Length of pelvic spine

16.0

19.1

19.8

17.2

15.8

15.5

Length of pelvic fin

49.7

37.0

28.5

37.6

43.2

—

Length of caudal fin

32.7

48.0

36.3

35.5

34.6

37.0

Caudal concavity

11.8

24.1

14.0

13.7

12.5

16.2

preceding, rotenone, J. E. Randall, 27 September 1970. CAS 35679,
1(41.8 mm SL), Hawaiian Islands, Oahu, off Makapuu Point, collected
from salvaged vessel raised from 68 m, W. D. Madden, 14 October
1970; USNM 215285, 1(43.8 mm SL), same data as preceding; BPBM
15058, 1(24 mm SL), Hawaiian Islands, Hawaii, South Point, large
boulders and adjacent sand, 28 m, rotenone, L. Taylor and P. M. Allen,
21 May 1973.

DIAGNOSIS: Dorsal rays X,16 to 18 (usually 17); anal rays
III, 9 (rarely 10); pectoral rays 15 (rarely 16); lateral-line scales
40 to 46; gill rakers 7 to 9 + 22 to 25; depth of body 2.37-2.94
in SL; no dorsal spines prolonged (fifth spine usually the longest,
but fourth to tenth nearly as long); front of upper lip of males not
thickened and pointed; no papillae along rim of posterior half of
eye; margin of subopercle serrate, and a few serrae may be pres-
ent on upper posterior edge of interopercle; pelvic fins long,
reaching well beyond origin of anal fin (entirely beyond base of
fin in adult males); a few small teeth on vomer in an approxi-
mately triangular patch; band of teeth on palatines narrow, con-
sisting of two or three irregular rows; a few auxiliary scales dor-
sally on body; mandible scaled; dorsal and anal fins scaled except
distally. Color as in figures 5 and 6, the most conspicuous feature
being the yellow of the upper half of the head with a few short
lines of violet or magenta behind and above the eye.

DESCRIPTION: Dorsal fin X,17 (X,17 except one paratype
with XI, 18, one with X,18, and one with X , 16); anal fin III, 9
(one of 19 paratypes with 10); pectoral rays 15 (one of paratypes
with 16), upper one or two and lowermost unbranched; pelvic fin

1,5; principal caudal rays 15 (upper and lower unbranched);
lateral-line scales 42 (40-46); scales above lateral line to origin
of dorsal fin 5; scales below lateral line to origin of anal fin 19
(18-20); circumpeduncular scales 22; gill rakers 8 + 25 (17-9 +
22-25); pseudobranchial filaments 11 (10-12); branchiostegal
rays 7; vertebrae 26.

Body depth variable, 2.65 (2.37-2.94) in SL; body moder-
ately compressed, the width 2.03 (2.04-2.25) in depth; head
length 3.37 (2.9-3.33) in SL; snout 4.02 (3. 9-4. 6) in head;
front of upper lip of males not thickened and not pointed (contour
of front of upper lip when viewed from above varying from
slightly convex to flat); diameter of orbit 3.81 (2.97-3.6) in
head; posterior edge of orbit without fleshy papillae; interorbital
space smoothly convex, the bony width 3.34 (3.33-3.75) in
head; least depth of caudal peduncle 1.98 (1.98-2.44) in head.

Mouth oblique and moderately large, the maxilla reaching
posterior to a vertical at center of eye and often to or slightly
beyond a vertical at posterior edge of pupil; lower jaw projecting
slightly when mouth closed; upper posterior comer of maxilla
more broadly rounded than lower; greatest depth of maxilla 2 in
orbit of holotype; no supplemental maxillary bone. Upper jaw
with inner row of villiform teeth of moderate size broadening to a
maximum of about 4 rows anteriorly, the innermost and most
medial of these teeth notably enlarged, lying nearly flat, and di-
rected inward; a canine laterally at front of upper jaw, followed
by a row of 13 teeth along side of jaw, which are about as long
as front canine but more slender; posterior ones pointing forward;

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 302:1-13.

Randall: Hawaiian Anthias

TABLE 6

Measurements of type specimens of Anthias (Pseudanthias) ventralis ventralis
expressed as a percentage of the standard length

HOLOTYPE

BPBM

16883

BPBM

13530

PARATYPES

BPBM

13530

BPBM

13530

BPBM

19895

Standard length (mm)

51.5

20.4

31.8

50.8

54.3

Depth of body

35.1

32.9

32.0

33.4

Width of body

16.5

15.7

16.7

16.2

Head length

31.5

36.8

33.4

31.8

33.0

Snout length

7.8

7.9

7.6

7.7

Diameter of orbit

10.3

12.5

11.3

9.6

Bony interorbital width

8.8

9.0

8.5

8.9

8.7

Length of maxilla

14.5

15.7

14.1

14.9

Least depth of caudal peduncle

14.4

13.7

14.6

14.3

14.2

Length of caudal peduncle

18.9

19.4

20.4

19.2

18.6

Predorsal length

33.2

34.8

33.1

33.2

32.3

Preanal length

61.0

63.7

63.1

60.3

61.7

Prepelvic length

33.2

36.6

34.0

33.0

33.2

Length of first dorsal spine

6.2

6.3

5.8

6.1

6.2

Length of longest dorsal spine

13.4

16.2

15.7

13.9

13.1

Length of longest dorsal ray

27.0

18.1

17.7

29.7

29.5

Length of dorsal fin base

60.2

53.9

56.8

61.0

58.2

Length of first anal spine

7.4

7.8

7.7

7.3

7.1

Length of second anal spine

14.4

15.7

13.8

14.0

Length of third anal spine

13.8

15.7

15.4

13.8

13.3

Length of longest anal ray

35.4

21.5

25.1

32.6

40.8

Length of anal fin base

21.4

22.5

22.1

22.0

21.8

Length of pectoral fin

32.4

30.6

32.1

33.4

32.6

Length of pelvic spine

16.7

19.7

17.9

17.1

16.4

Length of pelvic fin

39.3

29.4

29.2

44.7

54.3

Length of caudal fin

38.3

—

37.8

—

43.7

Caudal concavity

17.8

—

15.7

—

21.5

a pair of moderate canines anteriorly in lower jaw that project
slightly forward; two to three rows of smaller, incurved teeth
medial to anterior canines; four or five strongly retrorse canines
along side of anterior third of lower jaw; posterior to these are a
series of uniformly large teeth (six on one side of holotype, nine
on other) that point forward strongly, these closely associated
with an outer row of smaller more slender teeth. Teeth on vomer
small, in a patch of approximately triangular shape; a narrow
band of two or three rows of small teeth on palatines. Tongue
pointed. Gill membranes free from isthmus. Gill rakers slender
and long, about as long as longest gill filaments; length of the
longest contained 1.6 times in diameter of orbit of holotype.

Opercle with three flattened spines; the central one the largest
and most posterior; the upper spine obtuse and indistinct, slightly
anterior to lower; two lower spines acute, distance between their
tips generally about three-fourths distance between central and
upper spines. Lower edge of preopercle smooth; upper edge
finely serrate (25 serrae on holotype; number of serrae variable,
but tending to increase with size; paratypes from 24 to 42 mm SL
have 16 to 23 serrae whereas those over 50 mm have 22 to 36);
ventral edge of subopercle with 14 (4 to 14) serrae; interopercle
with 4 (0-4) serrae near upper end.

Anterior nostril in a membranous tube (higher dorsoposterior-
ly) directly anterior to middle of eye and about halfway between
edge of orbit and groove separating upper lip from rest of snout;
posterior nostril large, dorso-posterior to anterior one, roughly
semicircular, and without a rim; its greatest diameter nearly

equals its distance from anterior nostril (about 6 in diameter of
orbit of holotype).

Scales ctenoid; some auxiliary scales may be present on nape
and body dorsal to lateral line. Head scaled except throat, gill
membranes, extreme front of snout, and a broad zone of snout
from level of lower edge of eye to above nostrils. Dorsal and anal
fins with scales on about basal half (ignoring prolonged rays and
associated membranes); caudal fin with small scales extending
about three-fourths distance to posterior margin; pectoral fins
scaled on about basal third; pelvic fins scaled on medial surface,
the scales on first ray extending slightly beyond the pelvic spine.

Lateral line a little more strongly arched than upper contour of
body, reaching highest point below posterior half of spinous por-
tion of dorsal fin; last pored-scale of lateral line just anterior to
posterior edge of hypural plate. Pores of cephalic lateralis system
similar to those of A. bicolor.

Origin of dorsal fin above (or slightly anterior or posterior) to
upper end of gill opening; no dorsal spines prolonged; fifth dorsal
spine usually the longest, 2.3 (2.05-2.28) in head; a small cirrus
behind tip of each dorsal spine; twelfth dorsal soft ray (and to a
lesser extent adjacent rays) prolonged (except in small juveniles),
its length 3.1 (3.33 to 4.2) in SL; third anal spine 2.18 (2.02-
2.36) in head; fourth anal soft ray (and to a lesser extent adja-
cent rays) greatly prolonged, its length 2.49 (2.54-3.3) in SL;
caudal fin deeply emarginate, the caudal concavity 2.52 (1.43-
2.44) in head; longest ray of pectoral fins (ninth) 3.24 (3.0-
3.31) in SL; second pelvic ray (and to a lesser extent adjacent

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 302:1-13.

Randall: Hawaiian Anthias

rays) greatly prolonged, reaching beyond spinous portion of anal
fin and often posterior to base of fin, the ray length 2.01 (2.17-
3.51) in SL.

Color in alcohol pale with no dark markings. Color of holotype
and other large males when fresh (from field notes): anterior part
of body orange, dotted with magenta, shading to light yellowish
with a lavender wash on thorax and abdomen, and to violet pos-
teriorly. Upper half of head yellow with dotted lines of magenta
as follows: from snout in front of eye through upper edge of eye
and passing dorsally to nape, from behind eye along upper edge
of opercle, from in front of lower part of eye through lower edge
of eye and across middle of operculum to middle opercular spine;
lower half of head varying from light yellowish to pale lavender.
Dorsal fin orange, edged with magenta (except posteriorly); rear
basal part nearly colorless; anal fin with anterior part (including
prolonged rays) orange, posterior portion yellow, the two regions
separated by a magenta line; caudal fin yellow-orange, shading to
magenta in broad median posterior part of fin; pectoral fins pale
orangish with a large magenta-edged orange spot on base; pelvic
fins orange with a little magenta on leading edge of spine; isthmus
yellow-orange (see Fig. 5).

Color of females when fresh similar but body mostly yellow
instead of orange, the yellow unmarked dorsally, over posterior
part of caudal peduncle and most of caudal fin; ventral half of
body lavender; magenta lines on head more solid (and hence
more evident) on females than males; anal fin almost entirely
yellow, lacking the striking two-tone coloration with magenta di-
vision of the male; yellow spot at base of pectoral fin not as
conspicuous as the orange basal spot on the fin of the male (see
Fig. 6).

REMARKS: The Hawaiian population of Anthias ventralis is
sufficiently distinct to warrant subspecific recognition, hence
hawaiiensis. See “Remarks” of A. v. ventralis for discussion of
differences.

The Hawaiian subspecies was collected in the depth range of
26 to 68 m. Nearly all specimens were taken from caves or be-
neath well-formed ledges. They are more closely tied to the sub-
stratum than the other two Anthias in Hawaii and are often seen
swimming upside down near the roof of caves. They venture out
for feeding on zooplankton (mainly copepods, but also other
crustacean larvae, fish eggs, mollusk larvae, polychaete larvae,
etc.).

This species is a good aquarium fish, though not often caught
due to its deep-dwelling habits. It is hardier than the other two
Hawaiian Anthias in aquaria.

Named ventralis in reference to the striking prolongation of the
pelvic and anal fins.

The common name “Longfin Bass” is proposed for this
species.

Anthias (Pseudanthias) ventralis ventralis

NEW SUBSPECIES
Figure 7; Tables 1-3, 6

HOLOTYPE: BPBM 16883, 51.5 mm SL, male, Pitcairn Island,
patch reef on N side off Gannet Ridge, 40 to 44 m, rotenone, J. E.
Randall, D. B. Cannoy, I. R. Haywood, R. R. Costello, J. D. Bryant
and S. Christian, 6 January 1971.

PARATYPES: BPBM 13530, 8(18.2-50.8 mm SL), Tuamotu Ar-
chipelago, Gambier Group, Temoe Atoll, outside reef on N side, isolated
dead coral block on sand near fringing reef, 41.3 m, rotenone, J. E.
Randall, D. B. Cannoy, R. McNair and R. R. Costello, 16 December

1970; BM(NH) 1973.3.5.1, 1(29.4 mm SL), same data as preceding;
LACM 35586- 1, 1(21 mm SL), same data as preceding; USNM 215289,
1(29.8 mm SL), same data as preceding; BPBM 19895, 1(54.3 mm SL),
New Caledonia, false pass Uitoe (about 3 km N of Dumbea Pass), 55 m,
bottom mainly coral rubble, rotenone, P. Laboute and Y. Magnier, 5
November 1975.

DIAGNOSIS: As in Anthias ventralis hawaiiensis, except for
slightly lower gill-raker counts (see Table 3), more interopercu-
lar serrae, smaller size and color (see below).

DESCRIPTION: Dorsal rays X,17; anal rays III, 9; pectoral
rays 15 (upper one or two and lower one or two unbranched);
pelvic rays 1,5; principal caudal rays 15 (upper and lower un-
branched); lateral-line scales 44 (39-46); scales above lateral line
to origin of dorsal fin 5 (4 or 5); scales below lateral line to
origin of anal fin 19 (18-19); circumpeduncular scales 22 (20-
22); gill rakers 8 -I- 24 (7-8 + 21-24); pseudobranchial fila-
ments 10 (9-10); branchiostegal rays 7; vertebrae 26.

Body depth 2.85 (2.86-3.13) in SL; width of body 2.13
(2.0-2.09) in depth; head length 3.17 (2.72-3.14) in SL; snout
4.05 (4.14-4.67) in head; front of upper lip of males not thick-
ened and not pointed; diameter of orbit 3.08 (2.94-3.43) in
head; interorbital space convex, the bony width 3.58 (3.58-3.93)
in head; least depth of caudal peduncle 2.19 (2.22-2.67) in
head.

Dentition similar to A. v. hawaiiensis: 14 large teeth along one
side of upper jaw of holotype, and 15 on the other; more small
teeth posterior to large ones on side of lower jaw than in
hawaiiensis. Tongue, gill rakers, nostrils and head pores compa-
rable to hawaiiensis .

Opercular spines and serrae of preopercle and subopercle as in
A. v. hawaiiensis: holotype with 24 serrae on preopercle and 9 on
subopercle. There are, however, usually more serrae on the aver-
age on the interopercle of A. v. ventralis (4 on holotype and 1 to
6 on paratypes).

Squamation similar to A. v. hawaiiensis except there are very
few auxiliary scales on dorsal body scales, and small scales do
not extend as far out on the median fins.

Origin of dorsal fin above upper end of gill opening; no dorsal
spines prolonged; fifth to ninth dorsal spines the longest, 2.37
(2.13-2.52) in head; a small cirrus behind tip of each dorsal
spine; twelfth dorsal soft ray, and to a lesser extent the eleventh
and thirteenth, prolonged in adults, its length 3.7 (3.37-5.65) in
SL; third anal spine 2.28 (2.17-2.47) in head; fourth anal soft
ray, and to a lesser extent third and fifth rays, prolonged, 2.83
(2.45-4.65) in SL; caudal fin deeply emarginate, the caudal con-
cavity 1.77 (1.53-2.12) in head; longest ray of pectoral fins
(ninth) 3.09 (3.0-3.27) in SL; second pelvic ray, and to a lesser
extent adjacent rays, greatly prolonged, reaching beyond spinous
portion of anal fin and in some larger individuals posterior to rear
base of anal fin, the ray length 2.54 (2.24-3.42) in SL.

Color in alcohol uniform pale with no dark markings. Color of
holotype and other males when fresh: orange-yellow with small
spots of lavender dorsally, shading to lavender on sides (zone of
intergradation, especially anteriorly, with yellow streaks extend-
ing ventrally into lavender zone and vice versa), and to pale yel-
low ventrally. Snout, tip of chin, and postorbital part of head
above level of center of eye bright yellow; lower part of head and
all of thorax pale yellow; deep yellow streak mid-ventrally on
head; broken line of lavender running from upper lip to eye; hori-
zontal lavender band behind eye separating upper bright yellow
and lower pale yellow of head; an arc of lavender beginning on

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 302:1-13.

Randall: Hawaiian Anthias

opercle anterior to gill opening, rimming upper gill opening and
joining posterior end of horizontal lavender line of postorbital
head; broken lavender line on nape from upper edge of orbit to
origin of dorsal fin. Dorsal fin yellow with a few small lavender
spots basally; spines and adjacent membrane tips magenta; an-
terior margin of soft portion lavender; faint diagonal light blue
line in fin running in part on eleventh to thirteenth soft rays; anal
fin with a broad yellow-orange streak edged in lavender from
origin of fin to tip of prolonged fourth ray; remainder of fin light
yellow; caudal fin light yellow with lavender upper and lower
margins basally, shifting to light blue distally; posterior edge of
caudal fin in central part with faint light blue margin; faint wavy
vertical dark-edged light blue band near middle of caudal fin;
pectoral fins light yellowish with large semicircular yellow-
orange spot edged in lavender at base; pelvic fins yellow, deeper
yellow laterally, with lavender lateral margin. Iris yellow with
two broad diagonal streaks of blue (dorso-anterior and ventro-
posterior to pupil).

Color of a 33-mm female from the atoll of Temoe, Tuamotu
Archipelago when fresh: body light magenta dorsally, shading to
pale lavender ventrally and white on abdomen and thorax (laven-
der mid-ventrally), except for a zone of bright yellow above a
line of demarcation from mid-base of dorsal fin to lower base of
caudal fin (a deep orange streak separates the yellow and
magenta from below base of fifth soft dorsal ray to anterior
caudal peduncle). Tip of lower jaw and front of snout bright yel-
low; upper postorbital head light magenta; prominent V-shaped
mark of deep yellow, edged with orange from apex at origin of
dorsal fin to dorso-posterior edge of orbit of either side. Dorsal fin
bright yellow; spines tipped with magenta; margin of anterior soft
portion of fin lavender; anal fin similar but entire fin becoming
lavender posteriorly; caudal fin light yellow, the upper and lower
edges narrowly magenta; pectoral fins pale; pelvic fins pale with
a broad deep yellow streak on first to third soft rays and as-
sociated membranes.

REMARKS: A. v. ventralis has been collected in the Pitcairn
Group, Tuamotu Archipelago, and New Caledonia from 40 to 55
m. In addition, my associates and I obtained one specimen in
46 m from a cave in a vertical drop-off at the west end of
Enewetak Atoll, Marshall Islands on July 5, 1975. This specimen
is unusual in having five opercular spines on one side and four on
the other, instead of the usual three, and mainly for this reason it
is not regarded as a paratype. In other respects it is typical of the
subspecies.

Populations of reef fishes in the Hawaiian Islands often are
distinct from elsewhere in Oceania. The differences may be so
slight as to not warrant nomenclatural recognition, or they may
be of equal magnitude to those that separate many sympatric
marine species. In the case of A. ventralis , recognition of the
Hawaiian variant as a subspecies seems to be the best treatment.
A. v. hawaiiensis differs in having slightly higher gill-raker
counts, fewer interopercular serrae, a deeper body on the average
(depth 2.37-2.94 compared to 2.85-3.13 ford. v. ventralis),
larger size (largest 76.9 mm, compared to 54.3 mm for A. v.

ventralis; since three of the four localities ford. v. ventralis are
from higher latitudes than Oahu, one cannot explain the larger
size as a cool- water effect), and color. The females in Hawaii do
not have such a sharp demarcation of the yellow dorsoposteriorly
on the body from lavender or magenta on the side, and the males
in Hawaii have an orange-red wash over the middle and anterior
part of the body (except ventrally) plus more orange and red in
the fins. It is possible, however, that a larger male than any taken
thus far outside Hawaii will have more red and orange color.

LITERATURE CITED

Bleeker, P. 1873. Revision des especes indo-archipelagiques du
groupe des Anthianini. Ned Tijdschr. Dierk. 4:155-169.

Brock, Vernon E. and Theodore C. Chamberlain. 1968. A
geological and ecological reconnaissance off western Oahu,
Hawaii, principally by means of the research submarine
“Asherah.” Pacific Sci. 22(3):373-394, 10 figs.

Clarke, Thomas A. 1972. Collections and submarine observa-
tions of deep benthic fishes and decapod Crustacea in
Hawaii. Pac. Sci. 36(3):310— 317.

Fowler, Henry W. 1923. New or little-known Hawaiian fishes.

B. P. Bishop Mus., Occ. Pap. 8(7):375-392.

Gosline, William A., and Vernon E. Brock. 1960. Handbook
of Hawaiian Fishes. Univ. Hawaii Press, Honolulu, 372
pp., 277 figs.

Herre, Albert W. 1927. A new genus and three new species of
Philippine fishes. Philip. Jour. Sci. 32(3):4 1 3—419, 2 pis.
Jordan, David S. and Barton W. Evermann. 1905. The
aquatic resources of the Hawaiian Islands. Part I. The shore
fishes. U. S. Bur. Fisheries, Bull. 23(l):xxviii + 574 pp.,
229 figs., 65 pis., 73 col. pis.

Jordan, David S. and Robert E. Richardson. 1910. A review
of the Serranidae or sea bass of Japan. U. S. Natl. Mus.,
Proc. 37(17 14):421 — 474, 16 figs.

Katayama, Masao. 1975. Caprodon unicolor, a new anthiine
fish from the North Pacific Ocean. Japan. Jour. Ichth.
22(1):13-15, 1 fig.

Randall, John E. In press. A preliminary revision of the fish
genus Plectranthias (Serranidae: Anthiinae), with descrip-
tions of 12 new species. Micronesica, 137 ms pp., 26 figs.

and Roger Lubbock. In press. A revision of the ser-

ranid fishes of the subgenus Mirolabrichthys (Anthiinae: An-
thias), with descriptions of six new species. Sci. Contr. Nat.
Hist. Mus. Los Angeles County, 65 ms pp., 1 text-fig., 17
col. figs.

Strasburg, D. W., E. C. Jones and R. T. B. Iversen. 1968.
Use of a small submarine for biological and oceanographic
research. Cons. Intern. Explor. Mer, Jour. 31(3):410-426,
7 figs.

Accepted for publication May 14, 1977.

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 302:1-13.

SERIAL PUBLICATIONS OF THE
NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY

Prior to November 30, 1973, publications of the Natural History Museum have appeared under various titles — Leaflet
Series, Museum Graphic, Education Series, History Series, Science Series, Study Guides, Contributions in History, History
Bulletin, Science Bulletin, unnumbered catalogs of exhibitions, and other miscellaneous publications.

The publication program of the Natural History Museum continues to publish a series of reports that give accounts of
new discoveries in history arid science as well as new advances in museum knowledge. The theme of the Natural History
Museum serials will emphasize basic research through the continuing active series:

Contributions in Science
Education Series
History Series
Science Series

the world.

reviewers.

Exposition Boulevard, Los Angeles, California 90007.

Number 303
March 16, 1979

CONTRIBUTIONS IN SCIENCE

NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY

THORACICAN CIRRIPEDIA OF THE LOWER PLIOCENE PANCHO RICO
FORMATION, SALINAS VALLEY, MONTEREY COUNTY, CALIFORNIA

By Victor A. Zullo

Published by the NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY • 900 EXPOSITION BOULEVARD • LOS ANGELES, CALIFORNIA 90007

CONTENTS

Abstract 3

Description of Localities 3

Systematic Descriptions, Suborder BALANOMORPHA
Pilsbury 1916, Family BALANIDAE Leach 1817,

Genus BALANUS Da Costa 1778, Subgenus BALANUS Da Costa 1778

Balanus gregarius (Conrad 1856) 5

Balanus proxinubilus new species 7

Balanus addicotti new species 8

Balanus crenatus leipochoma new subspecies 10

Subgenus SOLIDOBALANUS Hoek 1913

Balanus proinus Woodring 1950 H

Acknowledgments 1 1

Literature Cited j j

ISSN: 0459-8113

Suggested Citation: Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 303:1-13.

Contributions in Science are articles in the earth and life sciences, presenting results of original research in Natural
History. Science Bulletin (Numbers 1—30; 28 June 1965 to 10 April 1978) and Contributions in Science (Numbers 1—300;
23 January 1957 to 16 June 1978) were merged into a single imperial octavo serial beginning with Number 301.

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 303:1- 13.

THORACICAN CIRRIPEDIA OF THE LOWER PLIOCENE PANCHO RICO
FORMATION, SALINAS VALLEY, MONTEREY COUNTY, CALIFORNIA1

By Victor A. Zullo2

ABSTRACT: Five thoracican barnacles, including Balanus gregarius (Conrad 1856), B. proinus Woodring
1950, and three new taxa, B. proxinubilus , B. addicotti, and B. crenatus leipochoma, were found in associa-
tion with a nearshore warm temperate molluscan fauna. Balanus proxinubilus, which also occurs in the Upper
Pliocene San Diego Formation, is related to the extant Pacific Coast species B. nubilus Darwin 1854. Balanus
addicotti is allied to the extant pantropical species B. trigonus Darwin 1854, and B. crenatus leipochoma
differs but slightly from extant B. crenatus Bruguiere 1789. The Pancho Rico occurrence of B. proinus repre-
sents the oldest known record of this southern California Pliocene species.

The Pancho Rico Formation is of interest to students of fossil
Cirripedia in that it is probably the type locality for Balanus
(Balanus) gregarius (Conrad 1856). As first described by Conrad
(1856, 1857a), this large and unusual barnacle was indicated as
having been collected in Monterey County by A.S. Taylor, an
early resident of Monterey. Except for its presence in the north-
ernmost outcrops of the Upper Miocene Santa Margarita Forma-
tion near Vineyard Canyon, B. gregarius is known in Monterey
County only from the Pancho Rico Formation (text figure 1). Its
conspicuousness and abundance at several easily accessible
localities in a part of the Salinas Valley that was populated during
Taylor’s lifetime strongly suggest that the original specimen
came from the Pancho Rico, rather than the more southerly and
limited exposures of the Santa Margarita. The type specimen,
which presumably would have been deposited in the Academy of
Natural Sciences of Philadelphia, has not been located and is
probably lost.

The Lower Pliocene Pancho Rico Formation as redefined by
Durham and Addicott (1964, 1965) is characterized by fine
grained sandstone and lesser amounts of conglomerate and mud-
stone overlying Late Miocene or older rocks and underlying the
Pliocene or Pleistocene Paso Robles Formation. According to
Addicott (in Durham and Addicott 1965: A 18) the molluscan
fauna of the Pancho Rico Formation suggests “that the local
early Pliocene marine climate was generally warmer than modem
climates at comparable latitudes off the central California coast”
and that “living representatives or modern analogs [of the
Pancho Rico molluscan fauna] are usually found in shallow near-
shore areas of the northeastern Pacific Ocean.”

The barnacles available from the Pancho Rico Formation sug-
gest environmental conditions similar to those proposed for the
molluscan assemblage. Besides B. gregarius, four additional taxa
were present in collections examined. Balanus proinus Woodring
1950 is an extinct species commonly encountered in younger
Pliocene deposits in the San Joaquin Valley, the Santa Maria dis-
trict, and in San Diego County. A new species related to B.
trigonus Darwin 1854 was found at two localities. Balanus

trigonus is an extant tropical species with Pleistocene records in
the Caribbean and on the Pacific side of Mexico, indicating its
presence in the tropical American fauna prior to the late Miocene
closure of Central American and northern South American sea-
ways. A second new species is closely related to the extant
northeastern Pacific B. nubilus Darwin 1854 that presently ranges
from southeastern Alaska to northern Baja California, and has a
fossil record extending back to at least the late Pliocene in north-
ern California. The new subspecies is represented by seven
specimens attached to a fragment of the test of a (?)schizasterid
echinoid, and differs but slightly from extant B. crenatus Bru-
guiere 1789, whose range on the Pacific Coast extends from the
Bering Sea to Santa Barbara, California.

DESCRIPTION OF LOCALITIES

Natural History Museum of Los Angeles County, In-
vertebrate Paleontology Collection (LACMIP)

4660. North side of Wildhorse Canyon, NW 1/4 of SE 1/4 of Sec. 8,
T. 20 S., R. 9 E., San Lucas Quadrangle. Collectors: J.A.
Wolfe and V.A. Zullo, 1961 (= USGS locality M1341).

United States Geological Survey (USGS)

M903. Sandstone bluff on west side of State Highway 198, 425 feet
south, 425 feet west of NE corner of Sec. 5, T. 21 S., R. 9 E.,
San Lucas quadrangle. Collectors: D.L. Durham, 1959; W.O.
Addicott and D.L. Durham, 1964.

M913. North side of Wildhorse Canyon, 2650 feet north, 1075 feet
east of SW corner of Sec. 8, T. 20 S., R. 9 E., San Lucas
quadrangle. Collector: D.L. Durham, 1960.

'Review Committee for this Contribution:

Warren O. Addicott
Dora P. Henry
Edward Wilson

2Research Associate in Invertebrate Paleontology, Natural History Mu-
seum of Los Angeles County; and Department of Earth Sciences, Uni-
versity of North Carolina at Wilmington, Wilmington, North Carolina
28401.

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 303:1- 13.

Zullo: Pliocene Cirripedia

Text Figure 1 . Generalized locality map of barnacle sites in the Pancho
Rico Formation, Monterey County, California (adapted from Durham and
Addicott 1965).

M977. Base of bluff on south side of Pancho Rico Creek, 1000 feet
north, 3100 feet west of SE corner of Sec. 11, T. 22 S., R. 10
E., San Ardo 15 minute quadrangle. Collector: D.L. Durham,
1960.

M982. Roadcut on west side of U.S. Highway 101, 150 feet north,
2525 feet west of SE corner of Sec. 25, T. 23 S., R. 10 E.,
Wunpost quadrangle. Collectors: J.G. Vedder and C.A.
Repenning, 1960.

M988. 3150 feet north, 325 feet east of SE corner of Sec. 28, T. 19

S. , R. 9 E., San Lucas quadrangle. Collector: D.L. Durham,
1958-59.

M1935. North side of Walker Canyon, 900 feet north, 850 feet west of
SE corner of Sec. 28, T. 22 S., R. 1 1 E., Wunpost quadrangle.
Collectors: W.O. Addicott and D.L. Durham, 1963.

M1970. Railroad cut at mouth of small gully, 425 feet north, 175 feet
west of SE corner of Sec. 25, T. 23 S., R. 10 E., Wunpost
quadrangle. Collectors: W.O. Addicott and D.L. Durham,
1964.

M2275. Cut on north side of Bull Canyon Road, 100 feet south, 2400
feet east of NW corner of Sec. 26, T. 19 S., R. 9 E., Hernan-
dez Valley quadrangle. Collector: W.O. Addicott, 1964.

M2279. Float collection near head of small gully on north side of Wild-
horse Canyon, 2500 feet north, 300 feet west of SE corner of
Sec. 8, T. 20 S., R. 9 E., King City 15 minute quadrangle.
Collector: W.O. Addicott, 1964.

University of California Museum of Paleontology
(UCMP)

A-3425. On ridge at head of Sargent Canyon, NW 1/4 corner of Sec. 9,

T. 22 S., R. 12 E., Priest Valley 30 minute quadrangle.

A- 4945. Same as USGS locality 1341, LACMIP locality 4660.

A-4947. Highest sandstone bed exposed on hill about 250 yards south-
east of the center of the east half of Sec. 33, T. 20 S., R. 9 E.,
Priest Valley quadrangle (1915 edition).

A-7570. East branch of prominent south draining gully; from diatomite
just below contact with sandy barnacle zone, about 15 feet
above lowest outcrop in cliff. NE 1/4 Sec. 32, T. 19 S., R. 9
E., King City quadrangle (1941 edition). Collected September,
1951.

A— 7571. Locality same as A-7570, but from higher bed in west branch
of gully just below top of cliff.

A-8477. Eight-tenths of a mile east of the junction of U.S. Highway 101

and State Highway 198. First outcrop on north side of Highway
198 towards Priest Valley.

SYSTEMATIC DESCRIPTIONS

Suborder BALANOMORPHA Pilsbry 1916
Family BALANIDAE Leach 1817
Genus BALANUS Da Costa 1778
Subgenus BALANUS Da Costa 1778

Balanus gregarius (Conrad 1856)

Figures 1-9

Tamiosoma gregaria Conrad 1856:315; 1857a:72, pi. 4, fig. 18 [Mon-
terey County, California]; Gabb 1869:61, pi. 18, figs. 22a-d; Dali
1902:5.

Balanus estrellanus Conrad 1 857b: 1 95, pi. 8, fig. 1 [Estrella, California];
1877:156.

Radiolites gregaria Conrad 1864:214.

Balanus H estrellanus Conrad 1876:273.

? Balanus aquila Pilsbry 1907:199, pi. 8, figs. 5-8; 1916:127, pi. 31,
figs. 1-2, 4a, pi. 32, figs. 2- 2c, text figs. 34a-b, 35a-c [Mon-
terey Bay, California],

Balanus gregarius (Conrad). Pilsbry 1916:126, pi. 28, figs. 1-3, pi. 29;
Zullo 1964:360; Durham and Addicott 1965: All, A14, pi. 1, figs.
2, 3, 6, 8 [probably not pi. 2, figs. 4, 9, referred to Balanus sp. on
legend, pi. 2]; Zullo 1969:6, figs. 3-7, 45.

Balanus (Tamiosoma) cf. B. (T.) gregarius (Conrad). Woodring, in
Woodring, Stewart and Richards 1940:96, pi. 36, figs. 2-5, 8-9.
Balanus concavus concavus Bronn. Ross 1962:14, figs. 6-7.

OCCURRENCE: USGS Iocs. M903, M913, M977, M982,
M 1 935, M 1 970; UCMP Iocs. A-3425, A-4945, A-7570,
A-7571, A-8477; LACMIP loc. 4660.

Distribution: Early Miocene — Late Pliocene, central and
southern California, Baja California. (?)Pleistocene — Recent,
central California to northern Baja California (as B. aquila Pils-
bry 1907; see Zullo 1964).

REMARKS: Balanus gregarius is the most commonly encoun-
tered barnacle in the Pancho Rico Formation, and locally forms
‘reefs’ similar to those described by Woodring (in Woodring,
Stewart and Richards 1940) in the Etchegoin Formation of the
Kettleman Hills, Kings County, California. Preservation is ex-
cellent, and opercular valves retaining purple coloration on the
beaks and in the medial part of the interior of the valve are often
found either in life position or at the bottom of the sediment filled
shell. About half of the Pancho Rico specimens collected lack an
elongate, cellular basis. In those possessing such a basis, length
(with respect to total height of shell) and density of cellular fill-
ing varied greatly. A few bases up to 35 cm in length were
found, but most were in the range of from 8 to 15 cm.

Opercular valves obtained from shells with flat bases were not
observed to differ from those found in shells with elongate bases.
Similarly, Pancho Rico valves could not be differentiated from
those previously illustrated from the Upper Pliocene San Diego
Formation (Zullo 1969), or from those of Pleistocene or extant
specimens of B. aquila. As suggested earlier (Zullo 1964), it is
probable that B. aquila is a junior synonym of B. gregarius. The
opercular valves of B. gregarius described and illustrated from
the Kettleman Hills by Woodring (in Woodring, Stewart and
Richards 1940) are also similar in form, but are not representa-
tive of the Etchegoin population. Bulk collections (which in-
cluded over 200 opercular valves) made by the author from

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 303:1-13.

Zullo: Pliocene Cirripedia

Figures 1-9. Opercular valves of Balanus ( Balanus ) gregarius (Conrad). (1) exterior of scutum, hypotype LACM1P
4926; (2) exterior of tergum, hypotype LACMIP 4927; (3) exterior of scutum, hypotype LACMIP 4928; (4) interior of
scutum, hypotype LACMIP 4929; (5) exterior of tergum, hypotype LACMIP 4930; (6) interior of scutum, hypotype
LACMIP 4931; (7) interior of tergum, hypotype LACMIP 4932; (8) interior of tergum, hypotype LACMIP 4933; (9)
exterior of tergum, hypotype LACMIP 4932 [Height of tergum, fig. 2, 21 mm; all valves to same scale.]

Contrib. Sci. Naiur. Hist. Mus. Los Angeles County. 1979. 303:1-13.

Zullo: Pliocene Cirripedia

Figures 10-17. Balanus (Balanus) proxinubilus, new species. (10) exterior of tergum, holotype LACMIP 4934, height, 18
mm; (11) interior of same; (12) three shells of ribbed form, paratypes LACMIP 4937, 4938, 4940, carinorostral diameter of
orifice, paratype LACMIP 4938 (center of photograph), 29 mm; (13) interior of scutum, holotype LACMIP 4934, height, 20
mm; (14) interior of scutum, holotype LACMIP 4934, height, 19 mm; (15) interior of tergum, holotype LACMIP 4934,
height, 19 mm; (16) exterior of scutum shown in fig. 13; (17) shells of smooth form, including holotype LACMIP 4934
(second from right) and paratypes 4935, 4936, 4941, 4942, carinorostral diameter of orifice of paratype LACMIP 4935
(second from left), 27 mm.

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 303:1-13.

Zullo: Pliocene Cirripedia

Woodring’s localities indicate that the Etchegoin population dif-
fers in two respects: 1) the tergal spur usually is flush with the
basal margin, rather than being elevated above the basal margin
by the closure of the spur furrow; and 2) the articular ridge of the
scutum usually is joined to the adductor ridge, rather than being
clearly separated. These differences are not constant, even in
specimens from the same lot, but they suggest the possibility of
the presence of a distinct race of B. gregarius in the San Joaquin
embayment during Pliocene time.

The tergum figured by Durham and Addicott (1965, pi. 2,
figs. 4, 9) from the Pancho Rico Formation was referred to B.
gregarius in the text and to Balanus sp. in the plate legend. The
.specimen does not appear to represent a tergum of B. gregarius.
The characteristic apical beak is missing, and the apex is neither
broken nor significantly worn. There is no pit entering the apex
on the carinal side of the articular ridge, the exterior surface of
the valve lacks radial striae (especially prominent on the scutal
side in B. gregarius ), and the tergal spur is shorter and broader.
In general form this tergum resembles those of either the B. paci-
ficus Pilsbry 1916 — B. concavus Bronn 1831 group, or of the
subgenus Megabalanus Hoek 1913. An undescribed species,
related to B. pacificus and with a similar tergum, occurs in asso-
ciation with B. gregarius in the Upper Miocene Santa Margarita
Formation in San Luis Obispo County, California (UCMP collec-
tion). The megabalanid B. wilsoni Zullo 1969 and some ques-

tionable specimens of B. pacificus are found with B. gregarius in
the Upper Pliocene San Diego Formation.

Balanus proxinubilus new species

Figures 10-20

Balanus n. sp.? Addicott, in Durham and Addicott 1965:14.

Balanus nubilus Darwin. Zullo 1969:8, figs. 8- 10.

DIAGNOSIS: Large, high conic Balanus with open parietal
tubes; scutum as in B. nubilus Darwin, but with smaller, circular
adductor muscle pit placed higher on valve, and broader, lower
adductor ridge; tergum as in B. nubilus, but with shorter, broader
tergal spur placed farther from the basiscutal angle, and resulting
marked prolongation of the scutal half of the basal margin, whose
juncture with the spur is angular.

DESCRIPTION: Shell large, high conic; orifice toothed, sub-
triangular; exterior of parietes nearly smooth to rugose; radii
moderately wide with oblique summits; sutural edges of radii
with complex pattern of primary and secondary denticulae; alae
broad, projecting slightly above radii; basis thick, porous, cling-
ing firmly to compartmental plates; interior of parietes ribbed
from sheath to basis; sheath one-half to two-thirds length of com-
partmental plates, with deep cavity under lower edge; parietal
tubes large, numerous, open throughout length.

Figures 18-20. Balanus (Balanus) proxinubilus , new species. (18-19) exterior of terga, paratype
LACMIP 4935, height, 20 mm; (20) basal view of paries, paratype LACMIP 4939, length (from
left to right), 20 mm.

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 303:1-13.

Zullo: Pliocene Cirripedia

Figures 21-22. Exterior of terga of Balanus (Balanus)
nubilus Darwin, illustrating lack of scutad prolongation
of valve (after Pilsbry 1916, pi. 31, figs. 3,4).

Scutum sculptured externally by prominent, widely spaced
growth lines; tergal margin not reflected; basitergal angle nearly
90 degrees; articular ridge moderately developed, reflected,
slightly over half length of tergal margin; articular furrow broad,
shallow; adductor ridge indistinct, low, merging with articular
ridge above to form low swelling between articular ridge and
adductor muscle pit; adductor muscle pit small, well defined, cir-
cular, placed high on valve; depressor muscle pit large, triangu-
lar, with distinct longitudinal ridge bisecting pit; occludent mar-
gin bearing prominent teeth formed by alternate growth incre-
ments; tergal margin one-third again as long as basal margin;
scutum slightly concave between basal margin and apex.

Tergum broad, triangular; articular ridge blunt, not reflexed,
long; articular furrow broad, shallow; scutal margin prominently
reflexed; crests for lateral depressor muscles well developed,
thin, long, about six in number; length of basal margin equal to
height of tergum excluding spur; width of tergal spur equals
three-eights length of basal margin; spur V-shaped, forming
angle slightly greater than 90 degrees with basal margin on scutal
side, less angulate on carinal side; spur placed centrally on basal
margin (about its own width from basiscutal angle), giving an
elongate appearance to scutal half of tergum; spur fasciole delim-

ited by grooves on either side.

DIMENSIONS:

Height

Carinorostral diameter
of orifice

Holotype LACMIP 4934:

37 mm

22 mm

Paratype LACMIP 4935:

50 mm

27 mm

Paratype LACMIP 4938:

54 mm

29 mm

DISPOSITION OF TYPES: Holotype and paratypes are depos-
ited in the Invertebrate Paleontology collection. Natural History
Museum of Los Angeles County.

OCCURRENCE: Lower Pliocene, Pancho Rico Formation,
LACMIP locality 4660 (type locality, = UCMP loc. A-4945,
USGS loc. M1341), and USGS loc. M913; Upper Pliocene, San
Diego Formation, LACMIP loc. 305.

ETYMOLOGY: A compound word from the Latin, meaning
nearest nubilus.

REMARKS: Balanus proxinubilus is quite similar in appear-
ance to the extant Pacific Coast species B. nubilus. The shells of
the two species are virtually identical in form. The scuta of B.
proxinubilus differ somewhat in having a lower, broader adductor
ridge and a smaller adductor muscle pit. The tergum of B. proxi-
nubilus, however, can be distinguished readily from that of B.
nubilus. The marked prolongation of the basiscutal part of the
tergum, the short, broad tergal spur, and the angularity of the
juncture between the spur and the basal margin on the scutal side
give to the tergum a distinctive appearance that is not found in
extant and fossil examples of B. nubilus.

The fragmentary terga from the Upper Pliocene San Diego
Lormation referred to B. nubilus in an earlier paper (Zullo 1969)
are similar in form to the Pancho Rico terga, and also appear to
represent B. proxinubilus . Balanus nubilus is known from depos-
its of Late Pliocene age on the Pacific Coast, but unlike B. proxi-
nubilus, the species is associated with a cool water fauna in
northern California. By Pleistocene time, B. nubilus was widely
distributed, as evidenced by the numerous examples available
from rocks of this age from Oregon south to Baja California.
Balanus proxinubilus thus appears to have been a southern, warm
water Pliocene species that may well have succumbed to the
southern advance of B. nubilus during the Pleistocene.

Balanus addicotti new species

Figures 23-32

DIAGNOSIS: Small, high conic, ribbed Balanus with broad
radii; external growth ridges of scutum cut by deep radial striae,
giving marked cancellate appearance to exterior of valve; scutal
adductor ridge short, stout and high; tergum with short, broad,
basally truncate spur and open spur furrow.

DESCRIPTION: Shell small, high conic; orifice untoothed,
subtrigonal to diamond shaped; parietes regularly ribbed; parietal
tubes small, rounded, numerous, without transverse septa, but
filled in upper half; radii broad, solid, thick, with horizontal
summits; sutural edges of radii coarsely denticulate; alae thinner,
with oblique summits; sheath about one-half height of shell,
lower edge dependent; interior of parietes sharply ribbed from
basis to lower edge of sheath; basis moderately thick, with small,
radiating non-septate tubes.

Scutum thick, narrow; exterior ornamented by closely spaced
growth ridges crossed by deeply incised radial striae that cut
growth ridges into a series of high nodes; tergal border reflexed
90 degrees; adductor ridge high, short and stout, situated in cen-
ter of valve, and with a small pit between it and the tergal mar-
gin; adductor muscle pit oval, shallow, situated close to occlud-
ent margin; depressor muscle pit large, deep; articular ridge two-
thirds to three-fourths length of tergal margin, broadly arched,
and reflexed over deep, narrow, articular furrow.

Tergum thin, broad; tergal spur short, broad, situated close to
basiscutal angle, and truncate basally; spur width from one-half
to two-thirds width of basal margin; juncture of spur with basal
margin angular on both sides; spur furrow broad, open; articular

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979 . 303:1-13.

Zullo: Pliocene Cirripedia

Figures 23-26. Balanus (Balanus) addicotti, new species, USGS loc. M1935. (23) Holotype USNM 248194 (center,
bottom) and paratypes USNM 248194/a-d, carinorostral diameter of orifice of holotype, 4.5 mm; (24) group of shells,
paratype lot USNM 248195, length of cluster, 21 mm; (25) group of shells, USNM 248196, maximum height of
photographed portion of cluster, 35 mm; (26) interior of shell from paratype lot USNM 248195 (fig. 24) showing basal tubes,
parietal ribbing, and sheath.

ridge low, short; articular furrow broad, shallow; depressor mus-
cle crests numerous, short, dependent below basal margin.

DIMENSIONS: Height Greatest Diameter

Holotype USNM 1 1 mm 9 mm

Paratype USNM 1 8 mm 12 mm

Paratype USNM 2 6.5 mm 10 mm

DISPOSITION OF TYPES: Holotype and described paratypes
deposited in the United States National Museum; other paratypes
deposited in the University of California Museum of Paleontol-
ogy-

OCCURRENCE: Lower Pliocene, Pancho Rico Formation,
USGS Iocs. M1935 (type locality) and M2275; UCMP loc.
A-7570.

ETYMOLOGY: This species is named for Warren O. Addi-
cott.

REMARKS: Balanus addicotti bears marked resemblance to
the extant tropicopolitan species B. trigonus Darwin. It differs
from this species in the less distinctly trigonal shell orifice, in the
cancellate rather than pitted sculpture of the scutum, and in the
stouter and more pronounced scutal adductor ridge. The cancel-
late ornamentation of the scutum is like that of 8. spongicola
Brown 1827, but this species differs in lacking a distinct scutal
adductor ridge, and in having a much narrower tergal spur and a
smooth shell wall. The opercular valves of the tropical American
species B. calidus Pilsbry, 1916 are similar to those of B. addi-
cotti, but the radial striae of the scutum of B. calidus are not
deeply incised and thus do not impart a cancellate appearance to
the surface of the valve. Also, the radii of B. calidus are much
narrower, and the shell wall is plicate rather than regularly
ribbed. Balanus addicotti is also similar to B. kanakoffi Zullo

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 303:1-13.

Zullo: Pliocene Cirripedia

Figures 27-32. Opercular valves of Balanus ( Balanus ) addicotti, new species, holotype USNM 248194,
USGS loc. M1935. (27) exterior of scutum, height, 3.8 mm; (28) exterior of tergum, height, 3.5 mm; (29)
exterior of scutum, height, 3.8 mm; (30) interior of scutum of fig. 29; (31) interior of tergum of fig. 28;
(32) interior of scutum of fig. 27.

1969, which is*known only from the Upper Pliocene San Diego
Formation of southern California. However, B. kanakoffi has a
plicate shell with narrow radii (the suture between the carina and
carinolaterals being linear or obscured), and a beaked, narrow
tergum. In addition the scutum of B. kanakoffi lacks a distinct
adductor ridge.

The extant and fossil distribution of B. trigonus in tropical
American waters indicates that this species ranged at least as far
back as the Late Miocene in the eastern Pacific, and was thus
probably present along the southern and central California coast
during the time of deposition of the Pancho Rico Formation. Bal-
anus addicotti may have been a geographically and temporally
localized derivative of B. trigonus that became extinct upon
withdrawal of the sea from the Pliocene Salinas Valley embay-
ment. Balanus addicotti may yet be discovered in rocks of Jaca-
litos age on the eastern side of the San Andreas fault, but it does
not appear to have been present in younger Pliocene warm water
faunas of the San Diego Formation (Zullo 1969) and of the
Etchegoin and San Joaquin Formation of the Kettleman Hills
(unpublished data).

Balanus crenatus leipochoma

NEW SUBSPECIES
Figures 33-37

DIAGNOSIS; Small, low conic shell with non-septate parietal

tubes and narrow, oblique radii; interior of parietes ribbed; basis
thin, nonporous; space behind dependent basal margin of sheath
bearing cellular secondary deposits; scutum broader than tall, ex-
ternally concave, with prominent articular ridge and indistinct
adductor ridge; tergum broad, with broad, short, subtruncate
spur.

DESCRIPTION: Shell small, low conic; orifice of moderate to
large size, diamond shaped; parietes thin, smooth, or plicate in
basal third; parietal tubes square, without transverse septa; radii
narrow, oblique; alae wider than radii and projecting above
summits of radii; sheath one-half to two-thirds length of com-
partment; space under lower edge of sheath crudely septate or
secondarily filled; interior of parietes ribbed; basis thin, solid.

External surface of scutum concave, ornamented solely by
growth increments; basal margin slightly longer than tergal mar-
gin; articular ridge prominent, nearly erect, occupying slightly
more than half of the tergal margin, articular furrow deep, broad;
adductor ridge indistinct, basically a calloused region between
adductor muscle pit and articular ridge; adductor muscle pit
prominent, deep, lenticular in shape; depressor muscle pit shal-
low, triangular.

Tergum broad, thin; tergal spur broad, short, near basiscutal
angle, and rounded basally; spur width about one-half that of
width of basal margin; juncture of spur with basal margin gently
curved on both sides; spur furrow broad, open; articular ridge
prominent, not reflexed, restricted to upper half of valve, artic-
ular furrow broad, deep; five short, well developed depressor

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 303:1-13.

Zullo: Pliocene Cirripedia

muscle crests depend slightly below basal margin.

DIMENSIONS: Height Greatest Diameter

Holotype UCMP 10929 4 mm 8.5 mm

DISPOSITION OF TYPES: Holotype and six paratypes depos-
ited in the University of California Museum of Paleontology col-
lection.

OCCURRENCE: Lower Pliocene, Pancho Rico Formation,
UCMP loc. A- 4946, on fragment of (?)schizasteroid echinoid
test.

ETYMOLOGY: Subspecific epithet a compound word from
the Greek leipo, ‘without’ and choma, ‘bulwark’.

REMARKS: It is perhaps inappropriate to characterize these
fossil specimens as distinct from typical B. crenatus. The oper-
cular valves are unremarkable and the shell, in all features but
one, is that of B. crenatus. However, examination of the parietal
tubes of several fragmental shells associated with the type lot,
and of the holotype itself revealed no trace of transverse septa; a
feature characteristic of typical B. crenatus and its known sub-
species.

The opercular valves and external shell form are easily con-
fused with those of B. proinus. However, none of the over two
hundred shells of B. proinus examined exhibited any trace of
parietal tubes or of secondary cellular deposits below the depen-
dent margin of the sheath.

Subgenus SOL1DOBALANUS Hoek 1913
Balanus proinus Woodring 1950

Figures 38-41

Balanus hesperius Pilsbry, var Woodring, in Woodring, Stewart and
Richards 1940:30, 97.

Balanus hesperius proinus Woodring, in Woodring and Bramlette 1950:
92, pi. 14, figs. 11, 15, pi. 16, figs. 1-3, 8- 12.

Balanus proinus Woodring. Zullo 1969:16, figs. 26-32.

OCCURRENCE: UCMP Iocs. A-4947, A-7570.

DISTRIBUTION: Pliocene, southern California: Lower Plio-
cene, Pancho Rico Formation, Salinas Valley; ‘Middle’ Pliocene,
Foxen Mudstone, Santa Maria district; Upper Pliocene, San Joa-
quin Formation, Kettleman Hills, Careaga Sandstone, Santa
Maria district (type locality), San Diego Formation, San Diego
County.

REMARKS: The specimens of B. proinus from the Pancho
Rico Formation represents the oldest recognized occurrence of
the species. It is apparently as common in Pliocene deposits of
southern California as is B. gregarius but, because of its diminu-
tive size, is not as well known. Shells are often found attached to
pectens and occasionally to clypeasteroid echinoids and gastro-
pods, but they have not been observed in association with other
barnacles.

Balanus proinus is the youngest known member of a Tertiary
tropical to warm temperate eastern Pacific group that goes back
to the Late Eocene (Zullo 1966). There are no known warm
water survivors, but the extant boreal Pacific species B. hesperius
Pilsbry 1916 appears to be a derivative of this lineage.

ACKNOWLEDGMENTS

I wish to thank Warren O. Addicott and Jack A. Wolfe of the
U.S. Geological Survey, Menlo Park, California for their help in
obtaining the specimens used in this study.

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 303:1-13.

LITERATURE CITED

Bronn, H.G. 1831. Italiens Tertiar-Gebilde und deren orga-
nische Einschliisse. K. Groos, Heidelburg, xii + 176 pp., 4
text figs.

BROWN, T. 1827. Illustrations of the recent conchology of Great
Britain and Ireland, with the description and localities of all
the species, marine, land and fresh-water. London, 6 pp.,
53 pis.

Bruguiere, J.G. 1789. Histoire naturelle des Vers. Encyclopedie
Methodique 1(1): 1-134.

Conrad, T.A. 1856. Descriptions of three new genera; twenty-
three new species Middle Tertiary fossils from California,
and one from Texas. Acad. Nat. Sci. Philadelphia, Proc.
8:312-316.

. 1857a. Description of the Tertiary fossils collected on

the Survey. Pacific Railroad Reports 6 (Geol. Rpt.
2):69— 73, pis. 2-5.

. 1857b. Report on the paleontology of the Survey. Ibid.

7:189-196, pis. 1-10.

.1864. Notes on shells, with descriptions of new fossil

genera and species. Acad. Nat. Sci. Philadelphia, Proc.
16:211-214.

. 1876. Note on a cirripede of the California Miocene,

with remarks on fossil shells. Ibid. 28:273-275.

. 1877. Note on the relations of Balanus estrellanus of the

California Miocene. Amer. Jour. Sci. (3)13:156- 157.

Da Costa, E. 1778. Historia Naturalis Testaceorum Britanniae,
vol. 1. London, 254 pp., 200 text figs., 17 pis.

Dall, W.H. 1902. On the true nature of Tamiosoma. Science
(n.s.)15:5-7.

Darwin, C. 1854. A monograph on the sub-class Cirripedia, 2.
Balanidae, Verrucidae. Ray Soc., London, viii + 684 pp.,
1 1 text figs., 30 pis.

Durham, D.L. and W.O. Addicott. 1964. Upper Miocene and
Pliocene marine stratigraphy in southern Salinas Valley,
California. U.S. Geol. Surv. Bull. 1194E: E1-E7.

. 1965. Pancho Rico Formation Salinas Valley, Califor-
nia.U.S. Geol. Surv. Prof. Paper 524A:i— iii , A1-A22 text,
figs. 1-7, pis. 1-5.

Gabb, W.M. 1869. Cretaceous and Tertiary fossils. California
Geol. Surv., Paleontology 2:i-xiv, 1-299, pis. 1-36.

Hoek, P.P.C. 1913. Cirripedia of the Siboga Expedition, B. Cir-
ripedia Sessilia. Siboga-Expeditie 31b: 129— 275, pis. 11-27.

Leach, W.E. 1817. Distribution systematique de la classe des
Cirripedes. Jour. Phys. Chim. His. Nat. 85:67-69.

Pilsbry, H.A. 1907. Cirripedia from the Pacific coast of North
America. Bull. Bur. Fisheries 26(617): 193— 204, figs. 1-4,
6-11.

. 1916. The sessile barnacles (Cirripedia) contained in the

collections of the U.S. National Museum, including a
monograph of the American species. U.S. Natl. Mus., Bull.
93:i-xi, 1-366, text figs. 1-99, pis. 1-76.

Ross, A. 1962. Results of the Puritan-American Museum of
Natural History Expedition to Western Mexico, 15. The lit-
toral balanomorph Cirripedia. Amer. Mus. Novitates
2084:1-44, figs. 1-24.

Zullo: Pliocene Cirripedia

Figures 33-37. Balanus (Balanus) crenatus leipochoma, new subspecies, UCMP loc. A-4946. (33) Shells of type lot
(holotype UCMP 10929 and paratypes UCMP 10930a-f), lateral paries of holotype ground to show non-septate tubes;
(34-35) tergum of holotype UCMP 10929, height, 2.1 mm; (36-37) scutum of holotype UCMP 10929, height, 2.2 mm.
Figures 38-41. Balamis ( Solidobalanus ) proimts Woodring, UCMP loc. A-4947. (38) exterior of tergum, hypotype UCMP
10931, height, 2.5 mm; (39) interior of scutum, hypotype UCMP 10931, height, 1.6 mm; (40) interior of tergum of fig. 38;
(41) group of shells on Lyropecten estreUanus (Conrad), hypotype lot UCMP 10932, height, 51 mm.

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 303:1-13.

Zullo: Pliocene Cirripedia

Woodring, W.P, and M.N. Bramlette. 1950. Geology and
paleontology of the Santa Maria District, California. U.S.
Geol. Surv. Prof. Paper 222:i-iv, 1-185, text figs. 1-9,
pis. 1-23.

. R. Steward and R.W. Richards, 1940. Geology of

the Kettleman Hills oil field, California. U.S. Geol. Surv.
Prof. Paper 195:1-170, text figs. 1-15, pis. 1-57.

Zullo, V.A. 1964. Re-evaluation of the late Cenozoic cirriped
“Tamiosoma” Conrad. Biol. Bull. 127(2):360 (abstr . ) .

. 1966. A new species of Balanus (Cirripedia, Thoracica)

from the Late Eocene Cowlitz Formation of southern Wash-
ington, U.S. A. Crustaceana 11:198-204, text figs. 1-2.

. 1969. Thoracic Cirripedia of the San Diego Formation,

San Diego County, California. Los Angeles Co. Mus., Con-
trib. Sci. 159:1-25, figs. 1-77.

Accepted for publication June 16, 1977.

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 303:1-13.

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Bv LouElla R. Saul

Published by the NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY • 900 EXPOSITION BOULEVARD • LOS ANGELES, CALIFORNIA 90007

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Suggested Citation; Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 304:1-8.

Contributions in Science are articles in the earth and life sciences, presenting results of original research in Natural
History. Science Bulletin (Numbers 1—30; 28 June 1965 to 10 April 1978) and Contributions in Science Numbers 1—300; 23
January 1957 to 16 June 1978) were merged into a single imperial octavo serial beginning with Number 301.

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 304:1-8.

A HOLLOW SPINED ANAPACHYDISCUS PENINSULARIS
WITH POSSIBLE MOSASAUR BITE IMPRESSIONS1

By LouElla R. Saul2

ABSTRACT: A specimen of Early Maestrichtian age Anapachydiscus peninsularis (Anderson and Hanna) from
Arroyo Santa Catarina, Baja California, Mexico, has nearly round holes punched in the shell so arranged that
they suggest mosasaur bite marks. The early whorls are ornamented with hollow lateral spines which could
have served as vertical stabilizers.

A concretion of roughly 20 cm diameter protruding from the
near vertical side of a gully cut into Rosario Formation siltstone
was overlooked by local entrepreneurs who, in 1968, collected
for use in the manufacture of lamp bases, etc., all exposed or
near surface ammonites from the well-known locality in Arroyo
Santa Catarina, Baja California, Mexico. A blow or two with
sledge and chisel split the concretion and freed from the rock a
well-preserved ammonite, Anapachydiscus peninsularis (Ander-
son and Hanna). Matrix plugs remained in the umbilicus, and a
lump of siltstone adhered to and filled what remained of the
broken body chamber. Four curiously round holes, two on each
side marred the specimen. Their shape, spacing and alignment
are reminiscent of those in the mosasaur-bitten ammonite
described by Kauffman and Kesling (1960). The siltstone matrix
at the apertural end was sawed away and most of the plug in the
umbilicus on the right side was removed so that the specimen
might be used to display evidence of interaction between ani-
mals. Removal of the matrix in the umbilicus exposed one more
roundish hole in the shell and, bordering the umbilicus, hollow
spines which have not been previously noted in descriptions of
this species (Anderson and Hanna 1935, p. 20, pi. 4, fig. 1; pi.
5, fig. 1-2; pi. 6, fig. 3-4; pi. 7, fig. 5; text-fig. 1; Anderson,
1958, p. 225; Matsumoto, 1959, p. 38).

THE ROUND HOLES

The holes are not truly round, but give the impression of hav-
ing been punched by round objects intersecting the curved,
ribbed surface of the ammonite shell. In addition to the five
roundish holes — two on one side, three on the other — the
specimen has a small angular hole (Fig. 6) on the abapertural side
of the umbilicus that could have been made by the tip of a tooth.
The shell is not depressed around the holes as in the specimen
described by Kauffman and Kesling (1960), but each hole has
cracks radiating from it. The holes are in the chambered portion
of the conch, and the edge of the shell along which most of the
body chamber broke off resembles a tear along perforations. If
the rounded embayments of this edge are considered to represent
half-tooth marks, there are ten tooth marks. Figures 1 and 2 dia-
gram two possible bites. The plump rounded whorls of A. penin-
sularis are not as advantageously shaped for displaying the bite
of a mosasaur as is the discoidal shape of the Placenticeras of

Kauffman and Kesling (1960), and other sortings of the holes
into bites can be argued for. Camp (1942) discussed two genera
of mosasaurs from the Maestrichtian of California, Plotosaurus
and Pleisiotylosaurus, but the limited record of these “bites”
(Figs. 1-3, 5-6) does not indict either. This probable mosasaur
may have regularly dined on ammonite as it seems to have been
more efficient about acquiring the benefits of its chase than did
the one inferred by Kauffman and Kesling. The Baja California
mosasaur apparently required only two bites to break off the body
chamber and thus free the edible soft parts rather than the 16
bites shown on the Placenticeras. An even more adept mosasaur
might bite just at the rear of the body chamber and leave as a
record of its predation only the torn-on-perforations shell edge.

THE HOLLOW SPINES

The spines on the specimen, UCFA cat no. 38765, are formed
by a thin outer layer of the shell (3rd abapertural spine of figure 5
has wall ,3mm thick) and are readily removed along with the
matrix leaving as their bases of attachment, bordering the umbil-
icus, the flat-topped tubercles of previous descriptions (Ander-
son and Hanna 1935, p. 20; Anderson 1958, p. 225). The maxi-
mum diameter of this specimen is 11.8 cm; the last flat-topped
tubercles occur at a diameter of about 7.3 cm. The spines and
spine bases have been best exposed on the last whorl of their
occurrence; here they are not evenly spaced; they occur on every
second to fourth rib. They appear to be more regularly spaced on
earlier whorls. The tallest spine (incomplete) is 8 mm high; it is
the next to the last adapertural one preserved (figure 5) and is
filled with crystalline calcite. The last spine is 6.5 mm high, has
the tip closed, and is probably nearly complete. The outermost
end of all the earlier formed ones were damaged in preparation.

'Review Committee for this Contribution
Richard Cowen
Michael A. Murphy
Edward Wilson

2Senior Museum Scientist, University of California at Los Angeles,
Department of Earth and Space Sciences; and Research Associate in
Invertebrate Paleontology, Natural History Museum of Los Angeles
County.

Contrib. Sci. Natur. Hist. Mus. Los Angeles County 1979. 304:1-8.

Saul: Ammonite ( Anapachydiscus )

TABLE 1

Measurements (in mm.) of Anapachydiscus peninsularis (Anderson and Hanna.)

cat. no.

maximum

last flat-topped tubercle
W D

W/D

CAS 4250

41.5

27.5

too poorly preserved

.66

paratype, poorly preserved

CAS 4253

(60)

paratype

UCLA 28717

107

29.5

(54)

.54

hypotype

UCLA 38765

118

67.5

(89)

.57

hypotype, with spines and holes

CAS 4249

120

(79)

.53

paratype, one spine showing?

CAS 4257

123

(85)

.58

paratype

CAS 4248

355

144

too poorly preserved

.41

holotype

D = diameter; (D) = diameter calculated from width of whorl; W = width of whorl; CAS = California Academy of
Sciences; UCLA = University of California, Los Angeles. The average diameter at which A. peninsularis apparently ceased
to generate spines was 73 mm.; however, two specimens, 4253 and 28717, ceased notably sooner at an average diameter of
57 mm. than the other three measurable specimens which ceased at an average of 84 mm. The gap in estimated size at which
they ceased to grow spines may result from measuring too few specimens or it may be an indication of sexual dimorphism.
The ratio W/D shows a loss of inflation with increase in size.

some because these structures were unexpected and the others
because they are so frail that they adhered to the matrix being
removed. All except the two abapertural spines are filled with
siltstone and thus were broken prior to entombment. Earlier
formed spines appear to have been longer relative to the size of
the ammonite shell than the later ones. The thin-walled, hollow
spines were sealed off at their bases by the formation of the next

Figures 1-3. Anapachydiscus peninsularis (Anderson and Hanna) with
aligned rounded holes. Holes of near side vertically lined, of far side
horizontally lined. Ammonite in presumed living position, arrows point
up. Because of the size of the ammonite and its inflation, too few “tooth
marks” are present to determine direction of bite, but two bites seem to
be required by the positions of the holes. Figures 1 and 2 diagram two of
several possible positions for the bites. Figure 1. Bite a caught the
ammonite and began to break off the body chamber. Figure 2. Bite b
broke off the body chamber. Figure 3. All holes indicated and body
chamber restored. Inferred body chamber length from Raup (1967, text-
fig. 16) is minimum length and results in most horizontal bite. A longer
body chamber rotates the diagrammed ammonite counter-clockwise and
suggests attack from above.

inner shell layer. The umbilical wall of the enveloping whorl
does not touch the spines; there is now a thin layer of siltstone
between spine and umbilical wall, and the spines apparently
stood free.

The original type lot of Parapachy discus peninsularis Ander-
son and Hanna comprises five specimens. Of these, three are not
well enough preserved to show spines or flat-topped tubercles.
The smallest specimen, California Academy of Sciences cat. no.
4250, has strong ribs with obvious tubercles bordering the umbil-
icus; but if the tubercles had rimmed flat-tops, weathering has
obliterated this detail. The specimen is, however, because of the
strength of the ribs, listed in Table 1 as having been spiny
throughout. With spines of equivalent length to those on UCLA
cat. no. 38765, it would have had a ventral profile like that in
figure 4. One of the paratypes, CAS cat. no. 4249, has a circlet
of shell that is probably a spine showing through the matrix in the
umbilicus. As can be seen from Table 1, the size at which A.
peninsularis ceased to grow spines was not consistent. UCLA
cat. no. 28717 (Popenoe 1954, fig. 3 (9) ) had a width of only
29.5 mm when its last spine formed; but UCLA cat. no. 38765
has its last flat-topped tubercle at a width of 51 mm. The differ-
ence between the two specimens of least diameter and the three
of larger diameter may reflect sexual dimorphism, but the sample
is too small to be significant.

An Anapachydiscus was collected by John Alderson from the
NW'A of section 6, T24S, R16E, Cholame 7 Vi min. quad., Mon-
terey Co., California (UCLA loc. 6337), or approximately 1 mile
westward along strike from LSJU (Stanford University) loc.
3354, from which Matsumoto (1959, p. 38; 1960, p. 127) identi-
fied A napachydiscus cf. A. arrialoorensis (Stoliczka). Alderson’s
specimen (UCLA cat. no. 57245) is plumper and has closer
spaced stronger ribs than specimens of A . peninsularis from Baja
California, and thus resembles Stoliczka’s (1865, p. 126, pi. 63,
fig. 2-4; pi. 64, fig. 1) figures of Ammonites arrialoorensis. It
differs from these figures in having more elliptical spine bases.

The outer whorl(s?) of Alderson’s Cholame specimen was
broken and eroded and discarded in the field leaving a specimen,
septate throughout, with diameter 90.5 mm and width 53 mm.
Most of one volution is present, the inner whorls being broken. A
flange of shell that formed the umbilical flank of the next outer

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 304:1-8.

Saul: Ammonite (Anapachydiscus)

volution remains on the right side of the specimen. The thickness
of this umbilical flank shell is about .6 mm. Broken hollow
spines are present at a diameter of 68 mm, width approximately
33 mm. The spine wall is almost one-third the thickness of the
flank wall. Spines appear to have been formed at a diameter of
87 mm, width 49 mm, but preservation and breakage are such
that I cannot be sure of their presence or absence. Also the spines
are less obvious than on UCLA cat. no. 38765 because in the
Cholame specimen the enveloping umbilical wall nearly engulfs
them. Considering the size to which some Anapachydiscus grew,
they seem to have had remarkably thin shells. Their shell walls
were about one-half as thick as those of Nautilus pompilius and
only a third thicker than that of an argonaut of similar diameter.

The spines were apparently secreted at the apertural edge as
part of the outermost layer of the conch. There is no evidence
that the leading side of the spine was open as is commonly the
case with spines arming the aperture of a gastropod. Compared to
spines on Mutex spp., the spines on Anapachydiscus have a wall
that is one-half to one-third as thick and is of equal thickness all
around. The diameter of the hollow center is approximately 3
times the thickness of the spine wall. Spines on Murex spp. have
adaperturally placed holes whose diameters are one-third or less
the thickness of the spine wall. The sealing off of the bases of the
Anapachydiscus spines probably took place soon after their
formation as the next inner strengthening layer of the conch was
deposited. Since the spines were apparently not open along their
leading edge and were probably soon sealed off at their base, it
seems unlikely that they functioned as supports for soft parts at
the aperture edge or as channels for current flow.

Teichert (1967, p. 204) included spines on ammonites in his
tabulation of adaptive devices to regulate buoyancy. Kennedy
and Cobban (1976, p. 28) suggested that spines might serve as
balancing aids when the shell rested on the bottom or as hori-
zontal stabilizers to prevent yawing. Westermann (1971, p 7)
has listed as functions of ornament on ammonites: strengthening
the relatively thin shell against implosion, protection against
impact, protection against smaller predators, decrease of drag
under conditions of turbulent flow, and camouflage. Arkell,
Kummel, and Wright (in Moore 1957, p. L122) suggested that
spines could have served as balancers or stabilizers or as a protec-
tion against enemies. It is this latter function that Westermann
inferred lateral and ventrolateral spines to have served. Cephalo-

Figure 4. Reconstruction of ventral view of young A. peninsularis .
Ammonite outline based on CAS 4250 and spines similar in length to
those on UCLA 38765 added. Spines produce an obviously unstreamlined
outline and effectively increase width of ammonite with respect to height.

pods of all kinds are noted for their varied camouflage techniques
(Lane 1960, p. 93-104; Cowen et al. 1973; Young and Roper
1976), and Cowen et al. (1973) suggest that the primary func-
tion of ammonite ornament was camouflage for a nekto-
benthonic organism.

Ornament to decrease drag is more likely to be ribbing or nod-
ing than long spines (Chamberlain and Westermann 1976,
p. 329). Such long frail spines were probably not secreted to
strengthen the shell. They could have offered some protection
against impact, absorbing some of the impact energy or perhaps
more importantly putting distance between the primary impact

FIGURES 5-6. Anapachydiscus peninsularis (Anderson and Hanna).
UCLA hypotype cat. no. 38765, x 3/4. (5) lateral view with three
rounded holes and “tom-on-perforation' ' edge to broken body chamber.
Twelve hollow spines show in umbilicus and are followed adaperaturally
by four flat-topped, rimmed tubercles. (6) opposite side of figure 5 with
two rounded holes and on opposite side of umbilicus one triangular hole
interpreted as being made by tip of tooth. Six flat-topped, rimmed tuber-
cles bordering the umbilicus considered to be bases of hollow spines.

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 304:1-8.

Saul: Ammonite (Anapachydiscus)

and the conch, but if this were their main function a more useful
placement would seem to be nearer the unprotected venter, espe-
cially if the ammonite was nekto-benthonic. Nor do they seem
well placed as defensive armaments. Gastropods arm their aper-
ture and anterior siphon; the bivalve Hysteroconcha has spines on
its corcelet which thus surround the siphons; but the spines on
young Anapachydiscus neither surround soft parts nor are they on
the outer perimeter of the animal to discourage attack. They are
thinner walled and even though liquid filled were apparently
frailer structures than the probably defensive spines of the gas-
tropods and bivalves. They would have increased the apparent
size of the young ammonite and so might have discouraged some
smaller predator. The need for camouflage (Cowen et al. 1973,
p. 211) was in part inferred from the paucity of ammonite speci-
mens showing pigmentation, but some few specimens have been
found (e.g., Arkell in Moore 1957, p. L92; Reyment 1958) and
some indicate the presence of Nautilus- like counter shading.

Surrogate ammonites were formed of sculpturing and investing
waxes. The investing wax was heavier, the sculpturing wax
lighter than water. As it has been suggested that ammonite and
shell were of near neutral buoyancy (Trueman 1941; Raup and
Chamberlain 1967), waxes were melted together empirically until
disks of approximately neutral buoyancy were obtained. These
disks were of similar diameter and width to the small paratype
CAS cat. no. 4250. Being homogeneous, they lacked the orient-
ing separated centers of gravity and buoyancy of actual ammo-
nites (Trueman 1941, figs. 14 and 15). Such a wax disk, lighter
than water, comes to rest floating with either lateral side up and
the venter directed horizontally. Disks of near neutral buoyancy
also usually settled toward a similar position, and a heavier disk
sank and usually lay flat on the bottom. The addition of lateral
spines gave the disks vertical stability, and they floated or sank
with the venter directed vertically. For the light disk the spines
had to be quite long to change its orientation in the water, at least
twice as long as those on A. peninsularis . Spines of similar
length to those on A. peninsularis gave the neutrally buoyant and
heavier disks vertical stability and greatly slowed the descent of
the heavier disk. Vertical stability could also be achieved by
carving out rough umbilici and adding the wax to the ventro-
lateral margin to produce a shape similar to young Tropites sub-
bullatus (Hauer). The greater breadth of young A. peninsularis
relative to adult (Table 1) may have also tended to increase ver-
tical stability in the young although the breadth of young A.
peninsularis is not great enough to produce vertical stability in
the absence of spines. Spines are not considered to have provided
all vertical stabilization. Well separated centers of buoyancy and
gravity would also lend vertical stability in addition to apertural
stability, but the somewhat depressed form and probably longer
body chamber (Raup 1967, fig. 16) of young A. peninsularis
would have caused the centers of buoyancy and gravity to be
closer together (Trueman 1941, p. 371). The spines would have
provided added stability. They were of appropriate length to pro-
vide a neutrally buoyant ammonite similar to the small paratype
with vertical stability.

A pelagic youth has generally been assumed for ammonoids
(Arkell, Kummel, and Wright, in Moore 1957, p. L100; Ken-
nedy and Cobban 1976, p. 34). Cowen et al. (1973, p. 211)
suggest that the sculpture of ribbed ammonites indicates that they
were necto-benthonic inhabitants of the photic zone, their sculp-
ture serving to hide them from the predator above, and their close
association with the bottom insuring that there was seldom a

predator below. That ammonites lived in the photic zone is also
suggested by the discovery that unlike the nocturnal Nautilus and
some deep water octopods, at least some ammonites had an ink-
sack (Lehmann 1971a, p. 1262). As the spines on young A.
peninsularis and A. cf. A. arrialoorensis would not have im-
proved the streamlining of a shell which was already poorly
streamlined (Kummel and Lloyd 1955; Chamberlain 1976),
swimming must have been at a rather deliberate pace, the
entrapment of food effected without much darting about. The
holotype of A. peninsularis is over 35 cm in diameter and much
less strongly ornamented on its outer whorls and similarly less
ornamented than specimens of less than 9 cm diameter. This
reduction in relief of ribbing can be considered to accord with
findings of Chamberlain and Westermann (1976, p. 328) that
ribbing can serve to reduce drag and conserve propulsive power
on smaller ammonites, but that large ones should be smooth.
Young A. peninsularis, however, additionally had spines, and it
does not merely become smoother as it approaches 10 cm in
diameter; it also becomes higher relative to width and reduces the
relative size of its umbilicus, thus improving streamlining slight-
ly

This complex of morphologic changes suggests that spiny
young Anapachydiscus peninsularis may have been virtually
planktonic and feeding on plankton. Ammonites have usually
been considered to share the macrophagous diet of other living
cephalopods, but a microphagous plankton-feeding mode of life
has been suggested for some (Morton and Yonge in Wilbur and
Yonge 1964, p. 49; Busnardo 1965, p. 110; Kennedy and Cob-
ban 1976, p. 35), and Lehmann (1971b, p. 339) reports the pre-
served crops of ammonites containing the remains of prey too

Figure 7. Hypothetical ammonite with eight webbed cirrated arms.
Webbed arms suggested by distribution of epizoans of Seilacher (I960)
and Meischner (1968). In both cases the epizoans were able to settle on
the living ammonite shell posterior to the aperture and on the body cham-
ber especially on the venter. They did not settle on the conch above the
aperture. If the ammonite had tentacles as does Nautilus this selective
infestation would not be expected, and long arms as in Argonauta would
be able to sweep the whole conch clean. But webbed arms should be able
to remove any larvae from the conch above the aperture while being
restrained from reaching the lower and rearward portions of the conch
Webbed arms could be used to trap food or also for propulsion as in
octopods (Lane 1960, p. 69).

Contrib. Sci. Natur. Hist. Mus. Los Angeles County .1979. 304:1-8.

Saul: Ammonite (Anapachydiscus)

small tor the application of jaws. Studies of the jaws themselves
(Kaiser and Lehmann 1971) indicate that the jaws of ammonites
were not capable of actual biting. Reconstruction of the living
ammonite is often based on analogy with Nautilus (e.g., Arkell
in Moore 1957. p. L82, fig. 124); but Nautilus may be a poor
model for ammonites (Mutvei and Reyment 1973; Kennedy and
Cobban 1976, p. 2). Lehmann (1971a and b) has shown that the
radula of ammonites is more like that of octopods and Vampyro-
morpha; and Flower (1955, p. 866) has argued for fewer arms,
than the 60+ tentacles possessed by Nautilus, in groups consid-
ered ancestral to ammonites (Sweet and Moore in Moore 1964,
p. K 101 , fig. 70). Mutvei and Reyment ( 1973, p. 632) propose a
long bodied ammonite without funnel. But their proposed long
ctenidia in narrow mantle cavity would require maintenance of a
current of water to ward off asphixiation, and as the funnel is
primarily a means of ejecting from the mantle cavity water which
has passed over the ctenidia, the funnel is here retained to aid
respiration. Figure 7 equips young A. peninsular is with eight
webbed cirri-bearing arms. The position of the arms back over
the upper part of the shell was suggested by the distribution of
epibionts described by Seilacher (1960) and Meischner (1968).
Webbed arms with cirri (as suggested by Kaiser and Lehmann
1971, p. 31) could have been used by young planktonic A.
peninsularis to gather in a planktonic feast as its outrigger spines
steadied it in the water.

The abandonment of spines, reduction of sculpture, increase of
height and decrease of umbilical size may mark the change to
greater swimming ability and more actively pelagic life. The
thinness of the shell suggests that this did not lead to the explora-
tion of great depths.

ACKNOWLEDGEMENTS

The other members of our safari, W.P. Popenoe, R.B. Saul,
and W.C. Cornell, made the collecting trip possible. Robert
Springfels donated sculpturing and investment waxes. Peter U.
Rodda granted permission to study the type specimens of Para-
pachydiscus peninsularis Anderson and Flanna at the California
Academy of Sciences. John Alderson became interested in the
spines and loaned a specimen of Anapachydiscus cf. A. arria-
loorensis (Stoliczka) for comparison. The manuscript has been
improved by the suggestion and needed criticism of R.B. Saul,
W.P. Popenoe, G.B. Cleveland, M.A. Murphy, and R. Cowen.
Final drafting of figures 1-4 was by Victoria Doyle Jones. To all
of the above I am grateful.

RESUMEN

Un especimen de la temprana epoca Maestrichtian Anapachy-
discus peninsularis (Anderson y Hanna) de Arroyo Santa Cata-
rina, Baja California, Mexico, tiene cavidades casi redondas
taladradas en la concha arregladas de tal ntanera que sugieren
muestras de las mordeduras del mosasaur. Las tempranas es-
pirales estan adornadas con huecas espinas laterales, las cuales
podrian haber servido de estabilizadores verticales.

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Saul: Ammonite (Anapachydiscus)

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Raup. D.M. 1967. Geometric analysis of shell coiling: coiling in
ammonoids. J. Paleontol. 41:43-65, 19 figs.

Raup, D M. and J.A. Chamberlain, jr 1967. Equations for
volume and center of gravity in ammonoid shells. J. Paleon-
tol. 41:566-574, 3 figs.

Reyment, R.A. 1957. Uber Farbspuren bei einigen Ammoniten.
N. Jahrb. Geol. Palaont., Monat.:343-351, 3 figs.

Seilacher, A. 1960. Epizoans as a key to ammonoid ecology. J.
Paleontol. 34:189- 193, 3 figs.

Stoliczka, F. 1863-66. Ammonitidae, with revision of the
Nautilidae, etc. In Blanford, M.F. and F. Stoliczka,
1861- 1866. The fossil cephalopods of the Cretaceous rocks
of southern India. India Geol. Surv. Mem.,Palaeont. Indica,
Ser. 3, 216 pp., 95 pis. [41-56, pi. 26-3 1 , 1863; 57- 106,
pis. 32-54, 1864; 107-154, pi. 55-80, pi. 66a, 1865;
155-216, pis. 81-94, 1866],

Teichert, C. 1967. Major features of cephalopod evolution. In
Teichert. C. and E. Yochelson [eds.j. Essays in paleon-
tology and stratigraphy. Univ. Kansas Press; Lawrence,
Kansas:pp. 162-210, 20 figs., 1 table.

Trueman, A.E. 1941. The ammonite body-chamber, with spe-
cial reference to the buoyancy and mode of life of the living
ammonite. Geol. Soc. London, Quart. Jour. 96:339-383,
17 figs.

Westermann, G.E.G. 1971. Form, structure and function of
shell and siphuncle in coiled Mesozoic ammonoids. Royal
Ontario Mus., Life Sci. Contrib. 78:39 pp., 17 figs.

Wilbur, K.M. and C.M. Yonge [eds.j. 1964. Physiology of
Mollusca. Academic Press, New York and London, 1:473
pp. , 11:645 pp.

Young, R.E. andC.F.E. Roper 1976. Bioluminescent counter-
shading in midwater animals: evidence from living squid.
Science, 191:1046-1048.

Accepted for publication July 5, 1977.

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 304:1-8.

Number 305
March 16, 1979

NATURAL HISTORY MUSEUM OF LOS ANGELES COUNT

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CHELODESMID STUDIES. VIII.

A NEW MILLIPED OF THE GENUS TRICHOMORPHA FROM COCOS

THE NEW TRIBE TRICHOMORPHINI (POLYDESMIJ&A: CHELODESMIDAE)

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Number 305
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CONTRIBUTIONS IN SCIENCE

NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY

CHELODESMID STUDIES. VIII.

A NEW MILLIPED OF THE GENUS TRICHOMORPHA FROM COCOS
ISLAND, WITH NOTES ON RELATED SPECIES AND THE PROPOSAL
OF THE NEW TRIBE TRICHOMORPHINI (POLYDESMIDA: CHELODESMIDAE)

By Richard L. Hoffman

Published by the NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY • 900 EXPOSITION BOULEVARD • LOS ANGELES, CALIFORNIA 90007

ISSN: 0459-8113

Suggested Citation; Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 305:1-7.

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 305:1-7.

CHELODESMID STUDIES. VIII.

A NEW MILLIPED OF THE GENUS TRICHOMORPHA FROM COCOS
ISLAND, WITH NOTES ON RELATED SPECIES AND THE PROPOSAL
OF THE NEW TRIBE TRICHOMORPHINI (POLYDESMIDA: CHELODESMIDAE)1

By Richard L. Hoffman2

Abstract; A new species of chelodesmid milliped, Trichomorpha hyla, is described from Cocos Island,
Costa Rica. It differs from T. folium (Brolemann), from the same island in reduced tergal setation, absence of
lateral paranotal dentation, lack of middorsal pink spots, and in a less complex gonopod structure. The new
tribe Trichomorphini is proposed to accommodate the genera Trichomorpha, Phylactophallus, Ancholeptodes-
mus, Belonodesmus, Allarithmus, Talamancia, and the new genus Loomisiola, based on Trichomorpha crini-
tapes Loomis, 1972, from Costa Rica. The new name Trichomorpha crucicola is proposed to replace T.
gracilis Loomis, 1974, preoccupied by T. gracilis Carl, 1914.

Dr. C. L. Hogue, Senior Curator in Entomology in the Natural
History Museum of Los Angeles County, recently placed in my
hands a small collection of diplopods picked up by him on Cocos
Island. Most of the specimens are female pyrgodesmoids and not
definitely identifiable to genus or species, but there is one adult
male of an undescribed species in the chelodesmoid genus Tri-
chomorpha, which is being described here in order to make the
name available for use in Dr. Hogue’s studies. The occasion is
taken to append some additional information on the nomenclature
and taxonomy of the genus and related forms.

Trichomorpha, endemic to the region between Ecuador and
Costa Rica, is apparently now in the expanding phase of its phy-
logeny, as the known species are very numerous and yet basically
similar in terms of general gonopod structure; specific differentia-
tion is most pronounced in various external non-sexual character-
istics. Synrpatry and apparent syntopy seem to be common. In
most features Trichomorpha appears to be a rather specialized
genus of chelodesmoids, apparently related to members of the
tribe Batodesnnni, which has a very similar geographic distribu-
tion. Along with several other genera of the northern Andean-
Pananranian region, Trichomorpha can be segregated into another
distinct tribe which is proposed later in this paper.

I wish to express here my thanks to Dr. Hogue for providing
the occasion for preparation of the following information about
this group of interesting Neotropical myriapods.

FAMILY CHELODESMIDAE

Genus Trichomorpha Silvestri

Trichomorpha Silvestri 1898, Boll Mus. Torino, vol. 12, no. 305,
p . 11. Type species; T. elegans Silvestri 1898 (Ecuador), by mono-
typy.

Eihophallus Chamberlin 1933, Pan-Pacif. Entom., vol. 9, p 20. Type
species, E. cervantes Chamberlin 1933 (Costa Rica), by original
designation.

Typophallus Chamberlin 1940, Bull. Univ. Utah (biol. ser.), vol. 5, no.
6, p. 10. Type species, T. evidens Chamberlin 1940 (Panama) by
original designation.

Desmaehrides Chamberlin 1940, op. cit . , p . 11 Type species, D
dichrus Chamberlin 1940 (Panama), by original designation.

DIAGNOSIS: A trichomorphine genus in which the gonopod
prefemur is short to moderate in length and the femur greatly
reduced or absent, the solenomerite appearing to originate at the
end of the prefemur; postfemoral elements simple to complex in
form, but usually partially enclosing solenomerite. Coxa with
prominent dorsal apophysis. Anterior legs of males with large
distal tibial pad, subtending the tarsus; in some species the coxa
and femur of the 7th pair of legs are lobed or otherwise modified.
Anterior sterna with or without paramedian processes or lobes.
Prefemora of some or all legs usually with apical spine in both
sexes. Antennae long and slender, extending back at least to
paranota of 3rd segment.

NOTES: About 40 specific names have been based upon
specimens referable to this genus, as well as a number of unjusti-
fiable generic names. Loomis (1964, 1972) has shown some of
these names to be synonyms and so reduced the total somewhat,
but a number of undescribed species at hand, and the small frac-
tion of the generic range that has been sampled thus far, suggest
that eventually as many as 200 species of Trichomorpha may be
discovered.

In 1972 Loomis merged the name Eihophallus with Tricho-
morpha, but in a later paper (1974) kept the two separate on the

‘Review Committee for this Contribution;

Charles L. Hogue
W. A. Shear
Rowland Shelley

“Professor of Biology, Radford College, Radford, Virginia 24142.

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 305:1-7.

Hoffman: Isla del Coco Milliped

basis of a supposed difference in proportions of several podo-
meres. Having the opportunity to study type material in the col-
lection of the late R. V. Chamberlin, I can verify that Ethophal-
lus cervanrcs is in fact only a species of Trichomorpha and the
‘difference’ in lengths of the tibia and tarsus entirely illusory. At
the same time I can confirm Loomis’s disposal of the two names
Typophallus and Desmachrides as strict junior synonyms of Tri-
chomorpha.

The name Trichomorpha gracilis, proposed by Loomis in 1974
for a Costa Rican species, is preoccupied by Trichomorpha
gracilis Carl 1914, from Colombia. On learning of my intention
to describe the Cocos Island species, Mr. Loomis asked me to
use the occasion to provide a new name for his gracilis. 1 here-
with propose the new name Trichomorpha crucicola, in refer-
ence to the type locality (Finca las Cruces, near San Vito, Costa
Rica).

Trichomorpha hyla new species

Figures 1-8

Holotype. — Adult male (LACM) collected at Wafer Bay, Cocos Is-
land, Costa Rica, 18 April 1975, by Charles L. Hogue.

DIAGNOSIS: A moderate-sized member of the genus character-
ized by the smooth metaterga, each with only two setae; presence
of tibial pads on legs 2-6 of males; denticulate posterior edge of'
most paranota; occurrence of paramedian processes on the sterna
between legs of the 2nd, 3rd, and 5th pairs, and of a prominent basal
lobe on the femora of the 7th pair; and by the relative simplicity of
gonopod structure, the acropodite set off by a distinct suture and
virtually without any lobes or processes.

DESCRIPTION: Body ca. 17.0 mm in length, 2.1 mm wide
over most body segments, W/L ratio about 12.4%. Body gener-
ally light brown. Metaterga pale testaceous brown, fading to
nearly colorless on the paranota, dorsal surface of prozona darker
brown, imparting a somewhat annulate appearance with low
magnification. Labrum and clypeus nearly colorless, basal anten-
nomere very pale brown, 2nd-6th antennomeres fuscous, 7th
nearly white. Podosterna and midventrum of prozona whitish,
basal four podomeres also nearly white, tibia light brown, tarsus
dark brown except for apical fourth which is white.

Head of normal appearance for the genus, front with sparse,
prominent, setiferous punctures; epicranial suture fine but distinct
down to level of antennae; genae convex, lateral margin narrow
but well-defined, with five or six marginal setae. Ventral edge of
labrum marginate except proximad to median teeth; 8-8 labral
setae. Interantennal space broad, wider than length of 1st anten-
nomere. Antennae long and slender, articles 5 and 6 with well-
defined, oval, white, sensory field on outer distal surface. Sen-
sory cones small, indistinctly grouped into two diads.

Collum about as wide as head, surface convex, smooth and
polished; lateral ends acute, forming about a 70° angle, lateral
edges set off up to level of mandibular articulation. 8 setae along
anterior margin, and 4 in a transverse row approximately at mid-
length.

Body segments not deeply telescoped, paranota thus well-sep-
arated, subsegments not strongly separated, stricture shallow, its
anterior edge sharply defined. Paranota well developed, set high
on body and nearly horizontal, middorsum somewhat convex,
surface of metaterga smooth and polished, with two paramedian
setae; transverse sulcus evident on segments 5- 17, dorsal surface
of paranota slightly convex basally. Posterior comers of all para-

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 305:1-7.

nota acute, becoming increasingly prolonged and subspiniform
back to about segment 17, posterior edge of most paranota dis-
tinctly denticulate, lateral edge with prominent offset at mid-
length on segments 2-7, thereafter nearly smooth and straight
(Figs. 1, 2); ozopores opening dorsolaterad, peritremata moder-
ately developed, elongate, not set off from remainder of para-
notal edge; anterior corners of paranota with a prominent denticu-
lation. Paranota of segment 19 small, curved mesad.

Epiproct of normal shape, but lateral and apical dorsal tuber-
cules a little more prominent than usual (Fig. 3). Paraprocts and
hypoproct without peculiarities.

Sterna of midbody segments broad (about equal to combined
length of coxa and prefemur), slightly elevated above prozona,
smooth and glabrous, with indistinct transverse groove, no sub-
coxal spines or lobes. Stigmata minute, without raised edges.
Side of metazona smooth. Pleurosternal carinae visible only on
first two or three segments. Prefemora with prominent apical
spine from 2nd to 18th pair of legs. Tibial pads present on legs of
2nd- 6th pairs, much the largest anteriorly where about 2/3rds
length of tarsus. Tarsal claw small and straight.

Gonopores opening flush with surface of coxae of 2nd pair of
legs. Sternum between 3rd pair of legs with two small para-
median lappets, that between 4th pair of legs with very large,
compressed, divergent, apically truncated processes (Fig. 4), that
between 6th pair of legs with two small contiguous acutely coni-
cal projections (Fig. 5). Coxae of 6th and 7th legs produced ven-
trad, the projection especially prominent on 7th legs, the femora
of which are provided with a large proximal lobe or process on
the ventral side (Fig. 5).

Gonopod aperture transverse, lateral and posterior edges pro-
duced ventrad, anterior edge unmodified; gonopods projecting
forward over sternum of 6th segment, of the form shown in fig-
ures 6-8. Coxae without median sternal remnant (Fig. 6), the
dorsal apophysis robust, with two setae; telopodite attached at a
right angle to coxa, nearly straight, setose prefemoral region
about one-third total length of telopodite, femoral region strongly
reduced and indicated only by the point of origin of the solenom-
erite; latter slender, nearly straight, its entire length visible in
mesal aspect (Fig. 8); postfemoral region a simple, spatulate,
apically concave process with a small thin lamella on the dorsal
side. Lateral side with a prominent postfemoral cingulum (Fig. 7,
C).

NOTES: Dr. Hogue supplied the following information on the
habitat of this species (in litt. . 19 September 1975): “All of this
material definitely came from bromeliads, although the collect-
ing occurred fairly close to ground level, approximately 4-6 feet.
The specimens were taken from very large plants growing on
downed logs and branches of Hibiscus tiliaceus, a mangrove
species. The ground beneath the site from which the millipeds
came was Hooded by high tide twice daily.’’

The relationships of this species are not easy to establish.
There are now about 35 presumably valid species of Tricho-
morpha, 22 of them described from Colombia, 5 from Panama,
and 6 from Costa Ricas. Attems (1938) gave a key to 20 South
American species, but was unable to include most of the forms
named by Chamberlin because of inadequate descriptions. An
additional member of this genus (T. folium Brolemann 1903) was
misplaced by Attems in Camptomorpha, where it has no affini-
ties whatever.

T. hyla will not trace through Attems' 1938 key at all, being
excluded by both of the choices in the first couplet (metaterga

Hoffman: Isla del Coco Milliped

with numerous irregular setae as opposed to setae in two or three
transverse rows). In this species each metatergum has only a sin-
gle row composed of two setae, placed paramedially behind the
transverse sulcus (increasing to four on the last few segments).
On the basis of “best fit’’ we select the second choice, T.
venusta Carl 1914, from Colombia, but hyla differs from this
form in a number of ways including a different gonopod struc-
ture.

Of the four Panamanian species keyed by Loomis ( 1964), hyla
runs out to T. nidicola Chamberlin, but differs in lacking a pale
middorsal line and lateral paranotal dentations (four to six prom-
inent teeth in nidicola).

Six Costa Rican species are keyed by Loomis (1972), of them
hyla comes closest to T. folium, coincidentally the Trichomorpha
already known from Cocos Island.

In light of present knowledge of the genus, it seems impossible
to confidently relate T. hyla to any mainland species, and for the
present it may be justifiable to consider it a valid species endemic
to Cocos Island. In this respect then, comparison needs be made
primarily with T. folium (Brolemann).

This species was supported by a detailed description and good
drawings, and the following points of difference may be noted:
folium is slightly longer and more robust than hyla. its W/L ratio
14% as opposed to 12%; the nretaterga in folium are reddish-
brown with a series of middorsal red spots and the legs are uni-
formly yellowish; although Brolemann did not allude to tergal
setation generally, he stated that the collum has three transverse
series of hairs and generally the arrangement of the collum per-
sists on the metaterga as well. In folium the paranota are laterally
dentate (nearly smooth in hyla)-, the femur of the 7th pair of legs
lacks the prominent basal lobe found in hyla; and the gonopod
telopodite is provided on its dorsolateral surface with three tri-
angular projections not present in hyla. There is no doubt what-
ever that they are quite different, despite some points of similar-
ity such as lobation of the anterior sterna and general gonopod
pattern .

Loomis (1972:192) suggested that perhaps T. evidens (Cham-
berlin) from the Canal Zone is a junior synonym of folium. 1 have
been able to study the type material of evidens and find it to be
quite different from folium; among other things evidens com-
pletely lacks processes on the anterior sterna and it is moreover
the least setose of known Trichomorpha species. I was unable to
locate either setae or setal sockets on any of the terga.

The present opportunity is taken to implement a long-standing
intention, to formally recognize a suprageneric taxon to include
Trichomorpha and related genera within the subfamily Chelo-
desminae:

Trichomorphini new tribe

DIAGNOSIS: Small (9-24 mm) chelodesmoids in which the
gonosternum is reduced or lost; gonopod aperture transversely
narrow with flared edges; gonopod coxae with prominent dorsal
apophyses; femoral region of gonopod very short and merged
with end of prefemur, solenomerite appearing to originate from
prefemur; without prefemoral process; solonomerite long and
slender, usually partly or entirely enveloped within the post-
femoral region, latter straight or strongly curved dorsad.

Body parallel-sided, none of anterior segments broadened or
narrowed, relatively slender; segments not deeply telescoped into

each other. Antennae long and slender, with sensory fields on
outer side of articles 5-7. Mandibles relatively large. Collum
hemispherical and convex. Paranota of moderate size and usually
set high on sides; at least some metaterga with transverse sulcus
(usually segments 5- 17), and most species with at least two or
three transverse rows of setae; pore formula normal, pores open-
ing dorsolaterad in elongated peritremata, latter continuous with
edge of paranota. Epiproct usually with enlarged lateral and sub-
terminal tubercules. Sterna relatively broad and flat, at most pro-
duced into low subcoxal cones. Anterior legs of males usually
with tibial pads subtending tarsi, often coxae and prefemora vari-
ously lobed or otherwise modified.

COMPONENTS: Trichomorpha Silvestri 1897; Ancholepto-
desmus Brolemann 1919; Belonodesmus Chamberlin 1918;
Allarithmus Attems 1933; Phylactophallus Pocock 1909; Tala-
mancia Loomis 1974; Loomisiola new genus.

DISTRIBUTION: Northern Cordilleran region, from Ecuador
to Costa Rica, east as far as Trinidad.

REMARKS: All of the generic names listed above are subject
to major change when sufficient material is at hand for a revision
of the group. Some may prove to be junior synonyms and Tri-
chomorpha itself may be divided into two or more smaller gen-
era. Loomis (1964, 1972) reduced the Chamberlinian names
Typophallus, Desmachrides. and Ethophallus to the synonymy of
Trichomorpha. and it seems quite possible, on the basis of pub-
lished information, that Ancholeptodesmus and Allarithmus may
suffer a similar fate.

I know the genus Phylactophallus only from Pocock's original
description, but it seems to be based on a trichomorphine species,
and to be a distinct genus. I think it is very likely that Isidrona
forficula Attems ( 1933) is a generic and probably also a specific
synonym of P stenomerus Pocock. In the “Checklist” of Cen-
tral America millipeds, Loomis (1968:15) — on the basis of a
suggestion made by me in I in. many years ago — combined the
names Phylactophallus and Allarithmus. I now suspect that my
opinion was based upon a confusion of two names (Allarithmus
instead of Isidrona). in any case it was egregiously incorrect.

Most of the species referred to Trichomorpha and its satellite
genera have the gonopods formed on a basically similar pattern.
One Costa Rican species, however, is so divergent that some
kind of recognition in a separate status seems desirable.

Loomisiola new genus

Type species. — Trichomorpha crinitapes Loomis 1972.

DIAGNOSIS: A trichomorphine genus differing peripherally
from Trichomorpha in having shorter and stouter antennae, and
paranota with less prominently produced posterior corners. The
gonopod prefemur is unusually long, distally broadened, and
provided apical I y on the lateral side with an oblique lobe set with
numerous long slender setae; postfemoral elements of gonopod
strongly reduced.

DISTRIBUTION: Loomisiola crinitapes is known so far only
from the original type material, taken at Cairo, Province Linton,
Costa Rica.

ETYMOLOGY: It is appropriate that this taxon commemorate
the name of H. F. Loomis, who has contributed most to our
knowledge of Central American and Antillean Diplopoda, in
numerous papers spanning four decades.

Contrib. Sci. Natur. Hist. Mas. Los Angeles County. 1979. 305:1-7

Figures 1-8. Trichomorpha hyla, structural details, from holotype. Figure 1, left paranota of segments 8 and 9, dorsal aspect. Figure 2, left paranota of
segments 13 and 14, dorsal aspect. Figure 3, posterior end of body, dorsal aspect. Figure 4, sternum and bases of right legs of segment 5, aboral aspect.
Figure 5, sternum and right legs of segment 6, aboral aspect (setation omitted from anterior leg). Figure 6, coxa and base of telopodite of right gonopod,
dorsal aspect. Figure 7, left gonopod, lateral aspect (c, cingulum). Figure 8, left gonopod mesal aspect. Figures 1-5 drawn X45, 6-8 X 90.

Contrib. Sci. Natur. Hist. Mus. Los Angeles Counts . 1979. 305:1-7.

Hoffman: Isla del Coco Milliped

RESUMEN

Se describe una especie de la familia Chelodesmidae, Tricho-
morpha hyla, que se encuentra en la Isla del Coco de Costa Rica.
Se diferencia de T. folium (Brolemann 1903) de esa misma isla
en la setacion dorsal reducida, en la ausencia de dentacion de los
paranota, y en una estructura gonopoda menos compleja. Le
nueva tribu Trichomorphini se propone para acomodar los
generos Trichomorpha, Phylactophallus, Ancholeptodesmus,
Belonodesmus , Allarithmus, Talamancia, y el nuevo genero
Loomisiola, basado en Trichomorpha crinitapes Loomis 1972, de
Costa Rica.

LITERATURE CITED

Attems, C. 1933. Neue Polydesmiden von Costa-Rica. Ann.
Naturh. Mus. Wien 46:257-269, figs. 1-20.

. 1938. Fam. Leptodesmidae, Platyrhacidae, Oxydes-

midae, Gomphodesmidae, in: Das Tierreich 69:1-487,
figs. 1-509.

Brolemann, H.W. 1903. Myriapodes recueillis a 1 ’Isla de Cocos
par M. le Professeur P. Biolley. Ann. soc. Ent. France
72:128- 142, text figs. 1-10, pi. I, figs. 1-18.

. 1919. Myriapodes, in: Miss. Arc Merid. equat.

Amer. Sud 10 (2):235— 275 , figs. 1-42.

Carl, J. 1914. Die Diplopoden von Colombien nebst Beitragen
zur Morphologie der Stemmatoiuliden. Mem. soc. Neu-
chatel. sci. natur. 5:821-993, figs. 1-262.

Chamberlin, R.V. 1918. The Chilopoda and Diplopoda of the
West Indies. Bull. Mus. Comp. Zool. 62:151-262.

. 1940. On a collection of centipeds and millipeds

from Barro Colorado Island, Panama. Bull. Univ. Utah,
biol. ser., 5(6):3 — 16.

Loomis, H.F. 1964. The Millipeds of Panama. Fieldiana: Zool.
47(1):1-136, figs. 1-13.

. 1968. A Checklist of the Millipeds of Mexico and

Central America. Bull. U. S. Nat. Mus. 266:1- 137.

. 1972. Millipeds from the Atlantic Lowlands of Costa

Rica. Florida Entom. 55:185-206, figs. 1-42.

. 1974. Millipeds from the southern Costa Rican high-
lands. Florida Entom. 57:169-187, figs. 1-29.

POCOCK, R.I. 1903- 1910. Chilopoda and Diplopoda, in: Biologia
Centrali- Americana, pp. 1-217, pis. 1-15 [fascicle con-
taining Phylactophallus issued in December, 1909].

Accepted for publication July 8, 1977.

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 305:1-7.

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Prior to November 30, 1973, publications of the Natural History Museum have appeared under various titles — Leaflet
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Edward Ostermeyer
Editor

Printed in the United States of America by International Printing Services, Inc.

Number 306
March 16, 1979

ONTRIBUTIONS IN SCIENCE

NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY

SYSTEMATICS OF THE SPECIES OF CRY PTORHOP ALUM
(COLEOPTERA: DERMESTIDAE) OCCURRING IN CALIFORNIA

By R.S. Beal, Jr.

Published by the NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY • 900 EXPOSITION BOULEVARD • LOS ANGELES, CALIFORNIA 90007

CONTENTS

Abstract 3

Systematics 3

Abbreviations 4

Generic Nomenclature and Description

Cryptorhopalum Guerin- Meneville 4

Key to Adults from California 6

Descriptions and Discussions of California Species

Cryptorhopalum triste LeConte 7

Cryptorhopalum rubidum new species 10

Cryptorhopalum uteanum Casey 12

Cryptorhopalum apicale (Mannerheim) 15

Cryptorhopalum fusculum LeConte 17

Cryptorhopalum filitarse Casey 18

Cryptorhopalum balteatum LeConte 19

Cryptorhopalum haplotes new species 20

Synonymies and Lectotype Designations

for C. reversum and C. pruddeni 20

Acknowledgments 21

Literature Cited 21

ISSN; 0459-8113

Suggested Citation; Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 306:1-22.

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 306:1-22.

SYSTEMATICS OF THE SPECIES OF CRYPTORHOPALUM
(COLEOPTERA: DERMESTIDAE) OCCURRING IN CALIFORNIA1

By R.S. Beal, Jr.2

ABSTRACT: The following described species are recognized as occurring in California: C. triste LeConte ( =
C. modestum Casey, C . fusciclave Casey, NEW SYNONYMS), C. uteanum Casey (= C. nephianum Casey,
C. aridum Casey, C. bakeri Casey, NEW SYNONYMS), C. upicale (Mannerheim) (= C. coloradense Casey,
C. tuckeri Casey, C.fontinale Casey, C. grisescens Casey, NEW SYNONYMS), C. fusculum LeConte (= C.
anthrax Casey, C. granum Casey, C. pumilum Casey, C. piceum Casey, NEW SYNONYMS, and C. affine
Casey, REVISED SYNONYMY), C. filitarse Casey, and C. balteatum LeConte. Two new species are
described: C. rubidurn, for which Yuma, Arizona, is the type locality, and C. haplotes, known only from
Jacumba, California. The species can best be distinguished by characters found associated with the aedeagus
and lateral lobes and with the eighth morphological abdominal sternum of the males. Redescriptions of existing
species are provided together with lectotype designations wherever necessary. A key to the species is provided
for forms of adults occurring in California. Flower hosts, geographic distribution, and synonymies are given for
each species insofar as known. In addition to the species above, synonymies are given and lectotypes
designated for two species closely related to C. balteatum but occurring outside California: C. reversum Casey
(= C. festivum Casey, C. balteatum Casey [non balteatum LeConte], C. pallens Casey, NEW SYNONYMS),
and C. pruddeni Casey (= C. insigne Casey, C. anthrenoides Casey, NEW SYNONYMS).

Adults of the genus Cryptorhopalum are small, ovate beetles,
almost always taken on flowers, and readily recognized as der-
mestids by the presence of a median ocellus and a hind coxa
grooved for the reception of the femur. The two-segmented
antennal club, which fits closely into a fossa on the underside of
the prothorax, and the covering of hairs rather than of scales eas-
ily separates them from all other Nearctic dermestids.

Relatively large numbers of specimens of the genus are found
in the collections of most insect museums. Nevertheless, no revi-
sion of the genus has been undertaken since that of Thomas L.
Casey in 1900. Because Casey worked from very small numbers
of specimens and described his species from a typological rather
than a population concept, his “species” in many instances are
simple variants of previously described species or of another of
his species. Consequently his key to the species is of little value.
There is a clear need to investigate the species from a population
concept, to develop a workable key for distinguishing them, and
to provide accurate descriptions for them. This paper is an effort
to accomplish these goals for those species presently known to
occur in California.

Other than records of flowers on which the adults are found,
very Little is known of the biology of species of Cryptorhopalum.
Larvae of some unidentified species have been intercepted at
border inspection stations on cheese and other food products from
Mexico. Nevertheless, no Nearctic species has ever been known
to occur as a pest of stored food products. Larvae of a very rare
Arizona species, C. poorei, have been found associated with

spider nests under the bark of dead ponderosa pine trees (Beal
1975), yet an association with spider nests does not seem to be a
likely natural habitat for larvae of most of the species, judging
from their numbers and the situations in which adults are often
found. Hopefully, the present study will stimulate further investi-
gations into the biologies of the species.

SYSTEMATICS

Species Characters

Adults of some species are readily distinguished by patterns
formed on the elytra by light and dark colored setae or by the
shapes of the antennal club. The form of the anterior tibia is
diagnostic for a few species, as is the density of punctation on the
pronotum. Some species, however, are virtually indistinguishable
from each other on the basis of external characters. Nevertheless,
apparently reliable characters for the recognition of all California
species and for many other Nearctic species are found in struc-

‘Review Committee for this Contribution:

Charles L. Hogue
John M. Kingsolver
Floyd G. Werner

2Northern Arizona University, Flagstaff, Arizona, 86011.

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 306:1-22.

Beal: California Cryptorhopalum

tures associated with the male genital apparatus.

The male genital tube is formed by the terga and the sterna of
the 8th and 9th morphological abdominal segments, the weakly
sclerotized tegmen (or basal piece) that lies beneath the base of
the aedeagus, and the lateral lobes and aedeagus. Except in copu-
lation, these parts are telescoped together within the abdomen.
Discriminating characters are found in the shape of the aedeagus,
in the shape of the lateral lobes and the “bridge” that connects
the two lobes, and most particularly in the shape of the apical
margin of the sternum of abdominal segment 8 and the form of
the setae that are inserted on it.

Species Identification

Any effort to identify species on the basis of external charac-
ters alone is complicated by two problems. One is the relative
paucity of such characters. The other is the relatively large
degree of geographic variation that occurs over the range of some
of the species. It is usually fairly easy to separate members of
any two species occurring within a limited geographic area. Yet
convergence of external characters may make quite difficult sep-
aration of specimens of one species from those of another from a
distance of only a few hundred miles. In the key that follows, an
effort is made to distinguish only those forms of each species that
are found in California. However, the formal description for each
is given for all variations within its entire range, as far as known.

Species Groups

Based on the total number of known adult characters, the
Nearctic species fall into three rather distinct groups. The Cali-
fornia representatives of these groups are the following.

TRISTE group
C. triste LeConte
C. rubidum, new species
C. uteanum Casey
C. apicale (Mannerheim)

C. fusculum LeConte
QUADRIPUNCTATUM group
C. filitarse Casey
C. balteatum LeConte
HAPLOTES group
C. haplotes, new species

The quadripunctatum group is characterized by the presence of
an appendage-like structure on each lateral posterior angle of the
8th (morphological) abdominal sternum of the male (Figs. 14,
15). Externally it is characterized by the presence of three bands
of light-colored pubescence on the elytron, a feature not found in
members of either the triste or haplotes groups. As far as the
California species are concerned, the quadripunctatum group is
further separable by the ovate outline of the male antennal club.
In distinction to this, members of the haplotes and triste groups
have a subcylindrical antennal club in the males.

In members of the triste group the 8th (morphological) abdom-
inal sternum of the male lacks lateral appendage-like structures or
sublateral marginal papillae but has a posterior median process at
the apex of which are various specialized setae. The one excep-
tion is C. rubidum, which has a median cluster of setae but no
process. The dorsal setae in members of this group are commonly
unicolorous, but if bicolorous they do not form three more or less

distinct bands on the elytron. (C. apicale usually has bicolorous
setae and these often form an oblique subbasai band and a sub-
apical spot, but there is no submedian band.)

The haplotes group, for which there is but a single Nearctic
representative, is distinguished from both of the preceding groups
by its more elongate form (Fig. 1). The 8th (morphological)
sternum of the male lacks a median apical process and lateral
appendage-like structures, but possesses sublateral papillae on
which are inserted several moderately long setae (Fig. 12). The
margin of the sternum has a cluster of setae inserted at the mid-
line. The dorsal setae are unicolorous, at least in this and the one
Mexican species I have studied.

C. rubidum is the least specialized of all the species consid-
ered, at least as far as the male genital structures are concerned.
Conceivably it could be placed in a separate group or attached to
any of the three groups as its most primitive member. However,
here it is placed in the triste group because of its very close
similarity in external characters to C. uteanum.

ABBREVIATIONS

In the discussion of species below, museums frequently
referred to are abbreviated as follows: CAS — California Acad-
emy of Sciences, San Francisco; MCZ — Museum of Compara-
tive Zoology, Cambridge, Massachusetts; LACM — Natural His-
tory Museum of Los Angeles County, California; NMNH —
United States National Museum of Natural History, Washington,
D.C.

GENERIC NOMENCLATURE AND
DESCRIPTION

Cryptorhopalum Guerin-Meneville

Cryptorhopalum Guerin-Meneville 1834 (1838), p. 67. LeConte 1854,
pp. 106-107. Reitter 1880 (1881), p. 43. Jayne 1882, p. 364.
Casey 1900, pp. 145, 155. Sharp 1902, p. 652. Rees 1943, p. 11.
Beal 1959, p. 101 (in key to genera). Beal 1975, pp. 228-233
(larval description).

Cryptorrhopalum Arrow 1915, pp. 437, 442.

Type of the genus: Cryptorhopalum quadripunctatum Guerin-Meneville
1838.

DESCRIPTIONS

ADULT MALES: Body ovate, moderately convex, covered on
dorsal surfaces with moderately short, subrecumbent hairs; scales
absent although some setae may be slightly ensiform. Head
slightly inflexed with front inclined 10 to 30 degrees from the
vertical; median ocellus present. Antenna 1 1 -segmented; club
2- segmented, ovate or subcylindrical; segment 10 subequal in
length to segment 11 or longer than segment 11. Pronotum
carinate on lateral margins but without sublateral carinae or
impunctate areas; basal lobe not raised (on same plane as poste-
rior margins), projecting posteriad moderately to strongly. Scutel-
lum visible. Elytra across humeri slightly wider than width of
pronotum at base; epipleuron inflexed with surface in approxi-
mately same plane as metepistemum. Prosternum with anterior
margin very weakly or not at all deflexed, arcuately rounding to
sides with area in front of lateral margin of procoxa somewhat
longer than area in front of middle of procoxa; process moder-
ately broad, received in mesosternal sulcus, and attaining meso-
sternum. Hypomeron with fossa closely conforming to size and
shape of antennal club; fossa occupying most of hypomeron and
margined behind by thread-like carina; posterior margin of

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Beal: California Crypt orhopalum

Figures 1-7. Figure 1: Dorsal aspect of C. haplotes. Figure 2: Dorsal aspect of C. apicale. Figure 3: Underside of prothorax showing antennal club in
place within antennal fossa of male of C . haplotes. Figure 4: Same of male of C. balteatum. Figure 5: Same of female of C. haplotes (hyp = hypomeron).
Figure 6: Same of male of C. rubidum. Figure 7: Same of male of C. fusculum.

Contrib. Sci. Natur. Hist. Mas. Los Angeles County. 1979. 306:1-22.

Beal: California Cryptorhopalum

KEY TO ADULTS FROM CALIFORNIA

1. Subbasal, subapical, and apical bands and sometimes basal band of light-colored pubescence present on elytron; basal and subbasal

bands sometimes coalesce. Basal half of 1st visible abdominal sternum with 2 oblique striae extending from beneath each trochanter
(leg may need to be relaxed and moved to observe character). Male antennal club ovate (Fig. 4). Female with 2 circular foveae on
disc of 5th visible abdominal sternum (Fig. 33) 2

— Elytral pubescence unicolorous, or with subapical spot of light-colored pubescence with or without subbasal band of light-colored

pubescence, or covered with light-colored pubescence with few to many intermingled dark hairs, particularly at apical third. Basal
half of 1st visible abdominal sternum with 1 oblique stria extending from beneath each trochanter. Male antennal club more or less
elongate, subcylindrical (Figs. 3, 5-7). Female without 2 circular foveae on disc of 5th visible abdominal sternum 3

2. Light-colored pubescence of elytron a deep gold color (orichalceous). Integument of pronotum generally appreciably darker in color

than elytron at base. Sternum of 8th (morphological) abdominal segment without cluster of setae at middle of apical margin; lateral
process with stout, curved seta at apex (Fig. 15) balteatum

— Light-colored pubescence of elytron white with slight golden cast. Integument of pronotum and base of elytron usually identically

colored. Sternum of 8th (morphological) abdominal segment with cluster of setae at middle of apical margin; lateral process with
slender seta at apex (Fig. 14) filitarse

3. Form elongate: ratio of width (across humeri) to length (of pronotum and elytra) 1:1.85 or longer (Fig. 1) haplotes

— Form more ovate: ratio of width to length 1:1.80 or shorter (Fig. 2) 4

4. Pubescence of elytron consisting of piceous to black hairs with subapical patch of light golden or whitish pubescence (rarely limited

to as few as 3 or 4 light-colored hairs) with or without subbasal band of golden or whitish pubescence, or (form limited to southern
deserts) elytron completely covered with whitish pubescence except for few scattered blackish hairs, mostly at apical VS. Both
segments of antennal club identically colored (ochreous to reddish) apicale

— Pubescence of elytron uniformly light golden or piceous but not whitish and without subbasal band or subapical patch of

light-colored hairs. Segments of antennal club variously colored 5

5. Protibia expanded at apex (Fig. 29). Pronotum with punctures of disc about 1 VS times diameter of facet of eye and separated by VS to

2 times diameter of single puncture. Male antennal club dark reddish brown to black triste

— Protibia gradually narrowing from middle to apex (as in Fig. 30). Pronotum with punctures of disc no wider than diameter of facet of

eye and separated by 3 or more times diameter of single puncture, or, if not separated by more than twice diameter of puncture, then
proximal segment of male antennal club light yellowish brown to ochreous. Male antennal club light or dark 6

6. Male with hypomeron continued behind antennal fossa; hypomeron about as wide at narrowest point behind fossa as width of

segment 3 of antenna. Male and female antennal clubs with segment 10 light yellowish brown or light ochreous and segment 11

usually somewhat darker, particularly on anterior side and at apex . Pronotum and elytra usually dark mahogany brown to black with
pronotum same color as elytra (lighter colored specimens probably teneral) fusculum

— Male with hypomeron behind lateral VS of antennal fossa obliterated or narrowed to no more than half width of segment 3 of antenna

(not including width of carina bounding fossa). Segments of antennal club in males and females concolorous or segment 11 darker
than segment 10, but when segment 11 darker, then segment 10 rufous rather than light yellowish brown or ochreous. Head and
pronotum usually black to dark mahogany with elytra rufous; elytra frequently dark mahogany at base and along basal half of median
suture and rufous apically and along sides 7

7. (Species indistinguishable by external morphological characters.) 8th (morphological) sternum more or less evenly rounded apically

with cluster of simple setae at middle and without extended median process bearing two recurved setae (Fig. 9); aedeagus without
proximally directed hook at genital pore (Fig. 22). Western California deserts from Inyo Mts. to Yuma rubidum

— 8th (morphological) sternum with distal process bearing tight cluster of acutely pointed setae and two, long, recurved setae at apex

(Fig. 13, 17); aedeagus with proximally directed hook at opening of genital pore (Fig. 24). Western Texas to SW Wyoming across
Utah and Nevada to California; in California in coastal range and along coast uteanum

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Beal: California Cryptorhopalum

hypomeron sharply angled with or without feeble, thread-like
carina. Mesostemal disc completely divided by deep and broad
sulcus for reception of prostemal process. Metastemal episternum
less than lA as wide as metasternum. Metastemal epimeron
broadly joined to lateral margin of metastemal coxa. Legs with
tibia spinose on dorsal margin but without teeth; tarsus of hind
leg with first segment subequal in length to or slightly longer
than second segment.

ADULT FEMALES: As males except antennal club about half
size of male club; antennal fossa correspondingly smaller.

DISCUSSION: Members of the genus are characteristically
dermestid-like, having a distinct median ocellus, the hind coxa
grooved for the reception of the femur, and the head hypogna-
thous and partially recessed within the pronotum. They somewhat
resemble members of the genus Anthrenus in size and shape, but
are covered with hairs rather than scales. The most distinctive
character is the 2-segmented antennal club, in which the seg-
ments are subequal or segment 10 is longer than segment 11.
Members of Orphinus Motschulsky, which in the United States
occur only in Florida and Hawaii, have a 2-segmented club, but
segment 1 1 is greatly enlarged and nearly round with segment 10
at most only a third as long.

The genus most nearly related to Cryptorhopalum seems to be
Hemirhopalum Sharp, a Neotropical genus. According to Sharp
(1902) members of Hemirhopalum lack a median ocellus, have
“a large, but ordinary, laxly-jointed” male antennal club, and
have “the sides of the thorax beneath” with “a large impression
which is shallow behind instead of a depression that exactly fits
the club of the antennae: this impression in front is broadly
open.” I have not seen any specimens with the particular charac-
ters that Sharp seems to have described. I have seen unidentified
Neotropical species that belong to the same species group as C.
haplotes. I suspect that some of the species now placed in Hemi-
rhopalum properly belong in the haplotes group within Crypto-
rhopalum. Obviously there is a definite need for a revisionary
study of the Neotropical members of both genera.

Aside from Hemirhopalum, species of Cryptorhopalum appear
to have the closest affinities with Orphinus and Thaumaglossa
Redtenbacher. This judgment is based on larval as well as adult
characters. Significantly, the larvae of all 3 genera have a cluster
of hastisetae inserted on the membrane behind each side of the
tergum of abdominal segment 7, but not behind any other seg-
ment. The endemic Hawaiian genus Lahrocerus also has larvae
that are quite similar to those of Cryptorhopalum (Beal 1975),
although the elongated form and the many-segmented antennal
club of the adults make them appear more closely related to
Megatoma or Trogoderma. Cryptorhopalum, Orphinus, Thau-
maglossa, and Labrocerus most likely share a more recent
common ancestry than any one of these genera does with any
other genus of Dermestidae for which larval characters are
known.

GEOGRAPHIC DISTRIBUTION: The genus is primarily Neo-
tropical with 120 species currently recognized from South and
Central America. Probably a total of 18 species occur within the
United States, including 3 as yet undescribed from states other
than California. The genus is found across the entire country with
the exception of the northern New England States. The greatest
proliferation of Nearctic species is in the southwest.

DESCRIPTIONS AND DISCUSSION OF
CALIFORNIA SPECIES
Cryptorhopalum triste LeConte

Cryptorhopalum triste LeConte, 1854, p. 111. Reitter, 1880 (1881), p.
44. Jayne 1882, p. 367. Casey, 1900, p. 158. Blatchley, 1910,
p. 594. Casey, 1916, p. 191. Hatch, 1962, p. 290.

Cryptorhopalum nigricorne LeConte, 1861, p. 344.

Cryptorhopalum picicorne LeConte, 1854, p. 111. Casey, 1900, p. 159.

Dillon and Dillon, 1961, pp. 375, 376. Kirk, 1969, p. 57.
Cryptorhopalum modestum Casey, 1900, p. 158 NEW SYNONYMY.
Cryptorhopalum fusciclave Casey, 1900, p. 158 NEW SYNONYMY.

DESCRIPTIONS

ADULT MALES: Dorsal pubescence uniformly piceous to
light golden brown. Dorsal integument immaculate, black to yel-
lowish brown. Antennal club subcylindrical.

Pronotum with punctures of disc about lVs times diameter of
facet of eye and separated by V2 to 2 times diameter of single
puncture. Antennal club light brown to black; ratio of width to
length of segment 10 varying from 1:1.2 to 1:1.7; ratio of length
of segment 11 to length of segment 10 varying from 1:1.1 to
1:1.8. Carina on lateroposterior margin of antennal fossa attain-
ing margin of hypomeron so that hypomeron not continuous
behind fossa or carina not quite attaining margin of hypomeron
so that plane of hypomeron continued very narrowly behind
carina with hypomeron at narrowest point no wider than twice
width of carina bounding fossa. Prosternal process without
median carina. Metasternum with long or short diagonal stria
originating at margin behind mesocoxa and directed toward
lateroposterior angle of metastemum. First abdominal sternum
with single oblique stria extending on each side from anterior
margin of segment beneath trochanter for about % length of seg-
ment. Tibia of front leg slightly expanded and widest at apex
(Fig. 29). Eighth (morphological) sternum with apical margin as
illustrated (Fig. 10); two dorsal setae of median process inserted
close together, their sockets separated by less than width of sin-
gle socket and positioned distad to insertions of setae of ventral
brush; setae of ventral brush erect, compact; each seta with
obtuse apex (Fig. 20). Aedeagus and lateral lobes as illustrated
(Fig. 19); base of aedeagus furcate with hinges widely separated.
Ratio of width (across humeri) to length (of pronotum and elytra)
varying from 1:1.55 to 1:1.73. Length ranging from 1.92 mm to
2.45 mm.

ADULT FEMALES: As males except antennal fossa occupy-
ing about half area of hypomeron. 5th visible abdominal sternum
without foveae. Length ranging from 2.06 mm to 2.83 mm.

LECTOTYPE DESIGNATIONS: The lectotype of C. triste is
herewith designated as the second of the two specimens standing
under the name in the LeConte collection in the MCZ. From the
original description, in which LeConte indicated a range of size
for the species, it is obvious that he had more than one specimen
before him when he wrote the description. The first specimen in
the series, one which bears a “type” label (probably placed on it
by Nathan Banks), is not C. triste as the species was interpreted
by Jayne, Casey, and Hatch, but is C. apicale Mannerheim. This
first specimen happens to bear a label with the species name on it
in LeConte’s handwriting, but judging from the pin holes in the
label, it has been removed at least four times and there is no

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Beal: California Crypt orhopalum

Figures 8-15. Apical margin of morphological abdominal sternum 8 of males of species of Cryptorhopalum. Figure 8 : fusculum. Figure 9: rubidum.
Figure 10: triste. Figure 11: apicale. Figure 12: haplotes. Figure 13: uteanum. Figure 14: filitarse. Figure 15: balteatum.

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 306:1-22.

Beal: California Cryptorhopalum

Figures 16-21. Figure 16: Ventral aspect of aedeagus and lateral lobes of C. haplotes. Figure 17: Detail of apical process of abdominal sternum 8 of C.
uteanum. Figure 18: Same of C. apicale. Figure 19: ventral aspect of aedeagus and lateral lobes of C. triste . Figure 20: Detail of apical process of
abdominal sternum 8 of C. triste. Figure 21: Ventral aspect of aedeagus and lateral lobes of C. balteatum.

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 306:1-22.

Beal: California Cryptorhopalum

certainty that it is now on the specimen on which it was orig-
inally placed by LeConte. Nomenclatural stability can best be
assured by designating the second specimen as the lectotype. The
type locality is San Jose, California.

The lectotype of C. nigricorne LeConte is herewith designated
as the specimen in the LeConte collection in the MCZ bearing the
type label No. 6879. The type-locality is California.

The lectotype of C. picicorne LeConte is herewith designated
as the specimen in the LeConte collection in the MCZ bearing the
type label No. 6878. The pink label on the specimen is
LeConte’s symbol for “Middle States.’’

The lectotype of C. modestum Casey is herewith designated as
the specimen in the Casey collection in the NMNH bearing the
type label No. 37572. The type-locality is Brownsville, Texas.

The holotype of C. fusciclave in the Casey collection in the
NMNH bears the type label No. 37573. The type-locality is
Texas.

GEOGRAPHIC DISTRIBUTION: The species is found from
northern New York State (and doubtlessly into Canada) to Flor-
ida and west to Kansas, Oklahoma, and Texas. No specimens
have been found in the Rocky Mountain states, but it is common
in the Pacific Coast states (Fig. 36). Whether the two populations
are continuous through Canada or (less likely) through Mexico is
not known at present. It is possible that the California population,
which appears quite homogeneous, is an introduction by early
emigrants. Collections of the species have been made in the Mex-
ican states of Nuevo Leon, Tamaulipas, Veracruz, and Puebla,
but, to the author’s knowledge, not on the western side of the
Sierra Madre Occidental.

DISCUSSION: This species can be recognized with moderate
ease, being distinguished by the slightly expanded apex of the
front tibia and the relatively dense punctation of the pronotum.
Some specimens of C. apicale have pronotal punctation almost as
dense as that found in C . triste. However, wherever these two
species are known to occur sympatrically, that form of C. apicale
is found in which most of the elytron is covered with dark pubes-
cence with an area of light colored pubescence present near the
apex of each elytron. The dorsal pubescence of C. triste is
always unicolorous. The two species are also distinguished in
California by the light ochreous to rufous color of the antennal
club in C. apicale and the dark reddish brown to black color of
the antennal club in C. triste. The longer, narrower shape of the
male antennal club and the much narrower rim of the hypomeron
behind the antennal fossa in the male of C. triste will further
serve to distinguish the species. C. triste is more difficult to dis-
tinguish from dark forms of C. uteanum and from C. fusculum,
but the denser pronotal punctation and the expanded apex of the
front tibia should make its separation positive.

INFRASPECIFIC VARIATIONS: Specimens from New Jer-
sey and Maryland are generally narrower than specimens from
California, but the ranges of ratios of width to length slightly
overlap. On the other hand, specimens from Kansas, Oklahoma,
and Arkansas have a range of ratios of width to length broadly
overlapping both those of the East Coast and the West Coast. The
light reddish brown color of the dorsal pubescence and the
piceous to black color of the antennal club characterize speci-
mens from both the East Coast, the West Coast and the Middle
Plains states. Specimens from Florida and along the Rio Grande
River in Texas have a somewhat lighter dorsal pubescence and a
light brown to reddish brown antennal club. Not enough speci-
mens have been available from Mexico to warrant conclusions

about variations in populations there. However, a series of ten
specimens from the State of Puebla in Central Mexico is scarcely
distinguishable from specimens from the Central Atlantic United
States.

ECOLOGY: Adults are collected on a wide variety of flowers.
The most extensive records of their occurrences on flowers have
been made by A.R. Moldenke (P.H. Raven, in litt.) for collec-
tions from San Mateo County, California. Why they come to
some flowers and not to others is not apparent, but it will be
noticed from the following list that their most frequent occur-
rences are on members of the lily family and on flowers clustered
into inflorescences. In the following list, flower species on which
three or more collections of C. triste have been made are fol-
lowed by an asterisk. Liliaceae: Brodiaea hyacinthina (Lindl.)
Baker*, B. peduncularis (Lindl.) Wats., Calochortus venustus
Dougl.*, C. luteus Dougl., Muilla maritima (Torr.) Wats.; Iri-
daceae: Sisyrinchium bellum Wats.; Ranunculaceae: Ranunculus
californicus Benth.; Cruciferae: “mustard”; Rosaceae: Amelan-
chier sp., Adenostoma fasciculatum H. & A.*, Prunus ilicifolia
Walp., Rosa sp., Potentilla sp.; Leguminosae: Melilotus sp.;
Polygalaceae: Polygala alba Nutt.; Rhamnaceae: Ceanothus
sorediatus H. & A., Ceanothus sp.; Cactaceae: Opuntia sp.;
Umbelliferae: Lomatium utriculatum (T. & G.) C. & R., Daucus
carota L., Heracleum lanatum Michx.; Cornaceae: Cornus sp.;
Asclepiadaceae: Asclepias sp.; Hydrophyllaceae: Phacelia sp.,
Eriodictyon californicum (H. & A.) Greene*; Scrophulariaceae:
Penstemon sp.; Compositae: Achillea borealis Bong.*, Baeria
sp., Bidens sp.*, Cirsium sp.. Coreopsis sp., Erigeron sp., Erio-
phyllum confertiflorum Gray*, Lasthenia chrysostoma (F. & M.)
Greene*. Rudbeckia hirta L., Solidago sp., Wyethia angustifolia
Nutt., W. glabra Gray. Blatchley (1910) recorded the species in
Indiana on yellow puccoon (Lithospermum sp., Boraginaceae),
red haw ( Crataegus coccinea L., Rosaceae), and goldenrod
( Solidago sp., Compositae).

Cryptorhopalum rubidum new species

DIAGNOSIS: 8th (morphological) abdominal sternum of male
without lateral processes, sublateral marginal papillae, or median
posterior process but with cluster of simple setae at midline near
apical margin; apex of aedeagus without proximally directed
hook.

DESCRIPTIONS

ADULT MALE: Dorsal pubescence uniformly golden brown.
Integument of dorsal surfaces with head, pronotum, and base of
elytra piceous; elytra becoming ochreous red on sides and on
apical half. Antennal club subcylindrical.

Pronotum with punctures of disc about % diameter of facet of
compound eye and separated by 4 to 5 times diameter of single
puncture. Antennal club with both segments fuscous; segment 10
with ratio of width to length 1:1.45; length of segment 11 to
segment 10 with ratio of 1:1.15. Carina on lateroposterior margin
of antennal fossa attaining margin of hypomeron so that plane of
hypomeron not continuous behind fossa (Fig. 6). Prosternal
process with slight transverse convexity and without median
carina. Metastemum with very short, almost transverse stria orig-
inating at posterior margin of socket of mesocoxa. First abdom-
inal sternum with single oblique stria on each side extending
about Vi length of segment from anterior margin of segment
beneath trochanter. Tibia of front leg widest at middle and taper-

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 306:1-22.

Figures 22-21. Figure 22: Ventral aspect of aedeagus and lateral lobes of C. rubidum. Figure 23: Same of C . filitarse . Figure 24: Same of C. uteanum.
Figure 25: Same of C. apicale. Figure 26: Same of C. fusculum. Figure 27: aedeagus and lateral lobes of C. fusculum in lateral aspect; for clarity, apex of
aedeagus is figured ventrad of bridge whereas it is normally positioned dorsad (inside) of bridge.

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 306:1-22.

Real: California Cryptorhopalum

ing very slightly toward apex. Eighth (morphological) abdominal
sternum with apical margin as illustrated (Fig. 9); no apical
median process present but dense brush of setae present at middle
of margin. Aedeagus and lateral lobes as illustrated (Fig. 22).
Ratio of width (across humeri) to length (of pronotum and
elytra): 1:1.67. Length: 1.92 mm.

ADULT FEMALE: Elytra entirely ochreous red. Ratio of
width to length of segment 10 of antenna 1:1.42; ratio of length
of segment 11 to length of segment 10, 1:1.11. Antennal fossa
occupying about half of area of hypomeron. 5th visible abdom-
inal sternum without foveae. Ratio of width (across humeri) to
length (of pronotum and elytra) 1:1.72. Length 2.57 mm.

RANGE OF OBSERVED VARIATIONS: Elytra entirely
ochreous red, or dark brown or black at basal 1/5 gradually
becoming reddish brown on apical half. Antennal club of male
with ratio of width to length of segment 10 varying from 1:1.4 to
1:1.9; ratio of length of segment 1 1 to length of segment 10 vary-
ing from 1:1.2 to 1:1.7. Plane of hypomeron continued behind
antennal fossa or not; if continued behind fossa, then no wider at
narrowest point than ‘A times width of segment 3 of antenna.
Ratio of width (across humeri) to length (of pronotum and elytra)
varying from 1:1.61 to 1:1.76. Length of males ranging from
1.90 mm to 2.26 mm; of females ranging from 2.09 mm to
2.95 mm.

TYPE OF MATERIAL: Holotype male, allotype female, 17 para-
types: Yuma, Arizona, April 13, 1955, Butler and Tuttle. Addi-
tional paratypes as follows: Palm Canyon [Kofa Mts., Yuma
Co.] Ariz., May 29, 1955 (G. Butler and D. Tuttle), 3 spm.;
Roll, Arizona, April 14, 1955 (Butler and Werner), 1 spm.; Inyo
Mountains, California, 7000 to 9000 feet elevation, July 7-11
(Wickham), 16 spms.; Potholes [near Laguna Dam], Imperial
County, California, April 9, 1923 (E.P. VanDuzee), 3 spms.
Holotype and allotype deposited in the CAS collection. Paratypes
distributed to collections of the CAS, LACM, MCZ, NMNH, the
University of Arizona, and the University of Wisconsin.

ETYMOLOGY: The specific epithet is a Latin adjective mean-
ing “reddish” or “brownish” in reference to the color of the
elytra.

DISCUSSION: The reddish orange (rarely reddish brown)
color of the apical V2 and often of the entire elytra contrasting
with the black head and pronotum superficially distinguish this
species from any California Cryptorhopalum except C. uteanum.
Occasional specimens of C. apicale may have elytra that are dark
at the base and rufous apically. These may be distinguished from
specimens of C. rubidum in that the elytral pubescence of C.
apicale is mostly dark with a distinct subapical spot of white
pubescence, or the elytral pubescence is whitish, rather than
golden brown. Specimens of C. uteanum usually have a darker
reddish brown elytral integument than specimens of C. rubidum,
but this is not invariably true. The author knows of no consistent
external character by which these 2 species may be distinguished.
However, the males are easily separated by the very distinctive
differences in the structure of the 8th (morphological) sternum.

ECOLOGY: The specimens from Yuma were collected on
mesquite ( Prosopis sp.), and the 3 females from Palm Canyon,
Arizona, were taken on the mint Hyptis emoryi Torr. in Ives.

Cryptorhopalum uteanum Casey

Cryptorhopalum uteanum Casey 1916, p. 195.

Cryptorhopalum nephianum Casey 1916, p. 196 NEW SYNONYMY.

Cryptorhopalum aridum Casey 1916, p. 196 NEW SYNONYMY.
Cryptorhopalum bakeri Casey 1916, p 197 NEW SYNONYMY.

DESCRIPTIONS

ADULT MALES: Dorsal pubescence uniformly whitish to
light golden brown. Integument of head and pronotum black,
integument of elytra entirely black or reddish with suffused black
area at base and along suture at basal third, or entirely reddish.

Pronotum with punctures of disc equal in diameter to diameter
of facet of compound eye and separated by 3 to 4 times diameter
of puncture. Antennal club black except for suffused dark reddish
area at base of segment 10, or segment 11 reddish black with
segment 10 ochraceous; ratio of width to length of segment 10
varying from 1:1.28 to 1:1.92; ratio of length of segment 11 to
length of segment 10 varying from 1:1.15 to 1:1.51. Antennal
fossa at lateroposterior margin reaching posterior margin of
hypomeron so that plane of hypomeron not continuous behind
fossa, or plane of hypomeron continued very narrowly behind
fossa with plane at narrowest point behind fossa no wider than
twice width of carina bounding fossa. Prosternal process trans-
versely convex without median carina. Metastemum with short
diagonal stria originating at margin behind mesocoxa and
directed toward middle of lateral margin of metastemum. First
(visible) abdominal sternum with single oblique stria on each side
extending from anterior margin of segment beneath trochanter for
about 34 length of segment. Tibia of front leg widest at basal
third and tapering very slightly to apex, or sides parallel from
basal third to apex. Morphological abdominal sternum 8 with
apical margin as illustrated (Fig. 13); two recurved dorsal setae
of median process inserted proximally to ventral brush of setae
with bases separated about twice width of one socket; setae of
ventral brush erect, compact, with acute apices (Fig. 17). Aede-
agus and lateral lobes as illustrated (Fig. 24); base of aedeagus
furcate with hinges widely separated. Ratio of width (across
humeri) to length (of pronotum and elytra) varying from 1:1.60
to 1:1.74. Length varying from 1.94 mm to 2.52 mm.

ADULT FEMALES: As males except that antennal fossa
occupying about V2 area of hypomeron and antennal club cor-
respondingly smaller. 5th visible abdominal sternum without
foveae. Ratio of width (across humeri) to length (of pronotum
and elytra) varying from 1:1.62 to 1:1.80. Length varying from
2.06 mm to 2.83 mm.

LECTOTYPE DESIGNATIONS AND NOMENCLATURE: A
male from Provo, Utah, bearing NMNH type No. 37562 is here-
with designated as the lectotype of C. uteanum. A male from the
same series was dissected and the genitalia found to be similar to
the structures figured for this species.

A female from Nephi, Utah, bearing NMNH type No. 73561
is herewith designated as the lectotype of C. nephianum. This
specimen and the 2 females next to it in the Casey collection
definitely appear to be synonyms of C. uteanum although con-
firming male genitalic characters are not available.

A male from 8,000 ft. elevation in the Inyo Mts. of California
bearing NMNH type label No. 37568 is herewith designated as
the lectotype of C. aridum. Another specimen from the same
series was dissected and found to have genitalic characters iden-
tical to those of C. uteanum.

The unique female from “Mts. near Claremont, Calif.” for
which Casey erected the species C. bakeri (NMNH type No.
37563) has here been placed in synonymy with C. uteanum,
since it seems to match this species better than any other. The
identification carries a certain degree of uncertainty.

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 306:1-22.

Figures 28-35. Figure 28: Posterior view of left front tibia of C. haplotes. Figure 29: Same of C. triste. Figure 30: Same of C. apicale. Figure 31: Lateral
view of apex of aedeagus of C. triste. Figure 32: Same of C. filitarse. Figure 33: Fifth visible abdominal sternum of female of C.filitarse. Figure 34:
Detail of apex of median process of abdominal sternum 8 of C . fusculum in lateral view. Figure 35: Same in ventral view.

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 306:1-22.

Beal: California Cryptorhopalum

DISCUSSION: This species and C. rubidum cannot be distin-
guished by any consistent external character. Males of C.
uteanum and C. rubidum are easily distinguished, however, by
those genitalic characters described for each. Males of C.
uteanum can be distinguished from males of C. fusculum and C.
apicale in that C. uteanum males have a very narrow or obliter-
ated extension of the hypomeron behind the antennal fossa. In
males of both C. fusculum and C. apicale the hypomeron behind
the fossa is at least % as wide as segment 3 of the antenna. Color
characters can usually be used to separate both males and females
of C. uteanum from those of C. apicale and C. fusculum. Speci-
mens of C. apicale usually have a subapical spot of white pubes-
cence on the elytron or are covered with mostly whitish hair with
some intermingled dark hairs, whereas the dorsal pubescence of
C. uteanum is always a uniform light golden white to golden
brown. C. fusculum is ordinarily piceous or black in color. The
integument of California specimens of C. uteanum always
includes a certain amount of rufous coloration on the elytron.

INFRASPECIFIC VARIATIONS: Moderate series of the
species have been taken from San Diego County, California,
southern Utah and northern Arizona, and the southeastern corner
of Arizona. Specimens from San Diego County tend to have
darker, mahogany red elytra in contrast to the lighter ochreous
red of specimens from southern Utah and northern Arizona.
Specimens from the Chiricahua Mts. in southwestern Arizona
and from the Davis Mts. in western Texas are entirely black.

DISTRIBUTION (See Fig. 37) ARIZONA: Cochise Co.:
Chiricahua Mts., July 9, 1959 (D.J. and J.N. Knull); same local-
ity, July 26, 1952 (D.J. and J.N. Knull). Coconino County:
Cameron, June 6, 1967 (C.D. Johnson); Cameron, Little Colo-
rado River, June, 1967 (C.D. Johnson); Soap Creek at Cliff
Dwellers, 7 mi. southwest of Marble Canyon, June 5, 1953
(G.D. Butler). CALIFORNIA: Contra Costa Co.: Berkeley, July
19, 1958 (J.A. Litsinger). Inyo Co.: Silver Canyon, White Mts.,
May 10, 1926 (J.O. Martin). Los Angeles Co.: Downey, May,
1945. San Diego Co.: 1 mi. south Del Mar, May 23, 1971 (M.
and A. Gilbert); Idyllwild, June 20, 1958 (P.S. Bartholomew);
Julian, June 12, 1963 (C.D. Johnson); 2 miles west Mt. Springs
(E.P. Van Duzee); San Diego, May 10, June 8, 9, 12, 1920 (E.
Schiffel). NEVADA: Elko Co.: Wendover, June 12, 1933 (J.T.
Howell). Nye Co.: Tonopah, June 1, 1914. NEW MEXICO: San
Miguel Co.: Las Vegas H.S., July 2, 4, 6, 9 (Barber and
Schwarz). TEXAS: El Paso Co.: El Paso, June 27, 1921 (C.D.
Duncan); Ft. Bliss, May 1, 1915 (J.I. Carlson). Jeff Davis Co.:
Davis Mts., July 6, 1936 and July 15, 1955 (D.J. and J.N.
Knull). WYOMING: Sweetwater Co.: Green River, July 2,
1920.

Four specimens in the CAS are labeled as taken at Seabrook,
Texas (Galveston Co.). One specimen in the collection of V.M.
Kirk is labeled as being from Yankton, S.D. I believe both of
these localities should be reconfirmed, since they are outside the
apparent range of the species.

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 306:1-22.

Beal: California Cryptorhopalum

ECOLOGY: Adults have been collected on the crucifer Stan-
leya sp.( on the garden stock, Matthiola incana (L.) R. Br., and
on carrot blossoms.

Cryptorhopalum apicale (Mannerheim)

Anthrenus apicale Mannerheim 1843, p. 258.

Cryptorhopalum apicale: Jayne 1882, p. 366. Reitter 1880 (1881), p. 44.

Casey, 1900, p. 157. Hatch 1962, p. 290.

Cryptorhopalum haemorrhoidale Horn 1894 (pars), p. 321.
Cryptorhopalum coloradense Casey 1916, pp. 192- 193 NEW SYN-
ONYMY.

Cryptorhopalum tuckeri Casey 1916, p. 192 NEW SYNONYMY.
Cryptorhopalum fontinale Casey 1916, p. 194 NEW SYNONYMY.
Cryptorhopalum grisescens Casey 1916, p. 195 NEW SYNONYMY.

DESCRIPTIONS

ADULT MALES. Dorsal pubescence entirely whitish, or
entirely light golden brown, or a mixture of varying amounts of
whitish with golden brown or blackish hairs, or almost entirely
blackish with a very few golden brown hairs; light colored hairs
of elytra often forming subbasal band and distinct subapical spot
with other hairs of elytra mostly black. Dorsal integument black,
immaculate, or black with subapical rufous spot on each elytron,
or with head and pronotum brownish black and elytra ochreous
except for brownish area at base and along median suture.
Antennal club subcylindrical.

Figure 37. Dots, distribution of C. uteanum. Circles, distribution of C.
rubidum .

Pronotum with punctures of disc about as wide as diameter of
facet of eye and separated by 1 to 3 times diameter of single
puncture. Antennal club ochreous to dark reddish brown; ratio of
length of segment 11 to segment 10 varying from 1:1.1 to 1:1.5;
ratio of length of segment 1 1 to segment 10 varying from 1:1.2 to
1:1.7. Antennal fossa at lateroposterior margin not attaining
posterior margin of hypomeron; plane of hypomeron continued
narrowly behind fossa with plane at narrowest point as wide as %
to 2 times width of antennal segment 3. Pronotal process without
median carina or with very narrow impunctate area suggesting
median carina. Metastemum with diagonal stria originating at
margin behind mesocoxa and extending % to Vi distance toward
lateroposterior angle of metasternum. First visible abdominal
sternum with single oblique stria extending on each side from
anterior margin of segment beneath trochanter to posterior mar-
gin. Tibia of front leg widest near middle and tapering gradually
toward apex. Eighth (morphological) abdominal sternum with
apical median process as illustrated (Fig. 11); two median dorsal
setae at apex of process recurved almost 270°, inserted distad to
ventral brush of setae with bases separated by distance about
equal to width of single socket (Fig. 18). Aedeagus and lateral
lobes as illustrated (Fig. 25); base of aedeagus not furcate; apex
of aedeagus with short, proximally directed hook; lateral lobes
rounding at apices. Ratio of width (across humeri) to length (of
pronotum and elytra) varying from 1:1.46 to 1:1.72. Length rang-
ing from 1.87 mm to 2.64 mm (Fig. 2).

ADULT FEMALES: As males except that antennal fossa
occupying about half of hypomeron with antennal club cor-
respondingly smaller. 5th visible abdominal sternum without
foveae. Ratio of width (across humeri) to length (of pronotum
and elytra) varying from 1:1.50 to 1:1.72. Length ranging from
2.12 mm to 2.69 mm.

LECTOTYPE DESIGNATIONS AND NOMENCLATURE:
Each of the species synonymized here with C. apicale is clearly
assignable to it on the basis of dorsal setal characters. The speci-
men in the NMNH from Tucson, Arizona, bearing type No.
37566 is hereby designated as the lectotype of C. grisescens
Casey. A male from the same series was dissected and found to
have genitalic structures typical of C. apicale.

The lectotype of C. fontinale Casey is herewith designated as
NMNH type No. 37560. It is one of a series of 7 specimens from
Jemez Springs, New Mexico. The genitalic structures of one of
the specimens are well exposed and serve to verify the identifica-
tion of the form with C. apicale.

The unique specimen of C. tuckeri Casey from Tucson, Ari-
zona (NMNH type No. 37555) has not been dissected, but is
clearly C. apicale.

The unique female type of C. coloradense Casey (NMNH type
No. 37556) from Golden, Colorado, lacks a distinct subapical
spot of light-colored pubescence on the elytron but does have a
pronounced subbasal band of light-colored hairs. It falls within
the range of variation of the form of C. apicale found along the
eastern slope of the Colorado Rocky Mts.

GEOGRAPHIC DISTRIBUTION: See the distribution map,
Fig. 29. Specimens have also been collected at widely scatterned
localities in Mexico as far south as Cuernavaca.

DISCUSSION: For most California specimens the presence of
a subapical spot of pale pubescence on the elytron is diagnostic.
Usually there is also a subapical rufous area in the integument
corresponding with the spot of light-colored pubescence. Speci-
mens from the San Francisco Bay area (Santa Cruz and Santa

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 306:1-22.

Beal: California Cryptorhopalum

Figure 38. Dots, Nearctic distribution of C. apicale. Star, Florida locality of closely related unnamed species or form conspecific with C . apicale.

Clara Counties north to Marin, Napa, and Solano Counties) often
have the apical spot reduced to a very few hairs and very rarely
are found lacking the apical spot. Males without a subapical spot
can be distinguished from other California species by the slightly
broader extension of the hypomeron behind the antennal fossa.
Both males and females can usually be recognized by the color of
the antennal club, which is a light yellowish brown to a medium
reddish brown, but never a dark brownish black. Furthermore,
both segments of the club are the same color. The base of seg-
ment 10 may be lighter in color than the apex of segment 11, but
there is no change of color between the apex of segment 10 and
the base of segment 1 1. Desert forms of C. apicale may be cov-
ered almost entirely with whitish pubescence, but forms from the
California deserts, in contrast to those from Southern Arizona
deserts, seem always to have a few intermingled dark hairs, par-
ticularly at the apical Vs of the elytra. In the case of a few speci-
mens collected near Yuma, Arizona, the “darker” hairs are a
light golden brown and are not quickly distinguishable from the
whitish hairs.

SUBSPECIFIC VARIATIONS: Because this is a highly poly-
typic species, it is not surprising that Casey should have
described 5 of its different forms as separate species. Each form,
however, intergrades more or less completely with neighboring
forms. In the north of California and extending south along the
Sierra Nevada to the San Gabriel Mountains, specimens are
found with the integument black except for a large rufous area on

the apical declivity of the elytron. The setae are brownish black
except for a few light golden hairs along the lateral and posterior
margins of the pronotum, a small lateral patch at the basal lA of
the elytron, and a prominent subapical patch. In the San Francis-
co Bay area the apical rufous area is much reduced or absent and
the number of light golden hairs is reduced considerably. In the
coastal ranges and valleys of Southern California, specimens tend
to have a large subapical elytral patch of light golden hairs with a
somewhat pronounced subbasal band of light golden hairs. The
most abrupt transition occurs between the montane and the desert
areas, where the light colored hairs become whitish rather than
golden and the areas of light-colored hairs expand to cover the
pronotum and most of the elytra.

ECOLOGY: Adults have been recorded from flowers of 14
families of plants. In comparison with the number of collections
that have been made of the species, plant records are too sparse
to draw any firm conclusions with respect to the factors that may
make one plant more attractive than another. In the following
list, any plant species on which C. apicale has been collected 3
or more times is followed by an asterisk. Liliaceae: Brodiaea hya-
cinthina (Lindl.), Calochortus venustus Dougl.*; Salicaceae:
Salix spp.*; Fagaceae: Quercus sp.; Cruciferae: Brassica sp.;
Capparidaceae: Cleome sp.; Rosaceae: Rubus sp., Adenostoma
fasciculatum H. & A.*, Holodiscus discolor (Pursh) Maxim.,
Physocarpus sp., Prunus ilicifolia Walp.*; Leguminosae: Meli-
lotus alba Desr.*, Astragalus sp.; Rhamnaceae: Ceanothus sore-

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 306:1-22.

Beal: California Cryptorhopalum

diatus H. & A.*, C. integerrimus H. & A.*, C. cuneatus (Hook)
Nutt., Ceanothus sp.*, Rhamnus crocea Nutt.; Malvaceae: cot-
ton; Tamaricaceae: Tamarix gallica L.*; Umbelliferae: Hera-
cleum lanatum Michx., Ligusticum apiodorum (Gray) C. & R.,
Angelica tomentosa Wats.*, “wild parsnip”; Asclepiadaceae:
Asclepias sp.; Boraginaceae: Cryptantha sp.; Rubiaceae: Galium
boreale L.; Caprifoliaceae: Sambucus mexicana Presl.; Composi-
tae: Baccharis glutinosa Pers.* Baccharis sp., Senecio salignus
DC.

In San Mateo County, California, where C. apicale occurs
sympatrically with C. triste, both species occur with moderate
frequency on Calochortus venustus and Adenostoma fasciculatum
(P.H. Raven, in litt.). C. apicale occurs with great frequency on
species of Ceanothus, on which C. triste is taken only occasion-
ally. On the other hand, although C. apicale occurs on various
Compositae elsewhere in its range, it has not been taken on any
composites in San Mateo County. Yet C. triste occurs frequently
in San Mateo County on a number of composites, particularly
Achillea borealis, Eriophyllum confertiflorum, and Lasthenia
chrysostoma.

At Granite Reef Dam, Maricopa County, Arizona, a moderate
series was taken on March 18, 1974, on catkins of Salix sp. and
one specimen on Tamarix gallica. However, no specimens could
be collected on other adjacent flowers, including Baccharis sp.,
Nicotiana glauca Graham, or Pluchea sericea (Nutt.) Coville.

Records of occurrence of adults have not been kept for any one
locality through a long enough period of time to discover any
correlations between dates of their appearance and environmental
factors. Across the entire range for the species, adults have been
taken between December 30 and October 23. In California west
of the Sierra Nevada, the greatest number of collections were
made in the two week period from April 22 through May 5. In
the deserts of California and Arizona, the greatest number of col-
lections were made in the two week period of July 29 through
August 11, no doubt correlations with blooming periods.

Cryptorhopalum fusculum LeConte

Cryptorhopalum fusculum LeConte 1854, p. 111. Reitter 1880 (1881),
p. 44. Casey 1900, p. 158.

Cryptorhopalum triste war. fusculum: Jayne 1882, p. 367.
Cryptorhopalum anthrax Casey 1900, p. 157 NEW SYNONYMY.
Cryptorhopalum granum Casey 1900, p. 157 NEW SYNONYMY.
Cryptorhopalum affine Casey 1900, p. 157 (non affine Reitter 1880
(1881) ). Sharp 1902, p. 657 REVISED SYNONYMY.
Cryptorhopalum pumilum Casey 1900, p. 158 NEW SYNONYMY.
Cryptorhopalum caseyi Dalla Torre 1911, p. 73 (nov. nom. proC. affine
Casey 1900).

Cryptorhopalum piceum Casey 1916, p. 198 NEW SYNONYMY.
DESCRIPTIONS

ADULT MALES: Dorsal pubescence uniformly light golden to
dark golden brown; integument of dorsal surfaces immaculate,
mahogany brown to black. Antennal club subcylindrical.

Pronotum with punctures of disc minute, % times as wide as to
equal in width to diameter of single facet of eye; punctures sepa-
rated by distance equal to 3 to 5 times diameter of single punc-
ture. Antennal club with segment 10 entirely light yellowish
brown or light yellowish brown at base becoming fuscous apical-
ly; segment 1 1 usually entirely fuscous but occasionally light yel-
lowish brown at base; ratio of width to length of segment 10
varying from 1:1.2 to 1:1.7 (Fig. 7). Plane of hypomeron con-

tinued behind antennal fossa with hypomeron at narrowest point
behind lateroposterior margin of fossa equal to or not less than
4/5 as wide as segment 3 of antenna (Fig. 7). Prosternal process
transversely flat to slightly concave without median carina or
rarely with low, thread-like carina on apical half. Metasternum
with short, oblique stria originating at medial margin of meso-
coxal socket and extending not more than Vi distance toward
middle of lateral margin of metastemum. First visible abdominal
sternum with single oblique stria on each side extending from
anterior margin of segment beneath trochanter Vi to % length of
segment. Tibia of front leg widest at middle and tapering slightly
toward apex. Eighth (morphological) abdominal sternum with
apical margin as illustrated (Fig. 8); two recurved setae inserted
dorsally on median process; setae of ventral brush inserted
apically on process, compactly arranged, with rounding apices
(Figs. 34, 35). Aedeagus strongly curved dorsoventrally; apex
without proximally directed hook (Fig. 27). Lateral lobes with
long, somewhat coarse setae at apices (Fig. 26). Ratio of width
(across humeri) to length (of pronotum and elytra) varying from
1:1.51 to 1:1.70. Length ranging from 1.63 mm to 2.50 mm.

ADULT FEMALES: As males except that antennal fossa
occupying about Vz of hypomeron and antennal club correspond-
ingly smaller. Fifth visible abdominal sternum without foveae.
Length ranging from 2.06 mm to 2.66 mm.

NOMENCLATURE: The female holotype of C. fusculum
(MCZ type No. 6880) from “Colorado River, California,”
seems quite certainly to be identical with the species described
here. The dark color of the dorsal integument practically excludes
its identity with C. rubidum. The only California species with
which it might be confused is C. uteanum, but this does not
occur where LeConte probably collected his specimen.

The female holotype of C. affine Casey (NMNH type No.
37565) from Benicia, Solano Co., California, is identical with
this species. On my advice, Mroczkowski (1968) synonymized
C. affine Casey with C. triste LeConte. A reexamination of the
specimen shows that I was in error. The apex of the front tibia is
definitely not expanded, as is true of specimens of C. triste.

Also identical with this species are the unique types of C.
anthrax Casey (NMNH type No. 37570), a female from Arizona,
C. piceum Casey (NMNH type No. 37564), a female from
Claremont, California, and C. pumilum Casey (NMNH type No.
37571), a female from Arizona. Although confirming genitalic
characters are unavailable, these all have setal characters
typical of C. fusculum. The male type of C. granum Casey
(NMNH type No. 37569) from Arizona was dissected and found
definitely to be this species.

GEOGRAPHIC DISTRIBUTION: See the United States dis-
tribution on the map, Fig. 30. In addition to the localities shown
on the map, the species has been found in Baja California, Mex-
ico, at the following localities: 10 miles south of Punta Prieta,
June 21, 1938 (Michelbacher and Ross), 44 km south of Tijuana,
June, 1952 (N.L.H. Krauss), 19 miles east of Rosario, June 17,
1938 (Michelbacher and Ross), and Isla Partida, March 23, 1953
(P.H. Arnaud). It was taken in Sonora, Mexico, at Cumbre del
Frente, 3 miles east of Guaymas, April 30, 1952 (J.P. Figg-
Hoblyn).

DISCUSSION: Within California this species is most likely to
be confused with C. rubidum or C. uteanum. Normally the dark
color of the integument will distinguish it from these species,
both of which, in California, at least, have elytra that are apically
rufous or entirely rufous or mahogany brown. Occasional speci-

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 306:1-22.

Beal: California Cryptorhopalum

mens of C. fusculum, which I take to be teneral, have rufous
elytra. In these cases males may be distinguished by the greater
width of the hypomeron behind the lateroposterior angle of the
antennal fossa in C. fusculum, but I know of no satisfactory way
of distinguishing such females.

ECOLOGY: In desert areas adults have been collected com-
monly on flowers of the legumes Prosopis juliflora (Swartz)
DC., Acacia greggii A. Gray, and Cercidium microphyllum
(Torr.) Rose and Johnston, on flowers of Tamarix gallica L., on
Eriogonum spp., and on mustards of the genus Stanleya. Across
the entire range of the species, single collections have been
recorded from the flowers of each of the following: Chenopo-
diaceae: Atriplex hymenelytra (Torr.) Wats.; Rosaceae: Pyra-
cantha sp.; Rhamnaceae: Condalia sp.; Asclcpiadaceae: As-
clepias sp.; Compositae: Aster spinosus Benth. It was also
recorded “in apricots” in Sacramento, California. In the Spring
Mts. of Clark County, Nevada (near Bonnie Springs), I collected
specimens in moderate abundance on flowers of Eriogonum sp.
but no individuals could be found on flowers of Salix sp. or Virus
sp. in the same vicinity.

Cryptorhopalum filitarse Casey

Cryptorhopalum filitarse Casey 1900, p. 156.

DESCRIPTIONS

ADULT MALES: Dorsal integument with head black to light
reddish brown, pronotum and elytra mahogany to light reddish
brown. Dorsal pubescence consisting of closely appressed, light
or dark golden brown hairs and golden white to whitish hairs.
Antennal club ovate, rufous.

Pronotum with golden white or whitish pubescence on sides
and on basal lobe and golden brown or mixed golden brown and
whitish hairs on disc; punctures of disc about as wide as diameter
of facet of compound eye and separated by 2 to 3 diameters of
single puncture. Elytra with golden-brown hairs, except for
light-colored hairs distributed as follows: few hairs or dense band
of hairs (as dense as submedian band) along basal margin, nar-
row submedian band, narrow or broad subapical band, light-
colored hairs along lateral margin between submedian and
subapical bands present or absent, small or large apical patch;
subapical band of light-colored hairs and apical patch occasion-
ally confluent. Antenna with ratio of width to length of segment
10 varying from 1:1.1 to 1:1.2; ratio of length of segment 11 to
length of segment 10 varying from 1:1.0 to 1:1.3. Process of
prostemum without median carina. Plane of hypomeron behind
antennal fossa as wide at narrowest point as IV2 times width of
antennal segment 3. Metasternum with or without short, oblique
stria originating at posterior margin of mesocoxal cavity. Front
tibia widest at middle and tapering gradually toward apex. Eighth
(morphological) abdominal sternum as illustrated (Fig. 14); lat-
eral process well developed with terminal seta slender; tuft of
simple setae present at apex of slightly expanded median area.
Aedeagus and lateral lobes as illustrated (Fig. 23); genital pore
simple with small dorsal lobe but without proximally directed
hook (Fig. 32). Ratio of width (across hunieri) to length (of pro-
notum and elytra) varying from 1:1.58 to 1:1.64. Length ranging
from 1.70 mm to 2.06 mm.

ADULT FEMALES: As males except that antennal fossa
occupying about half of hypomeron, and 5th visible abdominal
sternum with 2, circular, very slightly depressed foveae, each

with diameter about equal to half greatest length of segment and
each bounded by feeble, interrupted, thread-like carina (Fig. 33).
Ratio of width (across humeri) to length (of pronotum and elytra)
varying from 1:1.58 to 1:1.70. Length ranging from 1.74 mm to
2. 1 1 mm.

LECTOTYPE DESIGNATION: The NMNH specimen bearing
type label No. 37552, a female, is herewith designated as the
lectotype. The specimen bears a label with the letters “Cal,”
which are underlined. According to the original description, the
locality is Santa Barbara, California. One male and 5 females
next to it bear the same label. The male was dissected and found
to have genital structures similar to those figured here for the
species .

GEOGRAPHIC DISTRIBUTION: CALIFORNIA: Imperial
County: Santa Rosa Mts., July 4, 1946 (D.J. and J.N. Knull).
Inyo County: Inyo Mts., 7,000 to 8,000 ft. elev., July 7-11
(Wickham); Westgard Pass, July 22, 1961 (J.S. Buckett); 3 miles
southwest of Westgard Pass, July 18, 1964 (C.D. Johnson). Los
Angeles County: San Gabriel Canyon, August 8, 1960 (Fred G.
Andrews). Orange County: Fullerton, August 2, 1930 (Barthol-
omew). Riverside County: San Jacinto Mts., July 21, 1929 (L.D.
Anderson). Tulare County: Giant Forest, July 28, 1929 (R.H.
Bcamer). Tuolumne County: Elizabeth Lake [Yosemite National
Park], July 10, 1924 (W. Benedict). Ventura County: Ojai,
August 29, 1950 (F.W. Furry). See map. Fig. 31.

DISCUSSION: This species and C. balteatum are readily dis-
tinguishable from all the preceding species by the presence of 3

Figure 39. Nearctic distribution of C. fusculum.

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 306:1-22.

Beal: California Cryptorhopalum

or 4 distinct bands of light-colored pubescence across the elytra,
by the ovate rather than subcylindrical antennal club, and by the
presence of 2 round foveae on the 5th visible abdominal sternum
of the female. However, the 2 species are very difficult to sepa-
rate from each other on the basis of external characters. Genitalic
characters easily separate the males. The sternum of the 8th
abdominal segment of C. filitarse has a tuft of setae at the middle
of the apical margin, but these are lacking in C. balteatum. The
lateral process of the sternum of the same segment of C. filitarse
bears a slender apical seta, whereas the lateral process of C. bal-
teatum bears an apical seta that is about as wide as the apex of
the process itself. The aedeagus of C. filitarse has a genital pore
that is more or less apical with a small proximal lobe but without
a proximally directed hook as found in C. balteatum.

ECOLOGY: Adults have been collected on flowers of Erio-
gonum sp. and Asclepias sp.

Cryptorhopalum balteatum LeConte

Cryptorhopalum balteatum LeConte 1854, p. 111. Reitter 1880 (1881),
p. 43. Jayne 1882, p. 365.

DESCRIPTIONS

ADULT MALES: Dorsal integument with head and thorax

Figure 40. Dots, distribution of C. filitarse. Circles, distribution of C.
balteatum. Question mark, record of C. balteatum for county only.

piceous to black and elytra dark mahogany brown to rufous. Dor-
sal pubescence consisting of piceous and golden, depressed hairs.
Antennal club ovate, rufous.

Pronotum with golden hairs on sides and on basal lobe and
with piceous hairs covering disc; punctures of disc about as wide
as diameter of facet of eye and separated by 3 to 5 times diameter
of single puncture. Elytra with piceous hairs except for golden
hairs distributed as follows: few or none along basal margin,
patch on humerus coalescing with narrow submedian band, nar-
row or broad subapical band with none along lateral margin
between submedian and subapical bands except for single line of
hairs on lateral carina, patch on apex. Antennal club with ratio of
width to length of segment 10 varying from 1:0.8 to 1:1.5; ratio
of length of segment 11 to length of segment 10 varying from
1:1.2 to 1:1.5. Prosternal process without median carina.
Hypomeron behind antennal fossa at narrowest point about IV2
times as wide as antennal segment 3 (Fig. 4). Metasternum with
fine, sometimes short stria extending obliquely outward from
lateroposterior margin of mesocoxal cavity. Front tibia widest at
apical 3/5 and tapering gradually toward apex. Eighth (morpho-
logical) sternum with stout, evenly curving seta inserted at apex
of lateral process; seta terminating abruptly rather than gradually
becoming filamentous; middle of apical margin of sternum with-
out process or cluster of setae (Fig. 15). Aedeagus with ventrally
directed hook-like process at apex (Fig. 21); lateral lobes with
moderately broad bridge. Ratio of width (across humeri) to
length (of pronotum and elytra) varying from 1:1.50 to 1:1.65.
Length 1.79 mm to 2.28 mm.

ADULT FEMALES: As males except that antennal fossa
occupying about V2 area of hypomeron and antennal club cor-
respondingly smaller. Fifth visible abdominal sternum with 2,
circular, very slightly depressed foveae, each with diameter about
equal to % greatest length of segment, not or indefinitely mar-
gined by thread-like carina. Ratio of width (across humeri) to
length (of pronotum and elytra) varying from 1:1.54 to 1:1.66.
Length ranging from 2. 18 mm to 2.47 mm.

DISTRIBUTION: CALIFORNIA: Kern County, May, 1934
(A.T. McClay). Los Angeles County: Mint Canyon, June 24,
1949 (L.W. Isaac); 2 miles north of Valyermo, June 4, 1970
(R.S. Beal).

SYSTEMATICS: The female holotype (MCZ type No. 6875)
from San Diego, California, appears to be within the limits of the
species described here, but the overlapping range of external
characters between this species and C. filitarse leaves a small
margin of doubt. Two specimens stand under the name in
LeConte’s collection, but the second, which is C. pruddeni
Casey, is dated 1879, so is not part of a type series.

As with many other closely related allopatric forms in which
there is no evidence of a genetically controlled barrier to repro-
duction, C. balteatum and C. reversum Casey could be treated
with equal propriety as distinct species or as subspecies. There
are consistent differences between the two in setal color, but
these seem relatively insignificant in view of the variations of
setal color and pattern found across the range of C. reversum.
Nevertheless, the forms are treated here as distinct species, since
consistent differences, however small, do exist and since a rela-
tively wide barrier of desert seems to separate them. Future col-
lecting, particularly along the Mexican border, might necessitate
a new interpretation of their status. C. reversum presently is
known to occur from the Chiricahua Mountains in the southeast-
ern corner of Arizona east to Brownsville, Texas.

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 306:1-22.

Beal: California Cryptorhopalum

DISCUSSION: This species is in the same species group as the
sympatric C. filitarse. For a discussion of the differences
between the two, refer to the discussion under C. filitarse. Exter-
nally the species is quite similar to C. pruddeni Casey (1900),
which extends from Yavapai and Coconino Counties in Arizona
southeast to Brewster County, Texas, and northeast to Garfield
County, Colorado. Externally the two may be distinguished by the
fact that the dorsal integument of C. pruddeni is more or less
unicolorous, in distinction to the contrasting darker pronotal and
lighter elytral integument of C. balteatum. Also in C. balteatum
the light-colored dorsal setae are a deep golden color rather than
golden-white or whitish as in C. pruddeni. The 8th (morphologi-
cal) abdominal sternum of the males of each is provided with
lateral processes which bear at the apex a stout, curved seta. In
C. balteatum this seta is relatively stout for its entire length and
terminates rather abruptly. In C. pruddeni the seta is stout at the
base but gradually becomes finely filamentous.

C. reversion and C. balteatum have apparently identical male
genitalic characters. The light-colored dorsal setae of C. bal-
teatum are a deep golden color. Specimens of C. balteatum lack
light-colored setae between the subbasal band and the sub median
band of light pubescence. Most specimens of C. reversion have a
mixture of light and dark setae between these two bands as well
as a number of light-colored setae between the basal band and the
subbasal band. However, a few specimens of C. reversum ap-
proach the situation found in C. balteatum.

ECOLOGY: The only information is that adults were collected
2 miles north of Valyermo, California, in flowers of Yucca sp.

Cryptorhopalum haplotes new species

DIAGNOSIS: Ratio of width (across humeri) to length (of
pronotum and elytra) greater than 1:1.86. Eighth (morphological)
sternum of male with sublateral papillae on apical margin but
without lateral processes or median process; cluster of simple
setae inserted along apical margin at middle.

DESCRIPTIONS

ADULT MALE: Dorsal integument with head and pronotum
dark mahogany brown; elytra rufous; each elytron with suffused,
slightly darker area at about basal V4 and median Vs. Dorsal
pubescence mahogany brown, suberect; ventral pubescence
golden brown, recumbent.

Pronotum with punctures of disc minute, about % times as
wide as width of facet of compound eye and separated by 2 to 4
times diameter of single puncture. Antenna and hypomeron as
illustrated (Fig. 3); antennal club reddish brown. Antennal fossa
bounded behind by thread-like carina; width of plane of hypom-
eron at narrowest point behind fossa (exclusive of width of
carina) about as wide as V2 width of segment 3 of antenna. Pro-
sternal process without median or lateral carinae, transversely
flat. Metasternum without oblique striae. First visible abdominal
sternum with single, oblique stria originating beneath trochanter
and extending 3A distance to posterior margin of segment. Tibia
of foreleg as illustrated (Fig. 28). Eighth (morphological) abdom-
inal sternum with cluster of setae at middle of apical margin and
with small papilla-like sublateral process bearing long, slender
setae (Fig. 12). Lateral lobes with somewhat rounding, not trun-
cated, apices; bridge moderately narrow (Fig. 16). Aedeagus
with small, proximally directed hook at apex. Length (of pro-
notum and elytra): 2.38 mm. Width (across humeri): 1.27 mm.

ADULT FEMALE: Integument of head dark mahogany brown;
pronotum rufous with suffused darker areas across disc; elytra
rufous; each elytron with suffused darker area at about basal 14
and median Vs. Antenna and hypomeron as illustrated (Fig. 5).
Visible abdominal sternum 5 without foveae. Length (of prono-
tum and elytra): 3.07 mm. Width (across humeri): 1.54 mm.

RANGE OF OBSERVED VARIATIONS: Pronotum ochreous
to dark mahogany with or without light and dark mottled appear-
ance; elytra ochreous to light brown with or without darker, suf-
fused subbasal areas. Length (of pronotum and elytra) ranging
from 2.23 mm to 3.07 mm. Ratio of width (across humeri) to
length (of pronotum and elytra) varying from 1:1.86 to 1:2.07.

TYPE MATERIAL: Holotype male, allotype female, and 7
paratypes: Jacumba (San Diego County), California, June 19,
1951 (D.J. and J.N. Knull). Holotype and allotype deposited in
the collection of Ohio State University; paratypes deposited in the
collections of the CAS, the LACM, the NMNH, and the collec-
tion of the author.

ETYMOLOGY: The specific epithet is formed from the Greek
noun arrAoTT]? standing in apposition with Cryptorhopalum and
meaning “simplicity.” This is in reference to the simple structure of
morphological abdominal segment 8 of the male.

DISCUSSION: The elongated form will easily distinguish this
species from any other member of the Nearctic fauna. At least
one other species belonging to the same species group occurs in
Mexico in the state of TIaxcala. Whether this species is among
those presently described and assigned to Hemirhopalum is not
known. The Mexican species seems to be readily distinguishable
from C. haplotes by male genitalic characters as well as by the
color of the dorsal pubescence, the shape of the antennal club,
and other external features.

ECOLOGY: Quite a few pollen grains were found adhering to
the ventral surfaces of each of the specimens, indicating that they
were taken in flowers. An examination of the pollen showed it to
have'come from a plant within the order Rosales, but a species
for which the pollen characteristics appear to be undescribed.
Other details of the biological relationships of the species are
unknown.

SYNONYMIES AND LECTOTYPE
DESIGNATIONS

FOR C. REVERSUM AND C. PRUDDENI

Since these two species are very closely allied to C. balteatum
and since a further understanding of C. balteatum probably
depends in part on an understanding of its relationships to these 2
species, it seems appropriate to present synonymies that have
been discovered for them.

Cryptorhopalum reversum Casey

Cryptorhopalum reversum Casey, 1900, p. 156.

Cryptorhopalum festivum Casey, 1900, p. 156 NEW SYNONYMY.
Cryptorhopalum balteatum Casey, 1900, p. 156 (non balteatum LeConte,
1854) NEW SYNONYMY.

Cryptorhopalum pollens Casey, 1916, p. 193 NEW SYNONYMY.

A female from Ft. Wingate, New Mexico, bearing NMNH
type No. 37554 is herewith designated as the lectotype of C.
reversum .

A female from Brownsville, Texas, bearing NMNH type No.

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 306:1-22.

Beal: California Cryptorhopalum

37558 is herewith designated as the lectotype of C. festivum.

A female from El Paso, Texas, bearing NMNH type No.

37559 is herewith designated as the lectotype of C. pallens. The
specimen is badly abraded, so identification with C. reversum is
conjectural .

The single specimen which Casey identified in his collection as
C. balteatum is a female labeled only as collected in Arizona.

Cryptorhopalum pruddeni Casey

Cryptorhopalum pruddeni Casey, 1900, p. 156.

Cryptorhopalum insigne Casey, 1916, p. 191 NEW SYNONYMY.
Cryptorhopalum anthrenoides Casey, 1916, p. 194 NEW SYNONYMY.

Three females comprise the series from which Casey described
C. pruddeni, these taken in Arizona at the “Canon of the Colo-
rado River” (probably on the south rim of the Grand Canyon).
The specimen bearing NMNH type No. 37550 is herewith desig-
nated as the lectotype. It appears to be a teneral individual of the
species.

The female specimen bearing NMNH type No. 37553 from
Jemez Springs, New Mexico, is herewith designated as the lecto-
type of C. insigne.

The unique type of C. anthrenoides from the Santa Catalina
Mts., Arizona, is NMNH type No. 37557.

ACKNOWLEDGMENTS

This study could not have been completed without the gener-
ous cooperation of a number of individuals and institutions. I am
indebted for the loan or gift of specimens to William F. Barr of
the University of Idaho, the late Lutz Bayer of the University of
Wisconsin, Frank F. Hasbrouck of Arizona State University,
Charles L. Hogue of the Natural History Museum of Los Angeles
County, Paul D. Hurd, Jr., formerly responsible for the Califor-
nia Insect Survey of the University of California, Berkeley, C.
Daniel Johnson of Northern Arizona University, David H. Kava-
naugh of the California Academy of Sciences, John M. King-
solver of the Systematic Entomology Laboratory, U.S.D.A.,
National Museum of Natural History, Vernon L. Kirk of the Ento-
mology and Crops Research Division of the Agricultural
Research Service, Brookings, South Dakota, John F. Lawrence
of the Museum of Comparative Zoology, the late A.T. McClay
of the University of California, Davis, Scott McCleve of Doug-
las, Arizona, D.H. Pengelly of the University of Guelph,
Guelph, Ontario, P.H. Raven of Stanford University, H.J. Rein-
hard of Texas A. and M. University, E.P. Rouse of the Uni-
versity of Arkansas, Jerome G. Rozen, Jr., of the American
Museum of Natural History, M.W. Sanderson, formerly of the 111-
nois Natural History Survey, Joe Schuh of Klamath Falls, Ore-
gon, Terry L. Seeno of the California Department of Food and
Agriculture, C.A. Triplehorn of Ohio State University, Robert H.
Turnbow, Jr., of the University of Georgia, Floyd G. Werner of
the University of Arizona, and R.E. Woodruff of the Division of
Plant Industry of the Florida Department of Agriculture. My
thanks are extended to Richard H. Hevly of Northern Arizona
University for some plant and pollen identifications. Additional
thanks are due C. Daniel Johnson, J.M. Kingsolver, Floyd
Werner, and Terry L. Seeno for a critical reading of the manu-
script.

I am particularly appreciative of a grant from the Systematic
Entomology Laboratory, 1IBIII, ARS, U.S. Department of Agri-

culture, which allowed me as a visiting scientist to study type
specimens in the Thomas L. Casey collection and to study mate-
rial deposited in the collection of the United States National Mu-
seum of Natural History.

LITERATURE CITED

Arrow, G.J. 1915. Notes on the Coleopterous family Dermesti-
dae, and descriptions of some new forms in the British Mu-
seum. Ann. Mag. Nat. Hist., ser. 8, 15:425-451.

Beal, R.S., Jr. 1959. A key to the Nearctic genera of Dermesti-
dae. Coleopterists’ Bull., 13:99-101.

. 1975. Description of adult and larval stages of a new

species of Cryptorhopalum from Arizona and Mexico
(Coleoptera: Dermestidae). Proc. Entom. Soc. Washington,
77 (2):228-233.

BLATCHLEY, W.S. 1910. An Illustrated Descriptive Catalogue of
the Coleoptera or Beetles (Exclusive of the Rhynchophora)
Known to Occur in Indiana (The Nature Publ. Co., Indi-
anapolis), pp. 1-1386.

Casey, Thomas L. 1900. Review of the American Cory-
lophidae, Cryptophagidae, Tritomidae, and Dermestidae,
with other studies. Jour. New York Entom. Soc., 8:51-172.

. 1916. Some random studies among the Clavicomia.

Memoirs on the Coleoptera, 7:35-292.

Dalla Torre, K.W. von 1911. Dermestidae. Coleopterorum
Catalogus, 14 (33):39— 96.

Dillon, Elizabeth S. and Lawrence S. 1961. A Manual of
Common Beetles of Eastern North America (Row, Peterson
and Co., Evanston, 111.), pp. i-viii + 1-884 + plates
A-D.

GuErin-MEneville, F.E. 1838. Note sur le genre de Coleopteres
clavicornes, nomme par Latreille Globicornis, et description
d’une espece nouvelle de ce genre. Revue Zoologique,
1:135-139.

Hatch, Melville H. 1962. The beetles of the Pacific North-
west. Part III: Pselaphidae and Diversicomia I. Univ. of
Washington Pub. in Biology, 16:1-503.

Horn, George H. 1894. The Coleoptera of Baja California.
Proc. Calif. Acad. Sci., 4:302-449.

Jayne, Horace F. 1882. Revision of the Dermestidae of the
United States. Proc. American Philos. Soc., 20:343-377,
plates 1-4.

Kirk, Vernon M. 1969. A list of beetles of South Carolina, Part
1 — Northern Coastal Plain. South Carolina Agric. Experi-
ment Sta., Tech. Bull. 1033, pp. 1-124.

LeConte, John L. 1854. Synopsis of the Dermestidae of the
United States. Proc. Acad. Nat. Sci. Philadelphia,
7:106-113.

. 1862. New species of Coleoptera inhabiting the Pa-
cific district of the United States. Proc. Acad. Nat. Sci.
Philadelphia, 1861:338-359.

Mannerheim, Carl G. von 1843. Beitrag zur Kafer-Fauna der
Aleutischen Inseln, der Insel Sitkha und Neu-Califomiens.
Bull. Soc. Imp. Nat. Moscou, 16:175-314.

Mroczkowski, Maciej 1968. Distribution of the Dermestidae
(Coleoptera) of the world with a catalogue of all known

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 306:1-22.

Beal: California Cryptorhopalum

species. Ann. Zool. , Polska Akademia Nauk, 26
(3): 1 5— 191 .

Rees, B.E. 1943. Classification of the Dermestidae (larder, hide,
and carpet beetles) based on larval characters, with a key to
the North American genera. U.S. Dept. Agric. Misc. Pub.
No. 511, pp. 1- 18.

Reitter, E. 1880 (1881). Die aussereuropaischen Dermestiden
meiner Sammlung. Verh. Nat. Ver. Briinn, 19:27-60.

Sharp, David 1902. Dermestidae. In Biologia Centrali-
Americana. Insecta. Coleoptera, 2 ( 1 ):642— 669, plate 19.

Accepted for publication July 8, 1977.

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 306:1-22.

JjWj:

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Number 307
March 16, 1979

CONTRIBUTIONS IN SCIENCE

NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY

A NEW MONOPLACOPHORAN LIMPET
FROM THE CONTINENTAL SHELF OFF SOUTHERN CALIFORNIA

By James H. McLean

J0^ Published by the NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY • 900 EXPOSITION BOULEVARD • LOS ANGELES, CALIFORNIA 90007

CONTENTS

Abstract 3

Introduction 3

Supraspecitic Classification of Living Monoplacophorans 6

Systematic Account

NEOPILINIDAE Knight and Yochelson 1958 9

Neopilina Lemche 1957 9

Verna Clarke and Menzies 1959 9

Subgenus Laevipilina new subgenus 9

Verna (Laevipilina) hyalina new species 11

Discussion 12

Size and Structure 12

Habitat and Ecology 13

Radular Comparisons 15

Origins 18

Acknowledgments 18

Literature Cited 19

ISSN: 0459-8113

Suggested Citation. Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 307:1-19.

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 307:1-19.

A NEW MONOPLACOPHORAN LIMPET
FROM THE CONTINENTAL SHELF OFF SOUTHERN CALIFORNIA1

By James H. McLean2

ABSTRACT: A new subgenus and species of monoplacophoran, Verna (Laevipilina) hyalina, is described
from specimens with a maximum length of 2.3 mm taken on rocks from depths between 174-388 m on the
Santa Rosa- Cortes Ridge of the southern California continental borderland. The shell differs from other
living neopilinids in being nearly transparent and lacking clathrate sculpture. As in the genus Verna, there are
six pairs of gills and distinct postoral tentacles. Verna, on the basis of six pairs of gills is here regarded as
genetically distinct from Neopilina, which has five. On shell characters the new subgenus Laevipilina differs
from Verna in lacking concentric sculpture and in having its structural prisms of a depth equal to their diameter
rather than twice the diameter. Radular comparisons among neopilinids are here made for the first time; the
radula of the new species differs from that of three other neopilinids in having a more prominent first lateral
tooth. Verna (Laevipilina) hyalina is the first monoplacophoran to be verified as living on a rocky substratum
and the first to be found at continental shelf depths. Living specimens are accessible, suggesting that much will
soon be learned about its anatomy and life history.

INTRODUCTION

Until 1952 the monoplacophoran limpets were known only
from the Paleozoic fossils. On shell characters they differ from
modern gastropod limpets in having the muscle scar divided
metamerically. Paleontologists had regarded the fossil genera as
early patellaceans. No advance was made until 1938, when Wenz
(1938: 59) suggested that the symmetrically paired muscle scars
of the Silurian genus Tryblidium might correspond to the arrange-
ment of muscles in chitons. At that point he separated them from
the Patellacea and established the superfamily Tryblidiacea for
the group. Two years later he developed his idea further by con-
sidering them to represent untorted gastropods and distinguished
them from prosobranch gastropods at the subclass level (Wenz
1940). He was the first to mention the name Monoplacophora but
did not use it in a formal sense, stating that N. H. Odhner had
suggested the name to him (see Knight, Lernche, and Yochelson,
1958). Knight (1952) enlarged upon Wenz’s theory and used
Monoplacophora with ordinal rank equivalent to Polyplacophora,
the chitons.

Dramatic proof that monoplacophorans were untorted limpets
became available in 1952 when a living species was discovered
by the Danish Galathea Expedition at abyssal depths in the east-
ern Pacific off Costa Rica. The announcement of the discovery
took place five years later when the species was described by
Lernche (1957) as Neopilina galatheae. Its anatomy was thor-
oughly monographed by Lernche and Wingstrand (1959). Addi-
tional anecdotes about the original discovery were given later
(Lernche 1972).

Neopilina is an untorted limpet with a posterior anus and seri-

ally repeated muscles, gills, and other organs. Neopilina was the
most exciting malacological discovery of the century, a living
fossil — a relict of a once diverse group of mollusks. An addi-
tional living class of mollusks was recognized, now apparently
surviving only in the deep sea.

Further finds of Neopilina since the original discovery proved
that living monoplacophorans are more widely distributed than
was originally assumed. In recent years five more species of
Neopilina have been described and other records of unidentified
species have been published, all found at abyssal or hadal depths.
The second described species, Neopilina ewingi Clarke and
Menzies 1959, from the Peru-Chile Trench, differed from N.
galatheae in having six pairs of gills instead of the five pairs of
N. galatheae. Because of this difference it was made the type
species of the subgenus Verna Clarke and Menzies 1959. Further
discoveries brought the number of described species of Neopilina
to a total of five and those of Verna to two.

My involvement in the study of monoplacophorans began in
1966, when, in connection with my interest in gastropod limpets,
I was given the opportunity to work upon two small specimens in
the S. Stillman Berry Collection. The specimens, not exceeding
2.3 mm in length, had been taken on rocks snagged on hook and

‘Review Committee for this Contribution:

Eugene V. Coan
A. Myra Keen
Robert J. Lavenberg
M. Patricia Morse

2Curator, Section of Malacology, Natural History Museum of Los
Angeles County.

Contrib. Sci. Natur. Hist. Mus. Los Angeles Counts'. 1979. 307:1-19.

McLean: Monoplacophoran Limpet

line by a fisherman, Louis Zermatten, in the vicinity of the
Cortes Bank, due west of San Diego, California, in 1965. One
was from a depth of 95 fathoms (174 meters) and the other from
125 fathoms (229 meters). The rocks bearing the limpets were
saved for John E. Fitch of the California Department of Fish and
Game. He removed the associated mollusks and gave them to S.
Stillman Berry of Redlands, California.

The dried animal of the first specimen was sacrificed for a
radula preparation. Initial study of the radula (Figs. 20, 21) sug-
gested a new group in the Patellidae, with a radula characterized
by a narrow rachidian, three pairs of laterals and two pairs of
flaring marginals. A minute, deep water representative of the
Patellidae, otherwise known from robust intertidal forms, was
unexpected, but it seemed clearly to be the case, based on the
radular evidence. In the hope that more specimens would even-
tually be found, I delayed further work on this remarkable find.
Nine years passed but no additional material came to light.

In 1975 I again turned my attention to the specimens on hand.
Only then did I closely examine the specimen that still contained
the dried animal. Through the dorsal surface of the nearly trans-
parent shell I noted the circularly coiled intestine that is one of
the hallmarks of the described species of Neopilina (Fig. 1).
Then other monoplacophoran features such as metamerically
paired shell muscles and a posterior anus were observed. Upon
rehydrating the animal and examining it in fluid, the shell

Figure 1. Vema (Laevipilina) hyalina new species. LACM
19149, 2.16 mm in length, dorsal view prior to rehydration of
the soft parts and prior to removal of encrustations from the shell
surface. The anterior apex is visible near the top and the intestine
with four coils is seen through the nearly transparent shell
slightly posterior to the midpoint. Photograph by Solis.

became more transparent and its structural prisms visible (Figs.
2, 3). It thereupon became clear to me that this was a mono-
placophoran limpet. Contrary to all previous expectations for the
group, it had come from a rocky substratum in relatively shallow
water.

The rehydration of the specimen containing the dried animal
did not fully restore the features of the ventral surface. Structures
recognized were the head, mouth, velum extending laterally and
posteriorly to the head, radiating pedal retractor muscles, and the
posterior anus. There seemed to be no sign of gills or of postoral
tentacles that also characterize the group. Moreover, the true
shape of the foot could not be discerned. It seemed that it was
partially missing with nothing remaining but a stump in the cen-
tral area. An S-shaped structure that was clearly visible was
interpreted as the radular sac, considering that the radula
extracted from the other specimen was more than half the length
of the shell. The shell appeared completely smooth, devoid of all
traces of clathrate sculpture seen in the described species of Neo-
pilina. The radula also seemed to differ considerably from that of
N. galatheae , the only species for which a radula had been illus-
trated.

At that point in my studies I presented the preliminary findings
to the American Malacological Union — Western Society of
Malacologists joint meeting in San Diego, June 1975. Based on
the smooth shell, radular differences, and apparent absence of
gills, I considered that a new genus was indicated for the newly
discovered species, and so announced an intention to propose
one. The abstract resulting from my presentation was published
30 January 1976 (McLean 1976).

In the fall of 1975 an intensive offshore sampling program was
initiated by the U.S. Bureau of Land Management (BLM) on the
continental shelf of southern California, coinciding with the leas-
ing of offshore tracts for oil exploitation. A group of biologists
headed by Gilbert F. Jones of the University of Southern Cali-
fornia contracted to do the biological portion of the work.
Although most of the stations were made on soft bottoms, there
were some box core stations from rocky areas. Sorters and tech-
nicians were alerted to watch for the new microscopic mono-
placophoran, but after six months of sampling none were found.

Meanwhile, early in 1976 I decided to publish my preliminary
description of the species in the hope that this notice would
stimulate efforts to find other specimens. After more closely scru-
tinizing the rehydrated specimen with improved optics I noted
swellings on the mantle margin in the position where gills could
be expected. Contrary to my earlier observations, six pairs of
gills seemed to be present, although no clear configuration could
be seen.

No radular descriptions had been published on any mono-
placophoran species since the initial monograph of Lemche and
Wingstrand (1959). I was able to obtain specimens of two other
neopilinid species for radular comparisons. The results, discussed
herein, indicated that the two other species bridged the gap
between the radula of the new form and that of N . galatheae.

Although many questions remained unanswered, a draft of the
manuscript was completed and circulated for review. As I was
preparing to submit the paper for publication, two freshly col-
lected specimens were found in sediment from one of the BLM
stations. One had been picked from the residue by the sorters and
subsequently recognized by Patrick I. LaFollette, a member of
the BLM project. He reexamined the residue and found another.
This was the breakthrough I had hoped would occur, for I now

Contrib. Sci. Natur. Hist. Mus. Los Angeles Count y. 1979. 307:1-19.

McLean: Monoplacophoran Limpet

FIGURES 2, 3. Verna ( Laevipilina ) hyalina new species. LACM 19149, 2.16 mm in length, rehydrated and photographed in alcohol.
FIGURE 2, ventral view; FIGURE 3, dorsal view. Shell prisms show at the margin, arranged in curved rows; lighter and darker
concentric rings represent growth lines. Head with mouth near top center. Below the head is the S-shaped radular sac and foot stump.
Tubular rectum at bottom center. Seven pairs of pedal retractor muscles radiate from the center; the first pair is narrow, the second, third,
fourth and fifth pairs are long and broad, the sixth and seventh pairs are short. Beyond the lateral terminations of all but the first pair of
retractor muscles are 6 pairs of gills visible as swellings that blur the shell prisms. The dorsal view shows four dark coils to the intestine
in lower center. In the dorsal view, the light areas encircling the central visceral area are the terminations of the broad bands of the pedal
retractor muscles. Gills represented by the blurred lighter areas between the edge of the shell and the pedal muscle terminations.
Photographs by Draper.

had preserved specimens showing the true condition of the foot
and the clear presence of gills (Figs. 4, 5).

The two fresh specimens were somewhat smaller than the orig-
inal two, with a maximum length of 1 .75 mm. They came from a
box core station on the Santa Rosa-Cortes Ridge between San
Nicolas and Tanner Basins, at a depth of 388 m. Efforts to find
more specimens in sediment residues from other unprocessed
BLM stations from rock bottoms were unsuccessful.

Publication of the paper was withheld until I could learn more
about the anatomy of the species. Although the specimens had
not been fixed for histologic sectioning, it was hoped that sec-
tioning would be possible. 1 asked the help of M. Patricia Morse
at Northeastern University in Nahant, Massachusetts, who along
with her associate, Nathan W. Riser, had work already underway
on Neopilina ewingi.

Once again the material had limitations due to the lack of
proper fixation. Many sections made by Morse and Riser from
the smaller of the two specimens did not hold together and the
results were disappointing. However, they prepared the larger
specimen (Figs. 4, 5) by critical-point drying for viewing with a
scanning electron microscope (SEM). That effort produced some

highly satisfactory pictures on I I February 1977, which are
reproduced here (Figs. 6-8, 11).

In February 1977 further specimens were collected on a three
day cruise of the VELERO led by Heinz A. Lowenstam of the
California Institute of Technology, and assisted by LaFollette,
now a member of the Malacology Section of the Museum. Ef-
forts to recover the species with rock dredges and box cores were
successful at the same locality as the earlier BLM station. Six
living specimens attached to their rock substratum were obtained.
The animals survived for several weeks at ambient water tem-
peratures (Lowenstam 1977, abstract). A report on the behavior,
ecology, and shell ultrastructure will be published elsewhere by
Lowenstam. One of the living specimens was fixed for histologic
sectioning to be done by Morse and Riser, who will also report
separately on their results.

The rocks obtained on the cruise were examined aboard ship
for living specimens and then preserved in 70% ethanol. Further
examination of the rocks by LaFollette produced four more
specimens that became the type lot. One of these specimens (Fig.
9), designated the holotype, is in excellent condition and shows
most of the features now attributed to the species.

Contrib. Sci. Nalur . Hist. Mus. Los Angeles County. 1979. 307:1-19.

McLean: Monoplacophoran Limpet

FIGURES 4, 5. Verna (Laevipilina) hyalina new species. LACM 19150, 1.75 mm in length. FIGURE 4, dorsal view, showing the
regularity of the shell prisms, gills darkly outlined. FIGURE 5, ventral view, showing the 6 pairs of gills, U-shaped anterior lip of the
mouth, the postoral tentacles clearly visible between the mouth and the foot. Photographs by Morse and Riser, light microscope, 4 X
objective.

Supraspecific Classification
of Living Monoplacophorans

The Monoplacophora have been recognized as a separate class
of mollusks for a relatively short period. A revised classification
of the Paleozoic fossil representatives was given by Knight and
Yochelson (1958). Starobogatov (1970), and more recently,
Runnegar and Jell (1976), have offered other versions.3

These classifications have been based largely upon shell form
and the count and configuration of muscle attachment scars on
the shell. The modern monoplacophorans have thin shells that
lack readily visible muscle scars. Unfortunately, details of the
musculature are known only for Neopilina galatheae as given by
Lemche and Wingstrand (1959). There is therefore a poor basis
upon which to compare diversity in the living species with the
considerable diversity indicated in the fossil record.

When Lemche (1957) proposed Neopilina galatheae he placed
it in the family Tryblidiidae, a group otherwise unknown since
the Devonian. Knight and Yochelson (1958) established the sub-
family Neopilininae for the species. Subsequent authors followed
this scheme until Starobogatov (1970) disassociated the group
from the Tryblidiacea altogether by recognizing both a separate
family Neopilinidae and superfamily Neopilinoidea. However,
Runnegar and Jell (1976) retained Neopilina in the Tryblidiidae.

A consideration of the overall classification of Monoplaco-
phora is not within the scope of this paper; there are evidently
some controversial aspects that will not readily be settled. Cer-
tainly a Recent family Neopilinidae may be justified on grounds
in addition to the great disparity in age. The shells of neopilinids
are thin and lack the massive development of the nacreous layer
of the Devonian Triblidiidae (Erben, Flajs and Siehl 1968).

Generic criteria within the Neopilinidae are also lacking a
sound basis for comparison. Verna Clarke and Menzies 1959,
type species Neopilina (Verna) ewingi Clarke and Menzies 1959,
was proposed as a subgenus of Neopilina chiefly on the presence
of six pairs of gills, rather than five of Neopilina.

In the 18 years that have passed since the first two species
were proposed, four more species of Neopilina ( Neopilina ) have
been described, along with one more species of Verna. The num-
ber of gill pairs has proven to be a consistent character in species
of each group regardless of size or growth stage. The supplemen-
tary criteria for Verna, thinner shell and thinner periostracum.

3 While this paper was in its final stage of preparation, I received a useful
review paper on the Recent monoplacophorans (Cesari and Guidastri
1976). The article, in Italian, contains an extensive bibliography that
includes many titles omitted here. Also, an obituary of the late Henning
Lemche (Knudsen 1977) includes references to other papers by Lemche
on Neopilinia and its affinities.

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 307:1-19.

McLean: Monoplacophoran Limpet

Figure 6. Vema ( Laevipilina ) hyalina new species. LACM
19150, 1.75 mm in length, critical-point dried, SEM micrograph.
Lateral view of ventral surface, showing the 6 pairs of gills, the
anterior lip of mouth, the velar ridge extending laterally around
the mouth, the ridge between the mouth and foot bearing the
postoral tentacles. Shrinkage of the body has revealed the pallia]
line midway in the exposed portion of the shell, the nacreous
layer on the innermost portion thick enough to obscure the pat-
tern of hexagonal prisms. X 80.

Figure 7. Vema (Laevipilina) hyalina new species. LACM
19150, length 1.75 mm, critical-point dried, SEM micrograph.
Ventral aspect, enlargement of head area, showing the U-shaped
anterior lip of the mouth, the radula exposed on the odontophore
within. Postoral tentacles visible on the ridge below the mouth.
The velum is a narrow ridge anterior to the mouth and extending
laterally on both sides. X 190.

Figure 8. Vema ( Laevipilina ) hyalina new species.
LACM 19150, critical-point dried, SEM micrograph. Radular
ribbon exposed on the odontophore within the mouth cavity. This
is an enlargement of the area visible in Figure 7, oriented with
the anterior at the top as in the other figures. Rachidian and lat-
eral teeth as drawn in Figure 22. One of the fringed first mar-
ginals is fully exposed, showing depth to the comblike surface.
X 900.

have been maintained in the second reported Vema, N. (Vema)
bacescui Menzies 1968. The two species of Vema are known
only from the southern hemisphere in the eastern Pacific, where-
as the species of Neopilina have been shown to have a broader
distribution in abyssal depths at the base of the continental slope
in the eastern Pacific in both hemispheres and in the Indian
Ocean and mid-Pacific.

Inasmuch as the anatomy of Vema ewingi has not been
described in detail, there is no sound basis upon which to decide
whether the separation of the two groups should be at the sub-
generic, generic or familial level. Starobogatov (1970:301) stated:
“Vema can hardly be regarded as a subgenus of Neopilina.
Moreover it cannot be stated with confidence that Vema is a
member of the same family ... it is in any case appropriate to
await publication of detailed data on the anatomy of Vema."
Nevertheless, for a group in which serial repetition of key struc-
tures is fundamental, other internal organs in Vema can be
expected to have an arrangement differing from that of Neo-
pilina. Once the results of comparative anatomical studies are
available, there will, in my opinion, be ample reasons to consider

Contrib. Sci. Natur. Hist. Mits. Los Angeles County. 1979. 307:1-19.

McLean: Monoplacophoran Limpet

FIGURE 9. Verna (Laevipilina) hyalina new species. Holotype, LACM 1787, 1.94 mm in length, preserved and photographed in alcohol.
Shell prisms show at the margin on the left side where the pallium is lifted away. The pallial fold encircles the head area and borders the
outer side of the pallial cavity, in which six pairs of gills are visible. The anterior lip of the mouth is surrounded by the velum, which
extends laterally into the pallial cavity. Between the velum and the foot is the ridge bearing the postoral tentacles. Foot with thickened
margin, the two posterior lobes of the gonad visible through the transparent sole. The opaque area at the lower left is due to an encrusting
foraminiferan on the outer surface of the shell. Photograph by Draper.

Contrib. Sci. Natur. Hist. Mus. Los Angeles Count}’. 1979. 307:1-19.

McLean: Monoplacophoran Limpet

the two groups to be generically distinct. Although some might
argue that it is premature to make this distinction, it is necessary
to do so in order to recognize a subgenus within the six-gi lied
Verna lineage. There is a major dichotomy between the abyssally
occurring species with sculptured shells and the small, smooth
shelled, more shallowly occurring new species. I believe it pru-
dent to recognize this distinction now at the subgeneric level.

The outline that follows includes all of the taxa proposed to
date in the Neopilinidae4. As more continues to be learned about
the anatomy of the living species, the diagnoses can be enlarged.
Uncertainties now exist concerning the musculature and repeti-
tion of internal structures; discussion of these characters is there-
fore omitted.

SYSTEMATIC ACCOUNT
NEOPILINIDAE Knight and Yochelson 1958

NEOPILININAE Knight and Yochelson 1958:39; Starobogatov 1970:301
(familial rank).

DIAGNOSIS: Shell thin, cap shaped, apex anterior; shell
layers consisting of a thin periostracum, a dominant prismatic
layer, and a thin internal nacreous layer. Eyes lacking, mouth
bordered anteriorly by a velum, posteriorly by postoral tentacles;
foot sole thin, weakly muscularized; anus posterior to foot; gills 5
to 6 pairs, with simple lamellae. Radula with a narrow rachidian,
three pairs of laterals and two marginals, edge of first marginal
fringed.

The family is limited to the known Recent monoplacophorans.
It differs from the Paleozoic families in having an extremely thin
internal nacreous layer of the shell.

Neopilina Lemche 1957

Neopilina Lemche 1957:414. Type species: N. galatheae Lemche 1957.

DIAGNOSIS: Shell with radial and concentric ridges at least in
early stages, periostracum moderately prominent; gill pairs 5.

SPECIES: N. galatheae Lemche 1957 (Costa Rica); N. vel-
eronis Menzies and Layton 1963 (Baja California, Mexico); N.
adenensis Tebble 1967 (Indian Ocean); N. bruuni Menzies 1968
(Peru); N. oligotropa Rokop 1972 (mid-Pacific).

OTHER RECORDS: N. galatheae, Parker 1962 (off Cape San
Lucas, Baja California); unidentified, Menzies 1968 (Costa
Rica); unidentified. Rosewater 1970 (South Atlantic east of Falk-
land Islands); unidentified, Filatova, Vinogradova and Moskalev
1974 (Atlantic- Antarctic); same record identified as N.
galatheae, Filatova, Vinogradova, and Moskalev 1975.

DISTRIBUTION: Abyssal, Eastern Pacific: Baja California,
Costa Rica, Peru; east of southern tip of South America; mid-
Pacific; Indian Ocean.

REMARKS: Of the four species described, N. galatheae is the
only one known from material from other than the type locality.
Neopilina veleronis was described prior to the availability of
scanning electron microscopy. No intact shells of N. veleronis
are extant; shells of the remaining paratypes have been altered by
the preservative (Figs. 14, 15). Shells of the two original speci-
mens of N. oligotropa were thought to have been altered by the
initial preservative (Rokop 1972). Neopilina veleronis and N.
oligotropa were described from specimens not exceeding 3 mm
in length; N. veleronis was considered mature because ripe ova
were recognized by Menzies and Layton (1962); N. oligotropa

was presumed mature, having come from an oligotropic or
food-poor environment. Neopilina adenensis and A. bruuni were
described from single specimens, the latter species briefly diag-
nosed but not described in detail.

Verna Clarke and Menzies 1959

Verna Clarke and Menzies 1959:1027. Type species: Neopilina (Verna)
ewingi Clarke and Menzies 1959.

DIAGNOSIS: Shell thinner, periostracum thinner, sculpture
weaker than that of Neopilina: gill pairs 6.

Subgenus Vema s. str.

DIAGNOSIS: Moderate in size, sculptured with fine radial ribs
and raised concentric ridges especially strong in early stages;
depth of structural prisms twice that of surface diameter. Gill
pairs 6, gill lamellae 5-7.

SPECIES: V. ewingi (Clarke and Menzies 1959) (Peru); V.
bacescui (Menzies 1968) (Peru).

DISTRIBUTION: Abyssal, Pern-Chile Trench.

REMARKS: Vema ewingi (Figures 12, 13) has been collected
at a number of stations (Menzies 1968; Meenakshi et al. 1970)
off Peru, but V. bacescui is known from a single station, the
original number of specimens not mentioned. The latter species
was said to have a distinctive reticulate pattern on the shell sur-
face and a greater abundance of postoral tentacles. Further com-
parative details were not given.

Subgenus Laevipilina NEW SUBGENUS

Type species: Vema ( Laevipilina ) hyalina new species.

DIAGNOSIS: Small (under 3 mm in length), lacking concen-
tric and radial sculpture; structural prisms hexagonal, uniform in
size, their depth equal to their surface diameter. Gill pairs 6, gill
lamellae 2-3.

DISTRIBUTION: Continental Shelf, southern California.

ETYMOLOGY: The subgeneric name, like that of Neopilina,
is based on Pilina, a fossil (Silurian) monoplacophoran of the
family Tryblidiidae. The prefix laevi (from laevis, the Latin
adjective for smooth) emphasizes the unsculptured surface of the
shell in the new taxon.

4One published record may now be removed. Filatova, Sokolova and
Levenstein (1968) and Filatova and Zenkevich (1969) reported finding a
monoplacophoran at a mid-Pacific seamount northwest of Hawaii. The
apex was said to be close to the center of the shell rather than near its
anterior margin as in the known species. It was therefore considered to
represent a new genus in the Tryblidiidae. In response to my inquiry
about the current status of this specimen, she stated (Filatova, personal
communication, 12 July 1976): “As to our small specimen of Neopilina
(?) from Hawaii region it was very young one and it was difficult to
decide about its true systematic position, especially for it had the central
umbo, and it was only a single specimen."

Contrib. Sci. Natur. Hist. Mas. Los Angeles County. 1979. 307:1-19.

McLean: Monoplacophoran Limpet

Figure 10. Vema ( Laevipilina ) hyalina new species. LACM
19149, length 2.16 mm. Lateral profile, the anterior apex at the
right.

FIGURE 11. Vema (Laevipilina) hyalina new species.
LACM 19150, SEM micrograph, fractured edge of shell. The
narrow band below represents the periostracum. From an area
outside the pallial line and therefore lacking the internal nacreous
layer. This shows the hexagonal surface outline of the prisms and
the depth approximately equivalent to the diameter. X 1100.

FIGURE 13. Vema (Vema) ewingi. SEM micrograph, fractured
edge of shell near the outer margin. Some of the prisms have a
hexagonal surface, while others are more irregular and elongate.
Depth of prisms approximately twice the diameter of those with
hexagonal surface. Compare with same view of N. hyalina in
Figure 11, noting differences in magnifications. X 240.

Figure 12. Vema (Vema) ewingi (Clarke and Menzies 1959).
Anterior face of broken shell showing the bulbous apex at the
left; strong concentric and weaker radial sculpture is visible in
early growth stages. LACM 65-11, 6200 m, 1 10 mi. W of Cal-
lao, Peru, R/V ANTON BRUUN, 24 November 1965. Photo-
graph by Draper. X 45.

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 307:1-19.

McLean: Monoplacophoran Limpet

Vema ( Laevipilina ) hyalina

NEW SPECIES
Figures 1-11, 20-22

McLean 1976:60 (abstract, not named); Cesari and Guidastrl 1976:235
(review, McLean 1976); Lowenstam 1977:1076 (abstract, report-
ing collection of living specimens of “a new monoplacophoran
species”).

DIAGNOSIS: Vema hyalina differs from other 6-gilled neo-
pilinids in its smaller size at maturity, lack of concentric and
radial shell sculpture, shallow depth of the structural prisms and
reduced number of gill lamellae.

SHELL: Small (largest known specimen 2.3 mm in length),
thin, transparent, iridescent; surface smooth, clathrate sculpture
lacking. Periostracum thin, visible as a yellow sheen; prismatic
layer thick; internal nacreous layer thin. Pallial line visible at
one-fourth the distance from the margin to the center, inside of
which the thicker nacreous layer obscures the pattern of prisms.
Muscle scars not apparent. Staictural prisms visible under mag-
nification, irregular in shape, usually hexagonal, some pentag-
onal, not elongate on surface; depth of prisms approximately
equal to their diameter; prisms arranged in concentric rows corre-
sponding to growth lines at the shell margin and also showing a
pattern of curved lines radiating from the mid-dorsal region. Base
of shell in one plane, outline evenly elliptical, length 1.2 times
that of width; maximum shell height anterior to mid-point. Apex
smoothly papillate, positioned slightly behind anterior margin at
half the shell height, projecting and rendering anterior face
slightly concave. Apical area completely transparent, not show-
ing defined structural prisms, clear area extending for approxi-
mately 10% of shell length.

EXTERNAL ANATOMY: Foot large, thin, oval, nearly trans-
parent, its base outlined by a thickened ridge. Gills six pairs; in
mature specimens the first two pairs bilobate, next four with
three fleshy fingerlike lobes; located in pallial groove between
foot and pallial fold. Pallial fold a well-defined ridge in front of
head and extending on sides' halfway between foot and shell
edge, its position corresponding to pallial line on shell interior.
Pallium of thin mantle tissue extending to shell margin outside of
pallial fold. Mouth surrounded on front and sides by thick
U-shaped anterior lip. Velum a narrow ridge in front of anterior
lip, extending laterally on either side, terminating in large tri-
angular flaps that extend into pallial groove. Postoral tentacles
located on both ends of a ridge extending laterally between
mouth and foot; tentacles prominent and finely branched into at
least six branches. Anus posterior to foot, close to mantle mar-
gin.

INTERNAL ANATOMY: Gut coils four, visible through
transparent shell, coils filling space between midpoint and half
the distance to posterior edge of shell, final coil extending to
anus. Gonad visible through semi-transparent foot sole, divided
posteriorly into two broad lobes. Radula sac S-shaped, prom-
inent. Pedal retractor muscles tentatively seven pairs; first pair
narrow; second, third, fourth, and fifth relatively broad and long;
sixth and seventh pairs broad and short. Gills positioned just out-
side terminations of retractor muscles on shell, terminations of
first pair of muscles without corresponding gills.

RADULA: Ribbon of original specimen 1.6 mm long, about
70% of shell length, with 25 rows of fully developed teeth and 17
rows in developmental stages. Each inverted V-shaped row
approximately .08 mm wide, containing a rachidian, 3 pairs of

lateral teeth, and 2 pairs of marginal teeth. Rachidian tooth long
and slender, four times longer than wide, with a slightly swollen
and overhanging tip. Shaft of first lateral tooth of similar diam-
eter, its tip more than twice the width and with a smooth over-
hanging edge. Second lateral similar to the first, its free end
nearly three times width of the shaft. Third lateral smaller, free
end about twice its width, positioned below the second lateral.
First marginal large, positioned well below the lateral teeth,
broadly triangular, basal outline faintly visible, free edge broadly
curved and finely fringed; extending past midpoint of ribbon and
overlapping with opposite tooth, extending on outside well
beyond basal membrane. Second marginal small, free edge
smooth, extending beyond edge of basal membrane.

TYPE MATERIAL: Holotype, Los Angeles County Museum
of Natural History (LACM) 1787, 2 paratypes LACM 1788, 1
paratype National Museum of Natural History (USNM) 758556.
Other specimens from the vicinity of the type locality retained by
Heintz A. Lowenstan, California Institute of Technology.

TYPE LOCALITY: Santa Rosa-Cortes Ridge (32°59.0' N,119°
32.8' W), depth 373-384 m. Collected by Heinz A. Lowenstan
and Patrick I. LaFollette, R/V Velero IV sta. 25765, 14 February
1977.

ADDITIONAL RECORDS:

1) LACM 19148, radula slide (Figs. 20-22) and shell frag-
ments on SEM stub. One specimen. 95 fm (174 m), between
Cortes and Tanner Banks, California (32° 41 ' N, 119° 17.3' W),
February 1965, collected by Louis Zermatten, ex S. S. Berry
Collection, no. 33351 Shell used for SEM analysis by Lowen-
stam.

2) LACM 19149, rehydrated specimen (Figs. 1-3). One
specimen, 125 fm (229 m), Cortes Bank, California (32° 25.8'
N, 119° 13.5' W), April 1965, collected by Zermatten, ex Berry
Collection no. 33333. Specimens from these two localities were
attached to rocks brought up by hook and line; the hooks had
been snagged in bore holes made by pholad bivalves. The dried
specimens were removed by John E. Fitch and given to Berry.
Coordinates were supplied by Fitch in 1976 after consulting with
Zermatten. Both localities were productive fishing sites for the
red rock cod Sebastes miniatus and were regularly fished by
Zermatten.

3) LACM 19150, scanned specimen (Figs. 4-8, 11). Two
specimens, 388 m, Santa Rosa-Cortes Ridge, California (32°
58.85' N, 119° 33.05' W), 1 May 1976, R/V VELERO IV sta.
24904, BLM sta. 569 (Figure 16, bottom photograph). This was
the first record of the species at the type locality. The specimens
were found in the alcohol-preserved rock and gravel residue by
LaFollette. One specimen was used in an attempt at sectioning by
Morse and Riser, the other remaining specimen was critical-point
dried and examined with SEM.

DIMENSIONS:

length

width

height

LACM 1787 (holotype)

1.94

1.62

0.55

LACM 1788a (paratype)

1.54

1.22

0.41

LACM 1788b (paratype)

0.81

0.66

0.26

USNM 758556 (paratype)

1.94

1.62

0.59

LACM 19148 (radula)

2.16

1.70

0.70

LACM 19149 (rehydrated)

2.28

1.93

0.64

LACM 19150 (scanned)

1.75

ETYMOLOGY: The specific

name is an

adjective

based on

the Greek noun hyalos (glass), to denote the glassy, transparent
nature of the shell.

Contrib. Sci. Nolur. Hist. Mus. Los Angeles County. 1979. 307:1-19.

McLean: Monoplacophoran Limpet

DISCUSSION

The discovery of a monoplacophoran living at a depth consid-
ered to be the approximate boundary between the continental
shelf and the continental slope is remarkable and unanticipated on
the basis of previous knowledge of living monoplacophorans.

Prior to the discovery of this species, there had been none of
the abyssal species recovered alive, nor had it been possible to
obtain specimens without the use of an oceanographic vessel
capable of trawling in great depths. Now, 20 years following the
announcement that a group of moliusks supposed extinct in the
early Paleozoic survives today, it is possible to obtain living
specimens and observe them under laboratory conditions.

Verna hyalina is small and therefore is not an ideal laboratory
animal. Nevertheless, interest in the species should enable future
investigators to surmount the inherent difficulties.

Size and Structure

The maximum length known for the species is 2.3 mm. A suf-
ficient number of specimens are known of less than that size, so
that it now seems unlikely that larger ones will be found. The
sectioned specimen, which measured 1.6 mm in length, showed
mature sperm and was therefore considered sexually mature.

Shells of Verna hyalina have a mature look; they do not have a
disproportionately large apical area as would be the case with
immature specimens. The shell structure is so fragile that larger
specimens would be unlikely.

Verna hyalina, the third species of Verna to be described, is the
smallest of the genus, being about one-tenth the size of the
others. Two small-shelled species of Neopilina s. str. are known:
A. veleronis and A. oligotropa. That A. veleronis (Figs. 14, 15)
was described from mature specimens is reasonably certain from
the fact that mature ova were observed in histologic sections, as
noted in the original account of the species. Neopilina oligotropa
was considered likely to be mature because it came from a mid-
ocean environment with limited food sources. The other asso-
ciated fauna was represented by similarly small forms. Size dif-
ferences of this magnitude within genera are somewhat unusual
in moliusks, but are not unknown.

The lack of surface shell sculpture distinguishes Vema (Laevi-
pilina) hyalina from other described neopilinids. The shells of all
the other species are reinforced, at least in their young stages, by
concentric and radial sculpture, which strengthens without greatly
increasing weight or bulk. Although all neopilinid shells are thin
and fragile, they apparently provide sufficient support for the
moderately large species (A. galatheae reaches 37 mm in length)
living at abyssal depths where the physical conditions are rela-

Figures 14, 15. Neopilina veleronis Menzies and Layton, length 2.1 mm. Recent photographs of holotype in alcohol. FIGURE 14,
dorsal view. FIGURE 15, ventral view. The anterior apex is at the top. Five coils of the intestine are visible through the translucent shell.
The shell is now considerably decalcified; the visible nodular structures that remain are the intersections of the radial and concentric
sculpture. Photographs by Draper.

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 307:1-19.

McLean: Monoplacophoran Limpet

tively stable. A shallow water counterpart would be expected to
have a stronger shell, one better suited to an environment in
which physical stresses are greater. The habitat of V. hyalina at
the edge of the continental shelf is more variable than that of the
deep sea but is far less rigorous than that of intertidal or sublit-
toral zones. Verna hyalina has, however, the most fragile shell of
any of the species. Its existence in a somewhat more variable
environment seems to be at the cost of a major reduction in size.

Shell structure of Verna hyalina follows the basic neopilinid
plan, summarized most recently by Meenakshi et al. (1970).
Present are the periostracum, prismatic, and nacreous layers. The
periostracum is extremely thin. The prismatic layer forms the
major component of the shell, with the diameter and depth of the
prisms nearly equal. In Verna ewingi (Figs. 12, 13) there are
hexagonal prisms and also some prisms with a more elongate,
surface. The depth of the prisms in V. ewingi seems to be about
twice the diameter at the surface of the regular hexagonal prisms.
In V. ewingi there are also some prisms with surface area smaller
than in the regular hexagons. Small prisms are not found in V.
hyalina. These differences are regarded as more than specific dif-
ferences between the two species of Venia; the differences are
regarded as supraspecffically diagnostic of the subgenera Venia
and Laevipilina.

The internal nacreous layer of neopilinids is very thin (Meen-
akshi et al. 1970:21 1) and this is also true for V. hyalina. Shells
of V. hyalina show a pallial line corresponding to the position of
the pallial fold of the mantle, inside of which the nacreous layer
is thick enough to slightly obscure the pattern of prisms when
viewed from within.

The early developmental stages in neopilinid monoplaco-
phorans are largely unknown. Lemche and Wingstrand (1959:16,
64, figs. 34, 49) described and illustrated a coiled protoconch for
Neopilina galaiheae. No subsequent author has reported a coiled
protoconch in other specimens of N. galatheae or any other
species. Menzies (1968:7, figs. 8a-d) illustrated an unidentified
juvenile Neopilina shell, showing a bulbous, transparent proto-
conch with an abrupt transition to the adjacent shell area, which
is structured with prisms that are elongate on the surface. As
discussed by Menzies, the loss of such a protoconch and subse-
quent repair to the area would produce a circular scar like that
originally reported by Clarke and Menzies (1959: fig. Id) at the
apex of V. ewingi. The apical profile of V. hyalina (Fig. 10) is
similar to that of the unidentified species illustrated by Menzies
(1968: fig. 8c), and it is possible that the protoconch is missing
and the area sealed over. However, in V. hyalina there is no
sharp transition line between the apical tip and the area where
prismatic structure begins; the prismatic structure becomes appar-
ent, faintly at first, some distance away from the apex. It is to be
hoped that future workers will investigate the early development
of the species.

Habitat and Ecology

The localities known for Verna hyalina are on the western edge
of the “southern California continental borderland” (see Emery
1960). Unlike continental shelves in most other areas of the
world, the submarine topography is complex, with high and low
areas, and ridges and troughs running roughly parallel to the
adjacent land mass and its mountain ranges. Some of the topo-
graphic highs form islands, others form shallow banks, and the
lows form a number of deep, closed basins. Patterns of currents

at the surface and at intermediate depths are complex. The locali-
ties known for V. hyalina are on the Santa Rosa-Cortes Ridge, a
submarine range that includes Santa Rosa Island, Begg Rock,
San Nicolas Island, Tanner Bank and Cortes Bank. East of the
area is the San Nicolas Basin, to the south are the East and West
Cortes Basins and to the west the Tanner Basin. Further to the
west is another ridge of lesser elevation and beyond that is the
Patton Escarpment, a steep mud slope that drops to abyssal
depths.

Bottom temperatures at continental shelf depths in southern
California vary but little throughout the year. Joseph L. Reid of
the Scripps Institution of Oceanography informs me that at a
depth of 400 m at positions near the type locality the estimated
range of temperatures is 6.4° to 7.5° C, based on data of the
California Cooperative Oceanic Fisheries Investigations, summa-
rized in data reports from Scripps Institution. The shallowest of
the original records for the species was 174 m. Temperatures at
200 m in southern California are in the vicinity of 8° to 9° C
(Emery 1969: 98). The known bathymetric range for Verna
hyalina is 174 to 388 m; bottom temperatures correspondingly
range from a maximum of 9° to a low of 6.4° C.

It is unlikely that the species will be found at shallower depths,
considering the extent to which sampling at shallow depths has
been done in southern California. However, the lower limit of the
bathymetric range is unknown; we may have sampled only the
upper limits of the species’ bathymetric range. The scarcity of
rocky substrata at greater depths may well be the limiting factor.

There is considerable information available on the bottom con-
ditions at the type locality. When the first two specimens from
the type locality were collected on 1 May 1976 the station was
sampled with a box core and the bottom was photographed just
before the sample was taken (Fig. 16). Bottom conditions were
recorded as follows: “Smooth bottom with pebbles and small
rocks, sparse shell debris.” The photograph shows small rocks
somewhat obscured by fine sediment and some moderately large
invertebrates, including the seastar Rathbunaster californicus,
another seastar, the echinoid Allocentrotus fragilis. a gorgonian,
a massive sponge, and brittle stars. Rocks from the station are of
moderately hard shale and were partially encrusted with several
different species of bryozoans. The chiton Hanley e Ha oldroydi
was common.

The species composition is therefore diverse and includes mol-
luscan predators such as seastars and muricid gastropods ( Boreo -
trophon spp.) that bore through shells. Venia hyalina must have
some methods enabling defense against predators, whether it be a
rapid escape response as used by many shallowly occurring lim-
pets, or seclusion in deep crevices. Study of living animals
should answer these questions.

The histologic sectioning by Morse and Riser of one of the
specimens showed the presence of diatom frustules and sponge
spicules in the gut, suggesting that the species browses upon the
detrital material that accumulates in crevices. For neopilinids
from abyssal depths, Menzies et al. (1959:179) reported that: “A
fecal pellet removed from the hindgut of a specimen of Neopilina
(Verna) showed the presence of diatom frustules, radiolarian
skeletons, pelagic foraminiferal tests and innumerable bacteria-size
particles as well as sponge spicules. This evidence suggests that
Neopilina is a mud-ingesting animal or at least an unselective
detritus feeder.” Thus the feeding habits of all modern neo-
pilinids are probably similar. This is also suggested by the simi-
larity of radular structure among the neopilinids, as discussed in

Contrih. Sei. Nalur. Hist. Mus. Las Angeles County. 1979. 307:1-19.

McLean: Monoplacophoran Limpet

FIGURE 16. Bottom photograph near type locality taken just before sampling with box core. Depth 388 m, Santa Rosa- Cortes Ridge, California
(32° 58.85' N, 1 19° 33.05' W), 1 May 1976, R/V VELERO IV sta. 24904, BLM sta. 569. Bottom consists of small rocks obscured by sediment.
See text for description of the invertebrates pictured. Photograph courtesy of Gilbert F. Jones, BLM project.

detail below. The neopilinid radula has a feathery appearance,
and is not sufficiently robust to enable feeding by rasping or
piercing of prey organisms.

Verna hyalina is the first neopilinid definitely known to be
associated with a hard substratum. The first described species of
Neopilina and Verna were assumed to be free living on soft bot-
toms of ooze on the abyssal sea floor. No rocks or hard-surfaced
objects were reported in dredge hauls. Menzies, Ewing, Worzel,
and Clarke (1959) included bottom photographs of tracks pre-
sumed made by Neopilina, but this observation was later ques-
tioned by Wolff (1961), who suggested that the tracks were made
by a bivalve. No photographs are yet available that actually
reveal the life mode of any of the abyssal species of Neopilina or
Vema.

Not until 1972 with the description of N. oligotropa was the
availability of hard substratum documented along with the taking
of a neopilinid. Rokop (1972) reported the presence of man-
ganese nodules in the vicinity of N. oligotropa but had no evi-

dence to suggest that the species was attached to the nodules. In
1974 Filatova et al. reported that their unidentified Neopilina
from the Atlantic sector of the Antarctic came from a bottom of
mud with sand and pebbles. They considered it a foregone con-
clusion that all Neopilina require a hard substratum: “The hard
substratum to which Neopilina adheres (stones, manganese nod-
ules or pebbles) is dispersed in good quantity in the Antarctic
region by the action of floating ice.’’

Doubts about the habitat of the abyssally occurring species of
Neopilina will persist until some way is found to photograph or
observe living specimens. In view of the apparent absence of
hard surfaces in the abyssal depths off Costa Rica (type locality
of N. galatheae), in the Peru-Chile Trench (type localities of V.
ewingi, V. bacescui, and A. bruuni ) and in the Cedros Trench off
Baja California, Mexico (type locality of N. veleronis), it seems
to me that the weight of evidence would associate the abyssal
species with soft bottoms.

Yet this remains an anomaly because all other limpets (gastro-

Contrib. Sci. Natur. Hist. Mas. Los Angeles County. 1979. 307:1-19.

McLean: Monoplacophoran Limpet

pods of several families) are, without exception, attached to hard
surfaces. However, the foot of Neopilina is somewhat unusual
compared to that of gastropod limpets. Lemche and Wingstrand
(1959:31) prefaced their detailed description of the foot of N.
galatheae as follows:

“The circular foot occupies the central parts of the
ventral side of the animal. Being strongly contracted
in the preserved specimens, its diameter is about
half that of the shell. For descriptive purposes the
foot is here regarded as the central body wall under-
lying the large peri-intestinal blood sinus. Its central
part forms a circular membraneous disc, which, at
places, is little more than 0.1 mm thick and there-
fore somewhat transparent even in the preserved
material. The periphery of the foot is developed as a
prominent muscular foot margin all round the
organ.”

Thus the foot of Neopilina is characterized by its small diam-
eter, extremely thin sole, and highly muscularized margin.
Lemche (1957) had originally postulated that N. galatheae lived
upside down, but that idea was immediately challenged by Yonge
(1957), who considered it possible that the foot could indeed be
too weakly muscularized for creeping on a hard surface, but that
it could prevent the animal from sinking into the bottom ooze.
Clarke and Menzies (1959) reported that the original specimens
of V. ewingi were coated with mucus on the ventral surface and
this was considered to function as an aid to locomotion on soft
bottoms.

Perhaps the forthcoming histological work with V . hyalina will
yield some insight. If the foot of V. hyalina proves to be more
highly muscularized than that of the other species, it will suggest
that the other species are adapted for existence on soft bottoms,
whereas V. hyalina is better adapted to adhere to rocks.

Radular Comparisons

The neopilinid radula has not been discussed since the original
treatment by Lemche and Wingstrand (1959:27, fig. 88) of Neo-
pilina galatheae. Subsequent authors have apparently been hesi-
tant to sacrifice part of their material for whole-mount radular
studies. The radula of V. hyalina differs considerably from that
of N. galatheae. although the basic plan is similar. In order to
make further comparisons I prepared slides from a single speci-
men of V. ewingi in the LACM collection and a paratype of N.
veleronis from the Hancock Collection. Because of the scarcity
of material of all species, I had had to base my observations on
single specimens of each, using standard whole mounting meth-
ods. Future studies should use SEM with its much finer resolu-
tion and depth of field.

The neopilinid radula consists of a narrow rachidian and five
pairs of additional teeth arranged in an inverted V-shaped row
across the ribbon. Lemche and Wingstrand called all five pairs
lateral teeth but gave no reasons for doing so. I regard the inner-
most three pairs as lateral teeth and the outer two pairs as mar-
ginal teeth for two reasons: (1) the fourth tooth, here called the
first marginal, has a fringed edge unlike that of the preceding
laterals; (2) the marginals did not stain as darkly as the laterals in
my preparations using stained, non-resinous mounting medium,
suggesting that the marginals differ from the laterals in thickness
and composition.

Figure 25 herein is redrawn from Lemche and Wingstrand’s
illustration of the radula of Neopilina galatheae. The rachidian
tooth is slender, with no overhanging cusp, resembling the
reduced, vestigial central tooth of some Patellidae. The first lat-
eral tooth is only slightly larger, and has a blunt, overhanging
edge. The second lateral is longer and broader, with a broad
overhanging edge; and the third is similar but placed considerably
below the first. The next tooth, here called the first marginal,
was described as a “membranous tooth” or “comb tooth,” with
its overhanging edge “slit up into some 40 long and curved den-
ticles pointing aborally.” The final tooth, here called the second
marginal, is triangular and projects beyond the edge of the rib-
bon. It has a narrow, overhanging edge.

The radula of Verna ewingi (Figs. 17, 18, 23) differs from that
of N. galatheae in minor ways. One portion of the ribbon (Fig.
17) was stained and mounted in non-resinous medium after slight
treatment with NaOH in an effort to clean it. The teeth remained
aligned as in figure 17. Further cleaning was attempted on an-
other portion of the ribbon, but the teeth folded over in the result-
ing mount (Fig. 18), suggesting that the teeth are more delicate
than those of various gastropod radular ribbons with which I am
familiar. However, the latter preparation does show the teeth in
side view. In V. ewingi, as in N . galatheae, the first lateral is
small, although somewhat larger than the central tooth. The most
significant difference is that the second lateral of V. ewingi
extends well above the position of the first lateral, rather than to
approximately the same level. The second and third laterals seem
to be longer than those of N. galatheae, and show an overhang-
ing tip when viewed in the aligned position, whereas in side view
they are shown to be long and evenly curved (Fig. 18). It is
possible that Lemche and Wingstrand missed the overhanging tip
in their drawing of N. galatheae; the second and third lateral
teeth might therefore be larger than they indicated. The fringed
first marginal of V. ewingi resembles that of N. galatheae and the
second marginal is similar, except that its outer edge is thicker.

I had difficulty interpreting the radula of the only small-sized
species of Neopilina available, N. veleronis, because I was
unable to mount in an aligned position any of five separate pieces
of the ribbon. Here again, this may indicate that neopilinid rad-
ulae are more fragile than gastropod radulae of similar size.
However, all of the teeth may be recognized in the portion
photographed (Fig. 19), even though the rachidian and the
fringed first marginal are not clearly shown. In N. veleronis (Fig.
24) the first lateral is more prominent than in either N. gala-
theae or V. ewingi, and extends slightly above the position of
the rachidian. In addition, the fringed first marginal is broader
than in either N. galatheae or V. ewingi. These differences place
the radula of N. veleronis closer to that of V. hyalina than to that
of the two relatively large-sized neopilinids. However, the sec-
ond marginal is more than twice as long as that in any of the
other species. Neopilina veleronis is unique among the species
studied in the length of the second marginal.

A detailed description of the radula of V. hyalina is included in
the species description above. The whole- mounted ribbon of the
original specimen is shown in figure 20, focused on the shaft of
the lateral teeth, and in figure 21, focused on the fringed edge of
the first marginal. The teeth are drawn in figure 22. After the
drawings were finished I received an SEM view of the radula in
place within the mouth cavity (Fig. 8). The radula differs from
that of both A. galatheae and V. ewingi in having the first lateral
fully developed and about equal in size to the second lateral,

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 307:1-19.

McLean: Monoplacophoran Limpet

Figures 17-21. Neopilinid radular ribbons, magnifications adjusted to show a similar number of teeth rows. FIGURE 17, Vema (Verna)
ewingi, intact ribbon with teeth aligned (LACM 65 — 11, 6200 m, 110 mi. W of Callao, Pern, R/V ANTON BRUUN, 24 November
1965). FIGURE 18, Vema (Vema) ewingi, another portion of same ribbon with lateral teeth turned to the side. FIGURE 19, Neopilina
veleronis, intact ribbon of paratype, teeth not aligned (AHF 603, 2730-2769 m, 30 mi. W of Natividad Island, Baja California, Mexico).
FIGURE 20, Vema ( Laevipilina ) hyalina new species, intact ribbon with teeth aligned, focused on shafts of lateral teeth (LACM 19148).
FIGURE 21, Vema (Laevipilina) hyalina, same ribbon, focused on fringe of first marginal teeth.

instead of the highly reduced condition in these two species. Al-
though the first lateral of N . veleronis is somewhat larger than it
is in the other two species, that of V. hyalina is still the larger.
The fringed first marginal of V. hyalina is much broader than in
N. veleronis. Only in V. hyalina is the fringed tooth so broad that
it overlaps the opposite member in the central part of the ribbon.
The second and third laterals of V. hyalina are not significantly
different from those of the other three species, whereas the sec-
ond marginal is similar to those of N. galatheae and V. ewingi,
but not the exceptionally long second marginal of N. veleronis.

To summarize the radular differences noted among the four
species: Vema hyalina exhibits major differences in two of the
five teeth compared to the two rather similar species N. galatheae
and V. ewingi, whereas the condition of these two teeth in N.
veleronis is intermediate between these two species and V.
hyalina. The radula of N. veleronis is unique in the extreme
elongation of the second marginal.

The radular differences noted in the four species do not corre-
late with the existing generic division based on number of gill
pairs, five in Neopilina and six in Vema. The only correlation is
in size. Both large-sized species have similar teeth and the two

small-sized species have similar teeth. Radular differences among
the species examined are quantitative rather than qualitative, sup-
porting placement of the four species in the same family. A study
of the radulae of the other three living species of neopilinids
should reveal further specific differences.

The radula of neopilinid monoplacophorans is very similar to
that of chitons and patellacean limpets having the docoglossate
radula. In the three groups the radula has in common: a reduced
or absent rachidian; strong, hook-shaped lateral teeth; and few,
weakly developed marginal teeth. Lemche and Wingstrand
(1959) found major similarities in the radular supportive mech-
anism in Neopilina and chitons. Golikov and Starobogatov
(1975) discusssed the similarities in form and function of the
docoglossate radula with that of chitons and neopilinids.

The docoglossate radula has long been known to function in a
way that differs from that of other gastropods. Fretter and
Graham (1962:200) gave a detailed comparison of radular func-
tion in the two kinds of radulas. In the docoglossate radula there
is no longitudinal bending; the entire ribbon works as a rasp with
numerous rows functioning at once. In the rhipidoglossate and
other non-docoglossate radulae, rows of teeth bend longitudinally

Figures 22-25. Radular dentition of neopilinids, drawn from slides photographed in figures 17-21. FIGURE 22, Vema (Laevipilina)
hyalina new species. FIGURE 23, V. (Vema) ewingi. FIGURE 24, Neopilina veleronis. FIGURE 25, N. galatheae, after Lemche and
Wingstrand, 1959. Teeth from left to right are the rachidian; first, second, and third laterals; fringed first marginal; and the outermost,
second marginal. Drawings by Mary Butler.

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 307:1-19.

McLean: Monoplacophoran Limpet

Contrib. Sci. Natur. Hist. Mus. Los Angeles Count} . 1979. 307: 1-19.

McLean: Monoplacophoran Limpet

and the teeth are most effective at the “bending plane.”

Golikov and Starobogatov consider the rasp type of radula —
for which a more technical term has not been proposed — as the
most archaic in the Mollusca. Most other authors, such as Fretter
and Graham (1962), consider the rhipidoglossate radula as the
most primitive and the docoglossate radula derived from it in an
as yet unexplained way. The matter remains controversial
because there is no apparent affinity between these two kinds of
gastropod radulae.

A comblike fringe, similar to that on the neopilinid first mar-
ginal tooth, occurs on the marginal teeth of some genera of the
docoglossan limpet family Lepetidae. In the Neopilinidae the
fringe is found only on the first marginal tooth, but in the Lepeti-
dae the very similar appearing fringe is found on both pairs of
marginal teeth. The origin and significance of these comblike
teeth remains to be explained. Could the fringe be a clue to a
more direct affinity between the Monoplacophora and the Doco-
glossa?

The Paleozoic monoplacophorans are found in shallow water
deposits. They were probably grazing animals like modem patel-
lacean limpets and chitons. The large and robust radular teeth in
modern limpets and chitons are opaque and mineralized (Lowen-
stam 1967, 1971). The neopilinid radula is probably less robust
than that of its fossil predecessors, but is probably similar to what
was present in the extinct families of Monoplacophora. As in the
chitons and docoglossan limpets, the neopilinid radula is large in
proportion to body size. Its mineral content should be investi-
gated.

The radula of the Polyplacophora, the chitons, is not especially
diverse from family to family. I would expect Monoplacophora
as a whole, to have had a diversity similar to that of the Poly-
placophora, in which the level of organization is primitive, the
family distinctions are not profound, and the species are rela-
tively few in number.

Origins

When the discovery of Neopilina galatheae was announced,
who could have predicted that it represented but one of a small
number of species of this relict group? Can it be that the Laevi-
pilina branch in the Neopilinidae is represented elsewhere in the
world? The possibility should be considered by those who have
opportunity to dredge rocks at the edge of the continental shelf.
The offshore fauna of southern California is one of the most often
sampled and best known in the world. This discovery comes at a
time when there are rather few new species being discovered in
the area. Experienced collectors have no doubt had it on hand
before, but have missed it. Many years may pass before the
faunas of rocky bottoms on the outer continental shelves will be
sufficiently known to answer the question.

Unfortunately, a fossil record of the abyssal fauna is not acces-
sible. Many families of modem mollusks seem to have centers of
origin in which extensive speciation has taken place. One might
consider the eastern Pacific as the place of origin of the neo-
pilinid stock, since most of the species and records are from that
region. This might seem to be a foregone conclusion were it not
for those species described from single records in the Indian
Ocean and the mid-Pacific. The most likely explanation seems to
be that accepted by most authors (e.g. Parker 1962) that the neo-

pilinid line is a monoplacophoran offshoot that happened to
invade the deep sea and has existed there in the absence of severe
competition and predation since the Paleozoic. The shallow oc-
currence of Verna (Laevipilina) would thereby represent a reinva-
sion of the shallow water habitat from an abyssal stock.

Yet there is now an alternative theory: there is a possibility that
the monoplacophorans have continued to survive at intermediate
depths since the Paleozoic and that the deep-sea invasion is of
relatively recent occurrence. The discovery of other monoplaco-
phorans from intermediate depths would support this idea. How-
ever, the lack of eyes in Verna ( Laevipilina ) hyalina, a species
that lives at a depth where considerable light is available, sug-
gests that it is a derivative from forms existing in deep, lightless
environments.

ACKNOWLEDGEMENTS

The original discovery of this remarkable new species was due
to the combined efforts of Louis Zermatten, John E. Fitch and S.
Stillman Berry, those who collected, found, and curated the
specimens. I am especially grateful to Dr. Berry for enabling me
to work with the material.

The second find of the species was a supplementary dividend
from the project: Southern California Baseline Studies and
Analysis (FY 1975-1976), under subcontract to Science Applica-
tions, Inc., La Jolla, California (funded by the Bureau of Land
Management, U. S. Department of the Interior). I am grateful to
Gilbert F. Jones of the University of Southern California and the
project staff, especially Patrick I. LaFollette, for the efforts that
eventually produced the specimens.

I thank Heinz A. Lowenstam of the California Institute of
Technology, Pasadena, California, for the donation of four
specimens in the type series. He, along with my assistant Patrick
I. LaFollette, returned to the BLM locality on the R/V VELERO
IV and collected living specimens.

I am especially grateful to M. Patricia Morse and Nathan W.
Riser of Northeastern University, Nahant, Massachusetts, for sec-
tioning and scanning electron microscope study of the two
specimens from the second find. Their notes were a substantial
contribution to this paper. Credit for technical assistance with the
SEM work goes to Edward Seling (Harvard University) and Bart
Yatchmenoff (Northeastern University).

Many helpful suggestions have been received from Eugene V.
Coan, Myra Keen, Patrick I. LaFollette, Robert J. Lavenberg,
Heinz A. Lowenstam, Harold H. McLean, the late Robert J.
Menzies, M. Patricia Morse, and Ellis L. Yochelson.

Photographs are largely the work of Bertram C. Draper, Mu-
seum Associate, with several others by Armando Solis, Museum
Photographer. Radular slides were prepared and photographed by
Jo-Carol Ramsaran, Museum Volunteer, and drawn by Mary But-
ler, Museum Illustrator.

NOTES ADDED IN PROOF: 1) An account of the role of
N.H. Odhner in introducing the name Monoplacophora was pro-
vided by that author (Odhner 1961). 2) In a paper just received
Lowenstam (1978) has described the behavior and illustrated a
living specimen of Vema hyalina, which he referred to as “Mc-
Lean’s Vema sp.” 3) Recent efforts to find rocky bottom at the
two positions mentioned for the original specimens from the
Berry Collection have not been successful.

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979 . 307:1—19.

McLean: Monoplacophoran Limpet

LITERATURE CITED

Cesari, P. and R. Guidastri. 1976. Contribute alia conoscenza
dei monoplacofori recenti. Conchiglie, Milan 12: 223-250.

Clarke, A.H., Jr and R.J. Menzies 1959. Neopilina ( Vema )
ewingi, a second living species of the Paleozoic Class
Monoplacophora. Science 129: 1026-1027.

Emery, K.O. 1960. The sea off southern California: Wiley and
Sons, New York, 366 p.

Erben, H.K., G. FLAJS and A. Siehl. 1968. Uber die Schalen-
struktur von Monoplacophoren. Akad. Wiss. Lit. Abhandl.
Math.-Naturwiss. K. 1968(1): 1-24.

Filatova, Z. A., M. N. Sokolova and R.Y. Levenstein 1968.
Mollusc of the Cambro-Devonian Class Monoplacophora found
in the Northern Pacific. Nature 220: 1114-1115.

Filatova, Z.A., N.G. Vinogradova and L.I. Moskalev
1974. New finding of the ancient primitive mollusc Neo-
pilina in the Atlantic part of the Antarctic. Nature 249: 675.

. 1975 (English ed. 1976). The mollusk Neopilina

(Monoplacophora) in the Antarctic. Acad. Nauk USSR
Okeanologia 15(1): 101-102.

Filatova, Z.A. and L.A. Zenkevich. 1969. Present distribution
of ancient primitive Monoplacophora in the Pacific Ocean
and the fossilized Pogonophora in deposits of Cambrian
seas. Acad. Nauk USSR Okeanologia 9: 162-171 (in Rus-
sian).

Fretter, V. and A. Graham 1962. British prosobranch mol-
luscs. London, Ray Society, xvi + 755 pp., 317 figs.

Golikov, A.N. and Y.F Starobogatov. 1975. Systematics of
prosobranch gastropods. Malacologia 15(1): 185-232.

Knight, J.B. 1952. Primitive fossil gastropods and their bearing
on gastropod classification. Smiths. Misc. Coll. 117(13):
1-56.

Knight, J.B. and E.F. Yochelson. 1958. A reconsideration of
the relationships of the Monoplacophora and the primitive
Gastropoda. Proc. Malac. Soc. Lond. 33: 37-48.

. 1960. Monoplacophora, In R.C. Moore (ed.), Trea-
tise on Invertebrate Paleontology, Pt. I, Mollusca 1, Geol.
Soc. Amer., 77-84.

Knight, J.B., H. Femche and E.L. Yochelson 1958. Petition
requesting clarification of the date and authorship of the Or-
der/Class name Monoplacophora. Bull. Zool. Nom. 15:
555-556.

Knudsen, J. 1977. Obituary: Henning M. Lemche. J. Moll.
Stud. 43(3): 205-207.

Lemche, H. 1957. A new living deep-sea mollusc of the
Cambro-Devonian Class Monoplacophora. Nature 179:
413-416.

. 1972. The discovery of Neopilina, p. 195-201. In

M.M. Jenkins. The curious mollusks. Holiday House, New
York.

Lemche, H. and K.G. Wingstrand. 1959. The anatomy of
Neopilina galatheae Lemche, 1957 (Mollusca, Tryblidi-
acea). Galathea Rept. 3: 9-71.

Lowenstam, H.A. 1967. Lepidocrocite, an apatite mineral, and
magnetite in teeth of chitons (Polyplacophora). Science 156:
1373-1375.

. 1971. Opal precipitation by marine gastropods (Mol-
lusca). Science 171: 487-490.

1977. First recovery of live monoplacophorans. Geo-
logical Soc. Amer., 1977 Annual Meetings, Abstracts with
Programs. 9(7): 1076.

1978. Recovery, behavior and evolutionary implica-
tions of live Monoplacophora. Nature 273: 231-232.

McLean, J.H. 1976. A new genus and species of Monoplaco-
phora from the continental shelf of southern California (Ab-
stract). Bull. Amer. Malac. Union, Inc., 1975: 60.

Meenakshi, V.R., P.E. Hare, N. Watabe, K.M. Wilbur and
R.J. Menzies. 1970. Ultrastructure, histochemistry, and
amino acid composition of the shell of Neopilina. Anton
Bruun Rept. 2: 1-12.

Menzies, R.J. 1968. New species of Neopilina of the Cambro-
Devonian Class Monoplacophora from the Milne-Edwards
Deep of the Peru-Chile Trench, R/V Anton Bruun. Mar.
Biol. Assoc. India, Proc. Sympos. Mollusca, Symp. Ser. 3:
1-9.

Menzies, R.J., M. Ewing, J.L. Worzel and A.H. Clarke, Jr.
1959. Ecology of the Recent Monoplacophora. Oikos 10(2):
168-182.

Menzies, R.J. and W. Layton, Jr 1963. A new species of
monoplacophoran mollusc, Neopilina (Neopilina) veleronis
from the slope of the Cedros Trench, Mexico. Ann. Mag.
Nat. Hist., Ser. 13, 5: 401-406.

Odhner, N.H. 1961. Some notes of the classification of Gas-
tropada. Proc. Malac. Soc. Lond. 34: 250-254.

Parker, R.H. 1962. Speculations on the origin of the inver-
tebrate faunas of the lower continental slope. Deep-Sea Res.
8(3-4): 286-293.

Rokop, F.J. 1972. A new species of monoplacophoran from the
abyssal North Pacific. Veliger 15(2): 91-95.

Rosewater, J. 1970. Monoplacophora in the South Atlantic
Ocean. Science 167: 1485-1486.

Runnegar, B. and P.A. Jell. 1976. Australian Middle Cam-
brian molluscs and their bearing on early molluscan evolu-
tion. Alcheringa 1: 109-138.

Starobogatov, Y.I. 1970. Systematics of Early Paleozoic
Monoplacophora. Paleont. Journal 4: 293-302. (Trans,
from Russian, Paleont. Zhur. 1970(3): 6-17.)

Tebble, N. 1967. A Neopilina from the Gulf of Aden. Nature
215: 663-664.

Wenz, W. 1938. Gastropoda. In Schindewolf: Handb. der
PalaeQzool. vol. 6, pt. 1, Berlin.

. 1940. Ursprung und friihe Stammes-geschichte der

Gastropoden. Arch. Moll. 72: 1-10.

Wolff, T. 1961. Animal life from a single abyssal trawling.
Galathea Rept. 5: 129-162.

Yonge, C.M. 1957. Reflexions on the monoplacophoran Neo-
pilina galatheae Lemche. Nature 179: 672-673.

Accepted for publication August 1, 1977.

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 307:1-19.

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NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY

SERIAL PUBLICATIONS OF THE

Prior to November 30, i 973 . publications of the Natural History Museum have appeared under various titles — Leaflet
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Number 308
March 16, 1979

CONTRIBUTIONS IN SCIENCE

NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY

A NEW SALAMANDER, GENUS BATRACHOSEPS,
FROM THE INYO MOUNTAINS OF CALIFORNIA,
WITH A DISCUSSION OF RELATIONSHIPS IN THE GENUS

By Ronald William Marlow, John M. Erode, and David B. Wake

Published by the NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY • 900 EXPOSITION BOULEVARD • LOS ANGELES, CALIFORNIA 90007

ISSN: 0459-8113

Suggested Citation: Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 308:1-17.

Contributions in Science are articles in the earth and life sciences, presenting results of original research in Natural
History. Science Bulletin (Numbers 1-30; 28 June 1965 to 10 April 1978) and Contributions in Science (Numbers 1-300;
23 January 1957 to 16 June 1978) were merged into a single imperial octavo serial beginning with Number 301.

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 308:1-17.

A NEW SALAMANDER, GENUS BATRACHOSEPS,
FROM THE INYO MOUNTAINS OF CALIFORNIA,
WITH A DISCUSSION OF RELATIONSHIPS IN THE GENUS1 *

By Ronald William Marlow" John M. Erode,3 and David B. Wake 4

Abstract: A newly discovered species of plethodontid salamander from the Inyo Mountains in the north-
ern Mojave Desert, California, is described as Botrachoseps campi. This new species helps bridge the gap
between Botrachoseps and members of the tropical supergenus Bolitoglossa, and may well be the most general-
ized of all bolitoglossine salamanders. Botrachoseps campi and B. aridus are the only plethodontid salamanders
known to be restricted to localized mesic microhabitats surrounded by desert. The new species is very general-
ized in morphology. In some respects it resembles B. wrighti of Oregon, but it also shares some features with
B. aridus. On the basis of its distinctive morphology and distribution, B. campi is thought to be a survivor of
an early stage in the evolution of the genus. The Inyo Mountains region may have served as the habitat of the
species throughout much of the Tertiary. Recommendations are made for preservation of the fragile habitats in
which the species lives.

In 1970 Brame described the first terrestrial salamander
endemic to desert habitats. Botrachoseps aridus occurs in a
highly restricted, mesic microhabitat in the desert of southern
California. We now announce the discovery of a second species
of lungless salamanders, family Plethodontidae, known from
desert regions. The new species was discovered in the fall of
1973 in two canyons along the west slope of the Inyo Mountains
at the northern edge of the Mojave Desert near the Owens Val-
ley, California. This strikingly distinct form is so different from
other Botrachoseps in its robustness and coloration that we ini-
tially thought it to be a species of Hydromantes, or even of a new
genus. Upon further examination it proved to be a structurally
generalized member of the genus Botrachoseps, very similar to a
postulated ancestor of the entire genus and not much removed
from the ancestral stock of the tribe Bolitoglossini (including
Hydromantes and all of the neotropical plethodontid salamanders,
in addition to Botrachoseps). Like B. aridus, the species seems
to occur only in restricted, mesic microhabitats in a setting that is
seemingly harsh and inhospitable for salamanders. Yet, the
apparent absence of any populations even remotely similar to this
species in areas more favorable for salamanders suggests that the
Inyo Mountains have long been the home of the species. These
mountains are relatively ancient, and in fact may represent rem-
nants of the original crest of the Sierra Nevada before the modern
mountains were uplifted to the west (Bateman and Wahrhaftig
1966). Possibly these salamanders have been in the Inyo Moun-
tains throughout Cenozoic times.

The new species is named in honor of the late Charles L.
Camp, herpetologist, paleontologist, and historian, who con-
tributed greatly to our knowledge of the American West. Profes-
sor Camp discovered the genus Hydromantes in the New World,
described H. platycephalus, participated in the discovery and
description of H. shastae, and very early recognized the diversi-

fication within the genus Botrachoseps in his description of B.
major. The species shall be known as:

Botrachoseps campi new species

Figures 1, 6, 7, 8, 11

Suggested common name: Inyo Mountains Salamander.

HOLOTYPE: MVZ 122993, an adult female from Long John
Canyon, W slope of the Inyo Mountains, elevation 1695 m (5560
ft), 3.2 km (2 mi) (airline) N, 5.3 km (3.3 mi) E Lone Pine, Inyo
County, California, USA; collected by John M. Brode and Ron-
ald William Marlow on September 26, 1973.

PARATYPES: MVZ 122994-122996 (3 specimens), same
data as the holotype; MVZ 123011-123015 (5 specimens), same
locality as holotype, collected October 20, 1973; MVZ 122997 -
123009, 123017 - 123031 (28 specimens), LACM 122004 -
122011 (8 specimens), French Spring, W slope of the Inyo
Mountains, elevation 1829 m (6000 ft), 6.4 km (4 mi) N, 5.7
km (3.6 mi) E Lone Pine, Inyo County, California.

'Review Committee for this Contribution
Robert L. Bezy
Arden H. Brame
James F. Lynch
Jay M. Savage
John W. Wright

-’Museum of Vertebrate Zoology, University of California, Berkeley,
California 94720.

3Department of Fish and Game, 987 Jedsmith Drive, Sacramento, Cali-
fornia 95819.

4Research Associate in Herpetology, Natural History Museum of Los
Angeles County, Los Angeles, California 90007.

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 308:1-17.

Marlow, Brode, Wake: New Salamander (Batrachoseps)

Figure 1. Batrachoseps campi (MVZ 123023), an adult female (40.4 SL) from Long John Canyon, Inyo Mountains, Inyo County, California. Photograph
by James Hendel, Scientific Photography Laboratory, University of California, Berkeley. Scale is 25 mm.

DIAGNOSIS: A large, robust species of Batrachoseps (indi-
viduals identifiable to sex, 16 males: 41.3-53.3, mean 45.3 SL5;
23 females: 32.0-60.7, mean 45.5 SL) with a short tail, broad
head and large feet; distinguished from all members of the genus
by these characters and by its distinctive coloration (dark
brown-black ground color with moderate to large patches of light
gray iridophores on upper eyelids and irregularly spread over
head and anterior part of trunk, and lacking a dorsal stripe or
band that is lighter than the ground color); distinguished from B.
aridus by its much larger size (largest known B. aridus are less
than 40 mm SL), relatively shorter tail, and fewer vertebrae
(19-20 versus 17-19); further distinguished from species which
approach it in adult size by its much broader head, fewer trank
vertebrae (except B. wrighti) and much shorter tail. Batrachoseps
campi is the only member of the genus that has paired prefrontal
bones and divided premaxillary bones from early juvenile stages
throughout adult life.

DESCRIPTION: Batrachoseps campi is the most generalized
member of the genus in its morphology. The species is relatively
large and has a large, broadly rounded snout. Nostrils are small
and typical of the genus. There are no protuberances associated
with the distinct nasolabial grooves. No mental hedonic glands
have been observed in males. The head is very broad (SL
6.6-7. 1, mean 6.8 times head width in 6 males and 6. 3-7. 5,
mean 7.0 in females over 45 SL). Grooving patterns of head,
throat and neck are typical of the genus. Eyes are relatively large
and moderately protuberant. Vomerine teeth are borne in long
series (mean total 16.5); frequently the teeth extend nearly to the
lateral edge of the internal nares. Small maxillary teeth extend
beyond the center of the eyeball; they increase in number with
increasing size and are relatively numerous (mean total 46). Pre-
maxillary teeth are small and numerous (8-10 in adult males;
6-17 in adult females); they are in an advanced position in the
males, but barely protrude through the lip. Costal grooves
between appressed limbs (counting one each in axilla and groin)
range from 16- 18 with a strong mode of 17. The tail is short for
this genus and tapered; tail length in 20 adults over 45 SL is
0.62-0.88, mean 0.76 (± 0.06) times SL, with no discernible
sexual dimorphism. The tail has no basal constriction, and is

nearly round. However, an irregular row of enlarged glands is
present on the dorsal surface of the tail. These glands form
pustule-like protuberances that are more lightly colored than sur-
rounding surfaces. The postiliac gland is a small, grayish spot
that is not conspicuous. Limbs are long for the genus, and leave
from 2-5 intercostal folds uncovered when appressed (mean 3.66
± 0.89; n = 47). Standard length ranges from 4. 2-5. 3 (mean
4.8 ± 0.3) times hind limb length in 16 females, and from
4.5-5. 1 (mean 4.8 ± 0.25) in males. The hands and feet are
large for this genus. Webbing is insignificant, and the four fin-
gers and four toes are all discrete. The third digit of both manus
and pes is far longer than the others. All digits have bulbous tips
and large subdigital pads. Fingers in order of decreasing length:
3, 2, 4, 1; toes in order of decreasing length: 3, 2, 4, 1.

MEASUREMENTS (OF HOLOTYPE): Head width, 8.4;
snout to gular fold (head length), 12.8; head depth at posterior
angle of jaw, 2.9; eyelid length, 3.3; eyelid width, 2.0; anterior
rim of orbit to snout, 1.7; horizontal orbital diameter, 1.9; inter-
orbital distance, 3.8; distance between vomerine teeth and para-
spheroid tooth patch, 0.2; snout to fore limb, 5.7; distance sepa-
rating internal nares, 2.4; distance separating external nares, 2.6;
snout projection beyond mandible, 0.9; snout to posterior angle
of vent (standard length), 60.7; snout to anterior angle of vent,
56.1; axilla to groin length, 32.5; tail length, 48.5; tail width at
base, 4.3; tail depth at base, 3.7; fore limb length, 11.1; hind
limb length, 12.1; width of right hand, 3.4; width of right foot,
4.0.

COLORATION: This is a dark salamander that differs from all
other Batrachoseps in lacking a dorsal stripe at all ages. The
ground color is very dark brown-black in most individuals, but a
lighter brown-black in some. This is the result of a dense network
of melanophores, covering the whole body with few or no gaps.
The only additional color is a scattering of silvery iridophores
which when grouped together form patches with a slightly green-
ish hue. Patches of iridophores are concentrated on the upper

5SL = standard length, the distance from the snout to the posterior angle
of the vent; all measurements in this paper are in millimeters, unless
stated otherwise.

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 308:1-17.

Marlow, Brode, Wake: New Salamander ( Batrachoseps )

Figure 2. Type locality of Batrachoseps campi. Long John Canyon, Inyo Mountains, Inyo County, California. Camera is aimed southeast. Salamanders
were found inside the mouth of the canyon, under surface objects in the shaded area.

eyelids of most individuals, and in a series of spots of varying
size lying mainly on the posterior part of the head, the neck, and
the dorsum in the vicinity of the insertion of the hind limbs.
Elsewhere, including the nearly immaculate venter, the irido-
phores are scarce and small. These cells fade and disappear after
a short time in alcohol. The dorsal glands of the tail are prom-
inent near the tail base in many of the individuals. They appear to
be brown-tan in color, the result of a thinning of the ground
color. The iris is dark and featureless.

REMARKS: There is little sexual dimorphism in this species.
Mean SL of males and females in the sample is similar, but
females appear to reach larger sizes than males. Females have
vertically folded vent margins, whereas males have dense, short
papillae in the anterior half of the vent.

Some large females in the sample have enlarged, yellow
ovarian eggs. Testes of most adult males are large and unpig-
mented. Vasa deferentia are heavily pigmented in most adult
males. The two largest males (50.4, 53.3) have bilobed testes.
The smallest male with bilobed testes is 45.0 SL, and the largest
male with unilobed testes is 47.4 SL. Males 43.3 and 49.0 SL
have a second lobe forming, and a male 48.6 SL has two lobes.

When uncovered, individuals of B. campi displayed no special
defensive behavior of the kind observed in other species of the
genus (Stebbins and Lowe, 1947; Brame, Long, and Chiri,
1973). Apparently immobility is the sole defensive behavior.

HABITAT: The western slopes of the Inyo Mountains receive
approximately 22 cm of precipitation each year, much of it com-
ing in snowfall. Vegetation on the slopes is sparse and stunted
with extensive areas of open, rocky substrate. Long John Canyon
is narrow and winding with rock walls rising almost vertically for
about 150 m. At the type locality the canyon axis is east- west
and the canyon floor is about 10 m wide (Fig. 2). The floor of
the canyon is strewn with boulders and flood-carried rubble. A
permanent spring at the type locality provides enough moisture
for a dense, nearly impenetrable growth of woody shrubs (aver-
age height about 2 m) filling the canyon floor and extending
approximately 100 m. This appears to be the extent of the avail-
able surface habitat, though crevices and subterranean water
courses may provide extensive underground habitat.

Salamanders were found under rocks in fine, dark soil (a sandy
loam) that was wet enough that a cast formed in the hand with-
stood taps with the finger. As many as four animals were taken
under a single large rock (25 x 40 cm). This is a heavily shaded
canyon. On October 20 the sun shone on the canyon bottom at
10:30, but by noon the canyon was entirely shaded.

The flora of the canyon floor is dominated by dense stands of
Salix sp., Forestiera neomexicana and Rosa Woodsii. Also pres-
ent were Erodium sp., Artemisia Dracunculus, A. ludoviciana ,
Clematis ligusticifolia, Aquilegia formosa, Gutierrezia micro-
cephala, Philadelphus microphyllus, Chrysothamnus teretifolius

Contrib. Sci. Natur. Hist. Mus. Los Angeles County . 1979. 308:1-17.

Marlow, Brode, Wake: New Salamander (Batrachoseps)

Table 1. Classification

matrix

based i

on discriminant analysis

of 229 specimens of

nine

species of

Batrachoseps, including juveniles and adults of both sexes. Entries along the main diagonal are
matches of individuals to their source populations based on twelve morphological characters.

Number of Individuals Classified into Group

“correct”

Group

I aridus

2 stebbinsi

3 simatus

4 attenuatus

5 major

6 campi

8 wrighti

9 pacificus

and Haplopappus cuneatus.

The second locality, French Spring, is 6.4 km (4 mi) (airline)
north and 5.7 km (3.6 mi) (airline) east of Lone Pine, Inyo Co.,
California, at an elevation of about 1800 m. The water flow of
this spring is much more extensive than in Long John Canyon.
The flow from the spring cascades down approximately 300 m of
naturally terraced rocky hillside and approximately 300 m of
canyon floor, before disappearing in the sand. Vegetation is simi-
lar to that at the type locality (Fig. 3).

COMPARISONS: Both bivariate and multivariate morpho-
metric analyses have been used to compare B. campi with other
described species of Batrachoseps. For these comparisons the fol-
lowing juvenile and adult specimens were used (all deposited in
LACM and MVZ, catalogue numbers and measurements on fde
at MVZ): 14 B. aridas (type locality); 30 B. stebbinsi (type
series); 29 B. simatus (type series); 30 B. attenuatus (San Fran-
cisco, Calif.); 30 B. major (vicinity of Los Angeles, Calif); 46 B.
campi (type series); 20 B. wrighti (central Oregon); 30 B. pacifi-
cus (Santa Cruz Island). Only B. relictus, a highly variable
species currently under study by Kay Yanev, was omitted. The
latter species is small and slender, and is most similar in mor-
phology to B. attenuatus. Data for the following characters were
used: SL, head width, hind limb length, front limb length, limb
interval, total maxillary teeth, total vomerine teeth, tail length,
front foot width, hind foot width, head length, number of trunk
vertebrae. Bivariate plots of head width, limb length and foot
width against standard length were produced for comparison with
the results of Brame and Murray (1968), and tooth number and
tail length were also plotted against standard length. The results
of the bivariate analysis indicated that most characters distinguish
B. campi from some other species and some characters (head
width. Fig. 4) distinguish B. campi from nearly all other Batra-
choseps (compare with Fig. 9, Brame and Murray 1968). The
species differ greatly in adult size, thus making direct comparison
of individual features and even bivariate analysis difficult. How-
ever, the species of Batrachoseps are readily distinguished by
combinations of three characters or more, and we believe multi-
variate discrimination is the method of choice. Canonical variate
analysis was chosen to discriminate between the species. In par-
ticular, we wished to determine how distinct if. campi was rela-
tive to species it resembled most closely, based on bivariate
analysis. The University of California, Berkeley, version of the
BMD07M program for stepwise discriminate analysis was used
(see Lynch and Wake 1975, for description).

The main result of the discriminate analysis is the demonstra-

tion of the distinctiveness of B. campi relative to other species of
Batrachoseps. Even among the remaining species of the genus
there is a moderate amount of discrimination, but we emphasize
that we are not considering geographic variation here. A feature
of the program used is a classification matrix which gives the
number of "correct” and "incorrect" assignments of individuals
to their proper populations on the basis of morphological criteria
alone (see Lynch and Wake 1975). The classification matrix
(Table 1) assigned all 46 specimens of B. campi to the correct
population on morphological grounds. There are from one (B.
pacificus, B. aridus, B. wrighti) to six (B. simatus) mis-classifi-
cations in the other species, but overall 92 per cent of the classi-
fications are correct. The first three canonical variables account for
93 per cent of the observed interspecific variance. The first two
canonical axes are illustrated in Figure 5.

Along the first axis (accounting for 69 per cent of the interspe-
cific variance) species are separated mainly by factors related to
size (standard length, head width). Limb interval and tail length
also contribute importantly. B. campi is well separated from all
other species. B. aridus. B. stebbinsi. B. wrighti and B. pacificus
are relatively close to B. campi along this axis; B. attenuatus, B.
major, and B. simatus are relatively remote.

The second axis (accounting for 16 per cent of the total inter-
specific variance) separates B. wrighti from all other species, but
seems to add little to discrimination among remaining species.
Based on the magnitude of standardized coefficients of the canon-
ical variables, head width, numbers of trunk vertebrae and foot
width contribute most to discrimination along this axis.

The third axis accounts for only about 8 per cent of the total
interspecific variance. It allows separation of B. stebbinsi andfl.
simatus from all other species, with B. aridus being on the
extreme end of the axis from B. stebbinsi. Other axes contribute
little additional discrimination.

COMPARATIVE OSTEOLOGY: Osteological comparisons to
other Batrachoseps are based on study of an adult female (49.4
SL) and a juvenile female (37.6 SL) of B. campi. These were
cleared and stained using standard techniques. Radiographs of the
type series and both cleared and stained specimens and radio-
graphs of all other species of the genus were utilized.

Batrachoseps campi is similar to other species of Batrachoseps
in some features of skull structure, but it is highly distinctive in
others. The skull of B. campi is typical of the genus in propor-
tions (Fig. 6). The characteristic dorsal fontanelle between the
paired frontals and parietals is especially prominent. The fron-
tanelle is proportionately of similar size in all species of Batra-

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 308:1-17.

Marlow, Brode, Wake: New Salamander (Batrachoseps)

choseps, with a slightly smaller fontanelle in B. wrighti than in
other species. Only Batrachoseps among all genera of salaman-
ders has a fontanelle of this shape. In all Batrachoseps the skull
is relatively short and broad; total skull width is 70 to 75 per cent
of skull length, and B. campi falls about in the middle of the
range of measurements.

The relatively short maxillary bones seen in B. campi are
widespread in Batrachoseps . Characteristically the frontals are
narrowed, appearing “pinched” between the orbits, and this is
seen in B. campi. In all species the skull is lightly built and rather
weakly articulated. However, B. campi has notably stronger and
more extensive cranial articulations than other species.

Premaxillary bones of B. campi are separated in both juvenile
and adult specimens. In most of the radiographed individuals,
including juveniles, the divided premaxillaries can be seen
clearly. In some individuals the appropriate region of the skull is
obscure on the radiographs. Divided premaxillary bones are
found elsewhere in the genus only in large adult B. wrighti
(Bishop 1937, Stebbins and Lowe 1949, Wake 1966). Smaller B.
campi than those available to us (28.5 SL) probably have a single
premaxillary bone, the typical plethodontid situation; fused larval
and juvenile premaxillaries separate at metamorphosis or early in
life in the primitive plethodontid condition, but remain fused in a
variety of derived conditions (Wake 1966).

In B. pacificus, B. simatus, B. stebbinsi. B. major, B. aridus
and some populations of B . attenuatus and B. relictus , the frontal
processes of the premaxillary arise separately, then fuse for a
distance before again separating near their tips. We see no evi-
dence of such fusion in either B. wrighti or B. campi. The poste-
rior tips of the frontal processes diverge greatly in B. campi, to a
lesser degree in B. wrighti and only slightly in other species. As
in all species of Batrachoseps the premaxillaries of males are
larger and more protuberant than those of females, but sexual
dimorphism is apparently less marked than in such species as B.
pacificus and B. major, and probably less than in the other
species as well.

The maxillary bones are not well articulated with other bones
in Batrachoseps, but in B. campi the elements are relatively
stouter than in any other species of the genus. The facial lobe of
the maxillary in B. campi is relatively large, equalling the propor-
tions in such large species as B. pacificus. Palatal portions of the
maxillaries are very small. The large nasal bones are the domi-
nant facial elements. They have a narrow spinous projection par-
alleling the frontal processes of the premaxillaries in B. campi
and in other Batrachoseps, but these are especially pronounced in
B. campi. Rather well developed septomaxillaries are present in
all species of Batrachoseps.

Prefrontal bones have previously been reported only in B.
wrighti, in which individuals over 45 mm body size have very
tiny but discrete elements (Wake 1966). Well developed pre-
frontals are present in both the juvenile and adult B. campi, and,
while the radiographs are not suitable for finding these elements
in all instances, they can be seen in a number of the X-rayed
specimens. The bones are about the size of the facial lobes of the
maxillaries and are distinctly larger than the septomaxillaries. In
contrast, the prefrontals which develop in large B. wrighti remain
much smaller than the septomaxillaries.

Preorbital processes of the vomers are typically absent in Bat-
rachoseps. In B. campi small processes that extend to the lateral
margin of the internal nares (adult) or to the center of the nares
(juvenile) are evident. These tooth-bearing processes are prom-
inent in the radiographs. In other species of Batrachoseps the

Figure 3. French Spring, Inyo Mountains, Inyo County, California.
Camera is aimed southeast. Salamanders were found under rocks along
the stream course in the center of the photograph.

Table 2. Quantitative characteristics of the chromosomes of
Batrachoseps campi based on haploid set of Figure 12.

Chromosome

No.

Relative

lengthJ

Arm

.■ b
ratio

Centromere

index^

11.0

1.05

10.0

1.28

9.8

1.05

9.0

1.21

8.8

1.10

8.4

3.35

7.8

1.00

7.1

1.04

6.0

1.38

6.0

1.06

5.8

1.31

5.7

1.14

4.8

2.13

a Length of chromosome/total length of haploid genome x 100.
b Length of long arm/length of short arm.
c Length of short arm x 100/length of whole chromosome.

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 308:1-17.

Marlow, Brode, Wake: New Salamander (Batrachoseps)

6 -

Batrachoseps
x aridus
• campi
a stebbinsi
□ wrighti
o pacificus

••

x ** *

* • X*

□

D □ □

□ o dm

Q O A A

• •

•

• •

•

• •

• o

•• o

•

• • •

o o

•

O A

• O O O o

O a

•

o o

□

O A A

A O

□ o

□

O O A

□

A A
A DA
A

4 -

A A

“I 1 1 1 1 1 1

30 40 50 60

Figure 4. Relation of head width to standard length in the five species of Batrachoseps having the broadest heads in the genus.

processes are usually mere suggestions of bone growth behind the
nares (Fig. 7), but there are two exceptions. In a radiograph of
one very large B. wrighti (reported by Brame 1964) a small pro-
cess is clearly evident, and very small processes are evident in a
radiograph of one adult B. aridus.

The nasolacrimal duct extends anteroventrally from the corner
of the eye to the cartilaginous nasal capsule. The opening in the
nasal capsule lies in the gap between the facial process of the
maxillary and the nasal. Typically, slight evacuations in the lat-
eral margins of the nasals, or more rarely in the anterodorsal
margin of the maxillary, mark the route of the duct in species of
Batrachoseps. In B. campi the duct produces slight dorsal con-
cavities in the anterior part of the prefrontals as well.

Frontal bones in Batrachoseps contribute importantly to the
facial part of the skull, especially by means of a long attenuated
process that extends anteriorly below the frontal processes of the
premaxillary. In B. campi the facial portion of the frontal is rela-
tively less developed than in other species and there is no spinous
process. Further, only the tips of the premaxillaries overlap the
frontals. In other respects, the frontals inB. campi are similar to
those in other Batrachoseps. with the characteristic posterior
divergence of the bones. B. wrighti has relatively the widest
frontals. Frontals are about one-half the total skull length in B.
campi, B. wrighti and B. pacificus, but on the order of two-thirds

the total skull length in B. stebbinsi, B. aridus, and the other
species.

All species have small parietal bones, with no clear differentia-
tion among the species in terms of relative size. The bones
usually fall short of the narrow occipital bridge portion of the
skull. In all species a well defined lateral spur is present on the
parietal (see Wake 1966), and it is particularly stout in B. campi.

Squamosal and quadrate bones are small and have no special
features in any single species. Dentaries are slender and similar
in structure in all species. Prearticulars have low coronoid pro-
cesses. All species have columellar processes extending from the
opercular plate. These processes are well developed inB. campi.

Teeth of Batrachoseps typically occur on premaxillary, maxil-
lary, dentary, and vomerine elements, and in addition there is a
paired patch of teeth along the parasphenoid. The patches vary
greatly within and between species in size and number of teeth.

The hyobranchial apparatus is typical of the genus Batracho-
seps in all important respects (see Lombard and Wake 1977)
including the long, recurved radii, the absence of a lingual car-
tilage, the basibranchial of relatively uniform width, and the rela-
tively long epibranchial. The genioglossal muscles are slender
and elongate. These muscles have a unique arrangement in Bat-
rachoseps, originating from the mandible near the jaw articula-
tion and proceeding well forward before extending to the midline

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 308:1-17.

Marlow, Brode, Wake: New Salamander (Batrachoseps)

r~ — ' — — i — i — i i ” — T" — ’i i———h

-8 -6 -4 -2 0 2 4 6 8

First Canonical Axis

Figure 5. Plot of first two canonical variates for adults of eight species of Batrachoseps. The discriminant analysis includes 12 morphological variables.
The first variate (abscissa) accounts for 69 per cent of the observed interspecific variance. The cross marks the centroid for each species. All individuals for
each species are enclosed within the indicated perimeter except for B. campi, where all individuals are spotted. 1. B aridus, 2. B. stebbinsi, 3. B.
simatus, 4. B. attenuatus, 5. B. major, 6. B. pacificus, 7. B. wrighti.

and dorsally into the tongue (Piatt 1935).

Trunk vertebrae in Batrachoseps number from 16 to 23. but any
single population has a maximal range of variation of four. B.
wrighti, the species with the shortest trunk, has a modal number of
17. Some populations of B. relictus also have a modal number of 17,
but others have modal numbers as high as 21. In B. campi there is a
strong mode of 18, with a range of 17-19 (mean = 17.9 ± .42 SD).
All individuals from Long John Canyon have 18 trunk vertebrae. B.
aridus has a strong mode of 19, with a range of 19-20 (Brame,
1970). The remaining species all have higher numbers of trunk
vertebrae, with both B major and B. attentuatus having modes as
high as 22, and occasional individuals of these species and B.
relictus having as many as 23.

Genera of plethodontid salamanders have either two or three

caudosacral vertebrae, with the exception of Batrachoseps, in
which both numbers may occur in a single population (Wake
1966). B. campi is typical of the genus, some individuals having
two and others, three.

Complete tails are present in 32 of the type series of B. campi.
In these individuals postsacral vertebrae (including either 2 or 3
caudosacral and all of the caudals) range from 22-34 (mean
28.44 ± 2. 18), with numbers increasing from juveniles to adults.
This is a very low number for Batrachoseps, but accurate com-
parative data are not available for other species. The main prob-
lem in obtaining accurate counts is the high frequency of tail
breaks. However, numbers as high as 60 caudal vertebrae occur
in some of the species. Relatively few caudal vertebrae might
also be expected in the relatively short-tailed B. aridus, but our

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 308:1-17.

Marlow, Brode, Wake: New Salamander (Batrachoseps)

B. w right i

B. arid us

Figure 6. Dorsal views of the skulls of four species of Batrachoseps. The lines indicate the divided premaxillary bones and the prefrontal bones
only in B. campi and B. wrighti.

found

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 308:1-17.

Marlow, Erode, Wake: New Salamander (Batrachoseps)

1 1

B cam pi

B. stebbinsi

B. pad ficus

Figure 7. Ventral view of the anterior part of the skull of four species of Batrachoseps. The bones are the premaxillaries, maxillaries, and vomers. Teeth
not indicated. Same scale as Figure 6.

largest cleared specimen (41.0 SL) has 32 (a total of 34 post-
sacrals) and larger specimens reasonably could be expected to
have more. In B. wrighti, also relatively short-tailed, as many as
38 caudal vertebrae are present.

The limbs of B. campi are stouter than those of most other
Batrachoseps but are of about the same relative length as those of
B. stebbinsi. In osteological detail they are very similar to those
of the more generalized species of Batrachoseps (Fig. 8). The
tibial spur is well developed in B. campi. This structure is absent
in many populations of B. attenuatus, but is characteristically
present in other species. Wake (1966) reported that all species of
Batrachoseps except B. wrighti and some populations of B.
paciftcus (these particular populations are now referred to B.
major, following Brame and Murray 1968) have seven rather
than the primitive plethodontid number of eight carpal elements
as the result of fusion of the ulnare and intermedium cartilages.
We have found eight carpals in some B. major and in most
specimens of B. wrighti, but other members of these species have
seven. All populations of Batrachoseps have eight tarsal ele-
ments. B. campi has seven carpals and eight tarsals. The usual
phalangeal formula in Batrachoseps is 1-2- 3- 2 for both hand
and foot, but Wake (1966) and Brame (1970) report 0-2- 3- 2
rarely in B. attentuatus. Brame and Murray (1968) and Brame
(1970) also indicate that B. stebbinsi and B. aridus differ from

other species in lacking a phalanx on the first digit of the hand
and foot. A small terminal phalanx is present on the first digit
of the hand in four of five specimens examined of B. aridus, and
on the first digit of the foot in two of these specimens. Extremely
small phalanges that do not appear on radiographs are present on
the first digit of both hands and feet in all nine specimens of
cleared B. stebbinsi available to us. These are often only dots of
mineralized tissue, and in one foot in each of two specimens no
sign of a phalanx is seen. Terminal phalanges of the first digit are
also greatly reduced in B simatus, and in one foot of one indi-
vidual (of a total of 13) this phalanx is absent. In B. campi the
hands and feet are relatively large and stout. The generalized pha-
langeal formula is present. The first phalanx is well developed
and discrete. Phalanges are especially well ossified with rela-
tively little cartilage, especially in large adults (Fig. 8). Expan-
sion of the most terminal phalanx of each digit is characteristic of
all species of Batrachoseps, including B. campi.

EVOLUTIONARY RELATIONSHIPS: Those of us who discov-
ered B. campi were impressed immediately with its robustness
and its relatively short body and tail. We thought it might be a
representative of Hydromantes or some other genus, and did not
think of assigning the new species to Batrachoseps until we
noticed that it had but four toes. The species is hardly a typical
Batrachoseps, as that taxon has been understood, and its discov-

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 308:1-17.

Marlow, Brode, Wake: New Salamander (Batrachoseps)

B. stebbinsi

Figure 8. Feet of four species of Batrachoseps, drawn from cleared and stained specimens with aid of microprojector. Cartilage stippled.

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 308:1-17.

Marlow, Brode, Wake: New Salamander (Batrcichoseps)

ery forces us to re-evaluate the status of the genus.

In several morphological features B. campi approaches
hypothesized ancestral states for the entire tribe Bolitoglossini
(Batrachoseps, Hydromantes , and the tropical salamander
groups in the supergenus Bolitoglossa-Pseudoeurycea, Chirop-
terotriton, Oedipina, Thorius, Lineatriton, Parvimolge, and
Bolitoglossa). The species is more generalized than other Batra-
choseps in osteology and myology as well as in external form.
Features in which it retains ancestral conditions as compared to
existing conditions in Hydromantes include presence of pre-
frontals and of genioglossal muscles. It matches Hydromantes in
having paired premaxillaries. The paired premaxillaries distin-
guish it from all described species of the supergenus Bolitoglos-
sa. Only the absence of a fifth toe (including tarsal, metatarsal
and phalanges) and the presence of a fontanelle in the skull roof
remove B. campi from the presumed ancestral state for the tribe
Bolitoglossini (for comparative data see Wake 1966).

Perhaps B. campi is a very early derivative of the ancestral
stock of all boiitoglossines, representing a lineage that appeared
shortly after the differentiation of the Batrachoseps stock from a
more generalized ancestral group. The superficial similarity of B.
campi to members of such genera as Hydromantes, Chirop-
terotriton, and Pseudoeurycea may reflect the great conservatism
inherent in these relatively ancient lineages. Nevertheless B.
campi can rather easily be accommodated in the genus Batracho-
seps so long as B. wrighti is also included. The close relationship
of B. campi to other species of Batrachoseps rather than to other
boiitoglossines is evident, based on the number and nature of
derived features that are shared. It is in ancestral features that B.
campi resembles members of other genera.

The discovery of B. campi and knowledge of its structure tend
to make the tribe Bolitoglossini a somewhat tighter unit than
formerly perceived. However, no new insight is provided into
relationships of Batrachoseps as a unit relative to Hydromantes
and the supergenus Bolitoglossa. There is no reason to modify
the conclusions of Wake (1966) that Batrachoseps is cladistically
closer to the supergenus Bolitoglossa than to Hydromantes. The
single most important piece of evidence is the derived karyotype
of Batrachoseps and the supergenus Bolitoglossa, both groups
having a haploid chromosome number of 13 as opposed to the 14
chromosomes characteristic of Hydromantes and all other pletho-
dontids (Kezer 1964; Kezer and Macgregor 1971; Morescalchi
1975).

Certain specialized and presumably derived morphological
conditions are shared by B. campi with all other members of the
genus Batrachoseps. Accordingly we conclude that the species is
most appropriately associated with Batrachoseps. These features
include: (1) the hyobranchial apparatus and musculature, specifi-
cally a genioglossus muscle that takes its origin near the posterior
end of the mandible. This arrangement of the muscle is unique to
Batrachoseps (Piatt 1935; Wake 1966; Lombard and Wake
1977). Further, in all other features of the apparatus the species is
like other Batrachoseps rather than like any other genus. (2) A
distinct fontanelle is present in the skull roof, bordered by
markedly narrowed frontals and parietals. No comparable fon-
tanelle is found in any other salamander. Thorius has a smaller
fontanelle involving narrowed frontals but its parietals have pro-
portions typical of less specialized genera.

Some derived features occur in B. campi and other species of
Batrachoseps which either are less distinctive than those men-
tioned above, or are not exclusively present in Batrachoseps.

These add strength to the case for inclusion of B. campi in the
genus. The characters include: ( 1 ) A strongly developed lateral
spur on the parietal bone. This spur is shaped like that in other
Batrachoseps, but spurs of a somewhat different shape are also
present in the supergenus Bolitoglossa (Wake 1966). (2) The
number of caudosabral vertebrae may be either two or three in a
given population. Typically plethodontid species have either two
or three vertebrae in this region, and only in Batrachoseps does
intrapopulational and intraspecific variation occur (Wake 1966).
(3) Four digits present on the hind foot. All other plethodontids
have five digits on the hind foot, except Hemidactylium scutatum
and Eurycea quadridigitata, both distinctly different from Batra-
choseps in many features. (4) A haploid chromosome number of
13 (see Appendix).

Within the genus, B. campi is the most generalized species in
having superficial structure similar to that of other generalized
plethodontids and a number of osteological characters present in
ancestral states. These latter features include notably the paired
premaxillary bones, the prefrontal bones, and the preorbital pro-
cesses of the vomers. Only in large, presumably old B. wrighti
do these features appear in combination, and the first two charac-
ters are found only in B. campi and B. wrighti. All three charac-
ters are related to degree of morphogenesis (see Wake 1966, for a'
lengthy discussion of this issue). In plethodontids that have a
larval stage, all three features are associated with metamorphosis,
making their appearance at about that time. There is no meta-
morphosis in a technical sense in Batrachoseps and other sala-
manders with direct development, and the changes which ordinar-
ily occur during the relatively short metamorphic period are
spread out over the entire life span of these organisms. In most
species of Batrachoseps the metamorphic features listed above
are delayed in appearance to such an extent that the life span of
the organism is insufficiently long for them to appear. According
to this interpretation, metamorphosis of these structural features
occurs early in B. campi, late or not at all in B. wrighti, and
never in other species of Batrachoseps.

It can be argued that B. campi is primitive in respect to its
entire pattern of development (i.e. , it is the least paedomorphic),
and that one should not “count” three primitive states, but only
one. However one counts, B. campi seems to qualify as the most
primitive species. In addition to its primitive osteological charac-
ters, B. campi has external proportions that are more similar to
those of generalized plethodontids than are those of any of the
other species of Batrachoseps. Its broad head, long limbs, rela-
tively short trunk, and short tail are all generalized features.

It is unclear what the closest relatives of B campi might be.
Osteological and chromosomal comparisons suggest that B.
wrighti is the closest relative. In addition to the features dis-
cussed above, both species have relatively low numbers of trunk
vertebrae. Those features which separated, campi and B. wrighti
from other Batrachoseps are more striking than those which sep-
arate some other genera of salamanders within the family (Wake
1966). We considered placing B. wrighti and B. campi in the
resurrected genus Plethopsis (Bishop 1937; see also Stebbins and
Lowe 1949). However, Plethopsis would have to be founded on
the basis of a suite of primitive characters, and given the difficul-
ties in deciding whether we are dealing with one general meta-
morphic event or several independently evolving characters we
believe that it is advisable to keep the genus intact. In fact, there
are reasons for thinking that B. campi might be more closely
related to some other species than B. wrighti. B. aridus is much

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 308:1-17.

Marlow, Brode, Wake: New Salamander (Batrachoseps)

Figure 9. Map of eastern California showing the two localities of Batrachoseps campi. The closest locality for B. stebbinsi is indicated by an asterisk.
Deep Canyon, Riverside County, California, is the only other desert locality for plethodontid salamanders (B. aridus ) and it is indicated on the inset map.

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 308:1-17.

Marlow, Brode, Wake: New Salamander (Batrachoseps)

Figure 10. Aerial photograph of the two localities for Batrachoseps campi. Note the scant vegetation. FS, French Spring; LJC, Long John Canyon. Photo
courtesy of the USGS.

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 308:1-17

Marlow, Brode, Wake: New Salamander (Batrachoseps)

smaller than/?, campi, but in many proportions it is rather similar
(Fig. 4). Further, the color patterns in the two species are some-
what similar, and differ from those in all other species. Oste-
ology provides the only certain means of separating these two
species on other than size-related grounds. B. campi is matched
in some aspects of size by B. pacificus, but the species are highly
distinct osteologically. Further, B. pacificus has the characteristic
striped dorsal pattern that is seen in all species except B. aridus
and B. campi. Both B. stebbinsi and B. pacificus have propor-
tions more similar toB. campi than any species except B. aridus.

The presence of B. campi and B. aridus in remote and totally
isolated desert habitats raises questions concerning historical bio-
geography of this region. The animals survive in highly restricted
microhabitats surrounded by seemingly uninhabitable desert
(Figs. 9 and 10). Both species must utilize underground retreats.
Possibly they can travel for some distance through crevices and
solution channels in limestone formations. Brame (1970) reports
limestone at the type locality of B. aridus. While limestone out-
crops are not obvious in the two localities of B. campi, in parts of
the southern Inyo Mountains near these localities fossil-bearing
marine shales and limestones of Triassic age are found in sections
up to 1,800 ft (about 550 m) thick. These rocks are exposed in
many sites now and presumably they have been for long periods
in the past (Oakeshott 1971). The desert species of Batrachoseps
are unlikely to be Pleistocene relics, stranded in areas of favor-
able habitat which they reached during pluvial stages. More
likely, these species have been occupying habitats associated
with exposures of these ancient rocks for a long period, perhaps
since early Tertiary times. They may have maintained themselves
in favorable microhabitats through many cycles of heating and
cooling, moistening and drying. Had they entered desert habitats
during Pleistocene pluvial periods, they would have dispersed
from somewhere. There are no extant populations in California
that could have served as ancestors for either B. aridus or B.
campi, and it seems unlikely that these ancestors all would have
become extinct or given rise to such species as B. stebbinsi in the
relatively short time since even the beginning of Pleistocene.

Finally we note that salamanders recently have been collected
in a number of canyons in parts of the Inyo Mountains other than
those mentioned here. These populations are under intense study
currently.

PROTECTIVE MEASURES

Long John Canyon and French Spring are within the bound-
aries of National Resource Lands administered by the Bureau of
Land Management. Vehicle travel in the area currently is permit-
ted only on roads and trails established prior to November 1,
1973. Both springs are Public Water Reserves. Water cannot be
appropriated, and no camping is allowed within 150 m of the
water.

Better protection for these populations would be afforded if
Long John Canyon in the area of the spring were closed to vehi-
cles, and the habitat areas in both Long John Canyon and French
Spring were withdrawn from potential mining. Due to its small
range and limited habitat, B. campi qualifies as “rare” and
should be afforded protection under state law.

Feral horses and burros have caused massive destruction to the
vegetation and soils surrounding desert springs and water courses
in the Southwest. Burros are established in the Inyo Mountains
and the United States Fish and Wildlife Service and the Califor-

nia Resources Agency should assess their impact and take steps
to reduce or eliminate damage. Those springs in the area where
horses and burros presently occur should be carefully surveyed
for other populations of salamanders.

The California Resources Agency and other agencies should
re-evaluate the practice of opening and clearing desert springs,
often with explosives, for the enhancement of habitat for upland
game and other animals. The manipulation of fragile desert seep,
spring, and stream communities for any reason should proceed
only after rigorous investigation of the consequences of the an-
ticipated activities.

ACKNOWLEDGMENTS

We are grateful to Robert C. Stebbins for his advice and
review of the manuscript. Samuel S. Sweet and Kay P. Yanev
reviewed an early version of the manuscript, and James L. Patton
offered helpful criticism. Arden H. Brame, Jr. discussed
aspects of the research with us and presented information con-
cerning specimens collected by him. Materials from the Natural
History Museum, Los Angeles County, were kindly loaned by
John W. Wright and Robert L. Bezy. We thank James Kezer for
supplying information concerning his work on salamander
chromosomes, and for permitting us to report some of his obser-
vations. Gene M. Christman of the Museum of Vertebrate Zool-
ogy drafted the figures. Aspects of this work were supported by
the California Resources Agency, the Bureau of Land Manage-
ment, the Committee on Research of the University of California,
Berkeley, and the National Science Foundation (grant BMS
74-20922).

APPENDIX

Chromosomes of Batrachoseps campi

James Kezer has studied the chromosomes of a single B. campi
and has graciously provided the following information which he
has encouraged us to present as part of the description of this
species. For some years Prof. Kezer has been accumulating
information concerning the chromosomes of members of the
genus Batrachoseps, and comparative data derive from this work.

Chromosomes of B. campi were obtained from the testes of an
adult male. The testes were fixed in three parts absolute ethanol
and one part glacial acetic acid. Testes were squashed in 45 per
cent acetic acid and spermatocytes were examined for chromo-
somes. Unstained slides were photographed using phase-contrast
optics. Data were derived from anaphase cells of the second
meiotic division. Two haploid sets of chromosomes from one
such cell (Fig. 11) were the source of quantitative data (Table 2)
used to construct an ideogram (Fig. 12).

The haploid chromosome number of B. campi is 13. This is
the same number found in other species of Batrachoseps and in
the tropical salamanders of the supergenus Bolitoglossa (Kezer
and Macgregor 1971; Leon and Kezer 1978). All other pletho-
dontids that have been examined, including the remaining bolito-
glossine genus Hydromantes, have a haploid number of 14
chromosomes.

The karyotype of B. campi has two chromosomes having cen-
tromeres in subterminal regions (numbers 6 and 13, Fig. 12). In
B. attenuatus and B. major centromeres are all in median and
submedian regions, but B. wrighti has two chromosomes with

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 308:1-17.

Marlow, Brode, Wake: New Salamander (Batrachoseps)

Figure 11. Late anaphase II chromosome preparation of Batrachoseps
campi, prepared and photographed by James Kezer. Unstained squash
The chromosomes of this photomicrograph were used in preparing Table
2 and Figure 12.

Figure 12. Ideogram of haploid chromosome set of Batrachoseps campi
based on data in Table 2.

subterminal centromeres (see Kezer 1964, Fig. 7A.16). Informa-
tion is not available for other species of Batrachoseps. At present
not enough information is available to determine homologies of
salamander chromosomes. Should the two subterminal chromo-
somes of B. campi prove to be homologous with those of B.
wrighti, and should they represent derived conditions, the
hypothesized close relationship of the two species would gain
additional support.

LITERATURE CITED

Bateman, P.C. and C. Wahrhaftig. 1966. Geology of the
Sierra Nevada. Geology of northern California, Calif. Div.
Mines Bull. 190:107- 172.

Bishop, S.C. 1937. A remarkable new salamander from Oregon.
Herpetologica 1:92-95.

Brame, A.H., Jr 1964. Distribution of the Oregon slender
salamander, Batrachoseps wrighti (Bishop). Bull. So. Calif.
Acad. Sci. 63:165-170.

. 1970. A new species of Batrachoseps (slender sala-
mander) from the desert of southern California. Nat. Hist.
Mus. Los Angeles Co., Contrib. Sci. 200:1-11.

AND K.F. Murray. 1968. Three new slender salaman-
ders (Batrachoseps) with a discussion of relationships and
speciation within the genus. Nat. Hist. Mus. Los Angeles
Co., Bull. 4:1-35.

, M.C. Long and A. A. Chiri 1973. Defensive display

of the desert slender salamander. Herpeton, J. SW. Herpt.
Soc. 8:1-3.

Kezer, J. 1964. Meiosis in salamander spermatocytes, pp.
101-114 in Stahl, F.W. Mechanics of Inheritance.
Prentice-Hall, Inc., Englewood Cliffs, N.J. xii + 171 pp.

AND H.C. MACGREGOR 1971. A fresh look at meiosis

and centromeric heterochromatin in the red-back sala-
mander, Plethodon cinereus cinereus (Green). Chromosoma
(Berl.) 33:146-166.

Lombard, R.E. and David B. Wake. 1977. Tongue evolution
in the lungless salamanders, family Plethodontidae. II.
Function and evolutionary diversity. J. Morph. 153:39-80.
LeOn, P.E. and J. Kezer. 1978. Localization of 5S RNA genes
on chromosomes of plethodontid salamanders. Chromosoma
(Berl.), 65:213-230.

Lynch, J.F. and D.B. Wake. 1975. The systematics of the Chi-
ropterotriton bromeliacia group (Amphibia: Caudata), with
description of two new species from Guatemala. Nat. Hist.
Mus. Los Angeles Co., Contrib. Sci. 265:1-45.
Morescalchi, A. 1975. Chromosome evolution in the caudate
amphibia. Evolutionary Biology 8:339-387.

OAKESHOTT, G.B. 1971. California's Changing Landscapes.

McGraw-Hill Book Co., N.Y. xix + 388 pp.

Piatt, J. 1935. A comparative study of the hyobranchial appa-
ratus and throat musculature of the Plethodontidae. J.
Morph. 57:213-251.

Stebbins, R.C. and C.H. Lowe. Jr 1949. The systematic status
of Plethopsis with a discussion of speciation in the genus
Batrachoseps. Copeia 1949:116-129.

Wake, D.B. 1966. Comparative osteology and evolution of the
lungless salamanders, family Plethodontidae. Mem. So.
Calif. Acad. Sci. 4:1-111.

Accepted for publication October 17, 1977.

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 308:1-17.

Number 50.9
iii 16, 1979

I NATURAL HISTORY MUSEUM OF LOS ANGELES COUNT1!

NOTES FOR A REVISION OF THE ANT GENUS FORMICA.

. REIDENTIFICATIONS FOR SOME SPECIMENS FROM THE T. W. COOK
COLLECTION AND NEW DISTRIBUTION DATA
W (HYMENOPTERA: FORMICIDAE).

By Andre Francoeur and Roy R. Snelftng

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Pobllejhed by the NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY ■ TOO EXROSITI CN BOULEVARD * LOS ANGELES. OA UFO EM A 90007

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SERIAL PUBLICATIONS OF THE
NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY

Prior to November 30, 1973, publications of the Natural History Museum have appeared under various titles — Leaflet
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reviewers.

Number 309
March 16, 1979

CONTRIBUTIONS IN SCIENCE

NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY

NOTES FOR A REVISION OF THE ANT GENUS FORMICA.

2. REIDENTIFICATIONS FOR SOME SPECIMENS FROM THE T. W. COOK
COLLECTION AND NEW DISTRIBUTION DATA
(HYMENOPTERA: FORMICIDAE).

By Andre Francoeur and Roy R. Snelling

Published by the NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY • 900 EXPOSITION BOULEVARD • LOS ANGELES, CALIFORNIA 90007

ISSN: 0459-8113

Suggested Citation: Conlrib. Sci. Natur. Hist. Mas. Los Angeles Count}’. 1979. 309:1-7.

Contributions in Science are articles in the earth and life sciences, presenting results of original research in Natural
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23 January 1957 to 16 June 1978) were merged into a single imperial octavo serial beginning with Number 301.

Contrib. Sci. Natur. Hist. Mus . Los Angeles County. 1979.309:1—7.

NOTES FOR A REVISION OF THE ANT GENUS FORMICA.

2. REIDENTIFICATIONS FOR SOME SPECIMENS FROM THE T.W. COOK
COLLECTION AND NEW DISTRIBUTION DATA
(HYMENOPTERA: FORMICIDAE).1

By Andre Francoeur2
and Roy R. Snelling3

ABSTRACT: Specimens of the Formica neogagates and Formica fusca groups from the T.W. Cook collec-
tion are reidentified. New morphological variations are presented for F. longipilosa, together with a first
description of the male. Some of these data extend the geographical distribution of the less well known species
particularly in California.

“The Ants of California” was privately published by Thomas
Wrentmore Cook in 1953. This was Cook’s only published work on
ants and it does no credit to a many-faceted man. The book consists,
for the most part, of records and descriptive and biological notes
gleaned from previously published work of other authors. These
applied, for the most part, to California forms, but no effort was
made to correct records based on misidentified specimens. The
present paper reveals so many surprising misidentifications and
misinterpretations in Formica species or species groups, that it
becomes impossible to use any data from "The Ants of California”
without having the specimens at hand.

Furthermore, the drawings illustrating the book are misleading or
inaccurate: the head of Formica subnuda (p. 368) and the mesosoma
of F. occidua (p. 385) are erroneously proportioned; the "female”
head of F. subsericea (p. 368) is that of a male; the maxillary palpi of
F. sibylla (p. 392) are too short; the figure of F. subpolita cam-
ponoticeps (p. 392) is a worker of F. neoritfibarbis.

Numerous other deficiencies are pointed out by Brown and
Wilson (1953).

Cook died in 1962. In 1964 his collection was donated to the
Oakland Museum by his widow. The ants were transferred to the
Natural History Museum of Los Angeles County in 1971. There
were 4,397 pinned ant specimens and an undetermined number in
213 vials (Amaud and Wale 1973). Almost half of the pinned
material consisted of specimens from W. M. Wheeler’s collection
acquired during a brief stay at Harvard. Most of the remainder were
specimens from the collections of Stanford University, still bearing
the distinctive “L.S. Jr. U.” labels. The Stanford University
specimens were collected mostly by W.M. Mann. The specimens in
vials were, in large part, collected by Cook. Judging by the number
of specimens clearly collected by Cook and by the localities visited,
he had a very limited first hand acquaintance with the ants of
California.

The present note attempts first of all to clarify some of the
erroneous citations in "The Ants of California." We also cite new
distributional and morphological data, particularly on several

species in the F. fusca group described recently by the senior
author (Francoeur 1973). A few specimens, cited as various forms
of Formica, belong to other genera and are noted at the end of
this paper.

The following abbreviations are used: LSJU (Leland Stanford Jr.
University); TWC (T.W. Cook); WMM (W.M. Mann); WMW
(W.M. Wheeler).

FORMICA NEOGAGATES GROUP
Neogagates complex
Formica lasioides Emery

Formica neogagates lasioides: Cook, 1953, p. 360
Formica neogagates lasioides var. vetula: id. p. 361
Formica fusca fusca: id. (in part), p. 371
Formica subpolita var. camponoticeps: id. (in part), p. 397
Labelled neogagates lasioides var. vetula: 3 9 from Pacific Grove
(WMM), 3 9 from Lake Tahoe, 4 9 from Quincy (TWC), CALIF.
Labelled fusca: I $ , 3 9 from Lake Tahoe, CALIF.

Labelled subpolita camponoticeps: 19,59 from Glacier Point,
Yosemite, CALIF.

Unidentified: 4 9 from Eagle Creek (LSJU), 1 9 from Oswego
(Duncan), 2 9 from Rowena (LSJU), ORE.; 6 9 from Big Lagoon,
Humboldt Co., 5 9 from Lake Tahoe, 9 9 from Mirror Lake, elev.
4096 ft., CALIF.

1 Review Committee for this Contribution:

Jack C. Hall
Charles L. Hogue
G.C. Wheeler

2Universite du Quebec, Chicoutimi, Quebec, Canada G7H 2B1

3Section of Entomology, Natural History Museum of Los Angeles County,
900 Exposition Blvd. , Los Angeles, CA 90007

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 309:1-7.

Snelling: Ant Genus Formica

Formica neogagates Emery

Unidentified: 5 9 from Ithaca, N. Y.; 13 9 from Pullman, WASH.
(WMM).

Obtusipilosa COMPLEX

Formica perpilosa Wheeler

Labelled Formica: 4 9 from Santa Clara Co. , CALIF. The label is
the same as on other specimens from the Stanford University collec-
tion and they are surely from that collection.

FORMICA FUSCA GROUP

ClNEREA COMPLEX

Formica canadensis Santschi

Labelled fusca var. argentea: 1 9 from Colorado Springs, COLO.
(E.S. Tucker).

Formica pilicornis Emery

Formica cinerea var. lepida: Cook (in part), 1953, p. 373

Formica pilicornis: id. (in part), p. 385

Labelled pilicornis: 9 9 from South San Francisco and 18 9 from
San Francisco, CALIF. (TWC).

Labelled cinerea var. lepida: 16 9 from Saugus, CALIF.

Unidentified: 6 9 from Tejon Pass, CALIF. (TWC).

NEOCLARA COMPLEX

Formica neoclara Emery

Formica cinerea var. neocinerea: Cook (in part), 1953, p. 374.

Labelled fusca var. between neoclara and argentata: 3 9 from
Pullman, WASH. (LSJU).

Labelled fusca var. neoclara: 4 9 from Colorado Springs, COLO.
(E.S. Tucker).

Labelled fusca var.: 5 9 from Sisson, CALIF. (J.C. Bradley).

SUBPOLITA COMPLEX

Formica subpolita Mayr

Formica subpolita: Cook, 1953, p. 396.

Formica subpolita var. camponoticeps: id. (in part), p. 397.

Formica fusca var. marcida: id. (in part), p. 379.

Formica fusca var. neorufibarbis: id. (in part), p. 380.

Labelled subpolita: 8 9 from Pacific Grove (WMM), 8 9 from
Corte Madera Creek, 27 9 from Greenville, CALIF.

Labelled subpolita var. camponoticeps: 3 9 from Yosemite, 6 9
from Yosemite Falls Top, 3 9 from Yosemite Village, 19,39 from
Old Village, Yosemite Valley, 15 9 from Glacier Point, Yosemite, 2 9
from Mt. Tamalpais, CALIF.

Labelled fusca var. marcida: 5 9 from Point Joe, near Pacific
Grove, CALIF.

Labelled fusca var. neorufibarbis: 1 9 from top of Mount Wilson,
CALIF. (J.C. Bradley) together with 1 9 of Liometopum sp.

Lepida complex

Formica aerata Francoeur

Formica cinerea var. neocinerea: Cook, (in part), 1953, p. 374.

Labelled cinerea var. neocinerea: 3 9 from Three Rivers (J.C.
Bradley), 3 9 from Sacramento (H.S. Smith), 1 9 from Stanford
University, CALIF. (LSJU).

Labelled Formica: 3 9 from Stanford Univ. , CALIF.

Unidentified: 1 9 from Santa Clara Co., CALIF. (Harkins); 3 9

from 3.5 mi. south of Kirby, ORE.

Formica lepida Wheeler

Unidentified: 1 9 from Smith River Camp, Del Norte Co. , CALIF.
This is the second record for this species.

Formica longipilosa Francoeur

Figures 1-3

Unidentified: 20 9 and 2 6 from Ukiah, CALIF. (LSJU). These
specimens are the second series known to us. They provide new data
on the morphology of the workers. The male is described here for the
first time.

worker: The series fits the original description (Francoeur 1973,
p. 126-127). However, the size averages somewhat smaller: head
width 0.96-1.14 mm; head length 1.09-1.31; eye diameter 0.41-
0.47; scape length 1.07-1.27; trunk length 1.59-1.84. The dorsal
margin of petiole less convex, in one case feebly and narrowly
excised.

male: First description based on two androtypes (in LACM and
senior author’s collection). General morphology similar to F. trans-
montanis Francoeur, but averaging smaller. Measurements in mm:
head width 1.61-1.35; eye diameter 0.70-0.71; trunk length 3.05-
3.20; pronotum width 1.56-1.65. Head distinctly broader than long,
occipital margin slightly convex, ocelli small (0.12-0.14 mm);
clypeus bulging, faintly carinated, anterior margin convex; mandi-
ble rather narrow with only an apical tooth. Petiole low and thick in
profile, dorsal margin with large, weak concavity and rounded
comers. Genitalia: posterior half of parameres more acute and
larger, with long hairs; subgenital plate shining, when ventral sur-
face is seen from above, posterior end not produced as in transmon-
tanis, rather straight, with two concavities separated medially by
rounded process. Body concolorous brown, antennae paler, legs
yellowish brown; most parts with dense and long pubescence,
particularly on anterior dorsal half of head, pronotum, petiole and
gaster; pilosity distributed as in transmontanis but longer and finer
especially on head and thorax.

Formica pacifica Francoeur

Unidentified: 5 9 from Cannon Beach (LSJU), 3 9 from Goble
(LSJU), 1 alated $ from Umpqua River, Canyonville, ORE.

SUBSERICEA COMPLEX

Formica argentea Wheeler

Formica sibylla: Cook (in part), 1953, p. 393.

Labelled fusca var. argentata or var. argentea: 8 9 from Pullman,
WASH. (WMM).

Labelled fusca var.: 2 9 2 6 , & 1 9 from San Juan Island, WASH.
(WMM).

Labelled sibylla: 3 9 from Lake Tahoe, CALIF.

Unidentified: 1 9 from Boyd, 2 9 from Linsley, 1 9 from Rowena
(LSJU), 7 9 from Tygh Valley (LSJU), ORE.; 1 9 from Hewlett
Point, 1 9 from Prescott (L.R. Dice), WASH.

Formica podzolica Francoeur

Formica sibylla: Cook (in part), 1953, p. 393.

Labelled sibylla: 18 9 from Marysville, CALIF.

Formica subsericea Say

Unidentified: 6 9 from Ithaca, N.Y.

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 309:1-7.

Snelling: Ant Genus Formica

Figures 1-3. Formica longipilosa male (SOX). 1, head, frontal view, without pilosity; 2, petiole, lateral view; 3, paramere, lateral view.

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 309:1-7.

Snelling: Ant Genus Formica

Fusca complex

Formica accreta Francoeur

Formica fusca var. subsericea: Cook (in part), 1953, p. 384.

Formica sibylla: id., p. 393.

Labelled fusca var.: 26 9 & 1 $ from Quincy, CALIF.

Labelled fusca var. subsericea: 3 9 from Stinson Beach, CALIF.
(TWC).

Labelled sibylla: 5 9 from Mirror Lake, Yosemite Valley, 2 9 from
Glacier Point, Yosemite, 17 9 from Carmel (together with 1 6 of
Camponotus sp.), CALIF.

Unidentified: 4 9 from Patrick Creek, CALIF; 1 9 from Oswego
(Duncan), 2 9 from Eagle Creek (LSJU), 1 9 from Columbia
(LSJU), ORE.

Formica fusca Linne

Formica neogagates lasioides var. vetula: Cook (in part), 1953,
p. 360.

Formica fusca fusca: id. (in part), p. 369.

Formica cinerea var. neocinerea: id. (in part) p. 373.

Formica fusca var. argentea: id. (in part), p. 375.

Formica sibylla: id. (in part), p. 392.

Formica fusca: Francoeur, 1973, p. 189-201.

Specimens classified in the marcida form:

Labelled cinerea var. neocinerea: 3 9 from Mt. Tamalpais,
CALIF.

Labelled sibylla: 6 9 & 2 9 from Alta Meadow (WMW), 2 9,19
& 1 <5 from Lake Tahoe (WMW), CALIF.

Labelled neogagates lasioides var. vetula: 4 9 & 1 9 from Alta
Meadow, CALIF. (WMW).

Labelled fusca var. neoclara: 3 9 from Lake Tahoe, CALIF.
(WMW).

Unidentified: 2 9 from Yosemite, CALIF; 1 9 from Mt. Hood,
ORE. (LSJU).

Specimens classified in the subaenescens form:

Labelled fusca: 4 9 from Lake Tahoe, CALIF. (WMW).

Labelled /irstY/ var. argentea: 3 9 from Colorado Springs, COLO.
(E.S. Tucker).

Formica microphthalma Francoeur

Formica sibylla: Cook (in part), 1953, p. 393.

Labelled sibylla: 5 9 from Lake Tahoe, CALIF. (WMW).

Neorufibarbis complex

Formica neorufibarbis Emery

Formica subpolita: Cook (in part) 1953, p. 395.

Labelled rufibarbis var. gnava. 89,3 alated 9 & 1 $ from Lake
Tahoe, CALIF. (WMW).

Labelled subpolita: 18 9 & 3 9 Blue Lake, CALIF. (J.C. Bradley).

Labelled subpolita var. camponoticeps: 11 9 from Glacier Point,
CALIF. (WMW).

The workers of the above series are among the largest for the
species.

Labelled fusca var.: 2 9 from San Juan Isl., WASH.

Unidentified: 1 9 from Paradise Valley, 2 9 from Mt. Hood, 6 9
from Cannon Beach, ORE; 1 9 from San Juan Isl., WASH.

Rufibarbis complex

Formica foreliana Wheeler

Labelled fusca var. gnava: 3 9 from Ramsey Canyon, Huachuca
Mts. ARIZ. (WMM). These are topotypes.

Formica occidua Wheeler

Formica rufibarbis var. occidua: Cook (in part) 1953, p. 391.

Formica rufa obscuripes: id. (in part), p. 406.

Formica rufa propinqua: id. (in part), p. 409.

Formica microgyna californica: id. (in part), p. 413.

Labelled rufibarbis var. occidua: 1 9 from Stanford University, 21
9 from Napa (TWC), 6 9 from San Rafael, CALIF. (TWC).

Labelled rufibarbis var. gnava: 3 9 from Santa Cruz Isl., CALIF.
(R.V. Chamberlain).

Labelled rufa obscuripes: 1 9 from Marin Co. (TWC), 15 9 from
Del Monte (TWC), 9 9 from Santa Cruz Isl.. CALIF. (R.V. Cham-
berlain).

Labelled truncicola integroides var. propinqua: 4 from Pismo
(Beach), and 3 9 from Santa Cruz Isl., CALIF. (R.V. Chamberlain).

Labelled microgyna californica: 3 9 from Del Monte (TWC), 6 9
from Santa Cruz Isl., CALIF.

Unidentified: 3 9 from Cazadero (J.C. Bradley), 5 9 from Garber-
ville, 3 9 from Santa Cruz Isl. (R.V. Chamberlain), 2 9 from Santa
Clara Co. (Harkins), 1 9 from Berkeley, 20 9 from Davis, CALIF.

Sibylla complex

Formica sibylla Wheeler

Formica sibylla: Cook (in part), p. 393.

Labelled sibylla: 3 9 from Happy Isles, Yosemite Valley, 3 9 from
Alta Meadow, 3 9 from Lake Tahoe, CALIF.

Labelled fusca: 3 9 from Lake Tahoe, CALIF.

Without name: 4 9 from Glacier Point, CALIF.

SPECIES ERRONEOUSLY CITED
AS FORMICA
Subf. DOLICHODERINAE
Tapinoma sessile (Say)

Formica fusca var. marcida: Cook (in part), p. 379.

Labelled fusca var. marcida: 3 9 from Mt. Tallac, Lake Tahoe,
CALIF.

Subf. FORMICINAE
Lasius niger (Linne)

Formica neogagates var. lasioides: Cook (in part), p. 360.

Labelled neogagates var. lasioides: 3 9 from Lake Tahoe, CALIF.

Lasius pal li tar sis (Provancher)

Formica fusca var. blanda: Cook (in part), p. 377.

Labelled fusca var. blanda: 6 9 from Alta Meadow, CALIF.

Myrmecocystus mendax Wheeler

Formica subpolita camponticeps: (in part), p. 397.

Labelled subpolita var. camponoticeps: 6 9 from Hidden Springs
Canyon, CALIF. (Snelling, 1976).

Myrmecocystus mimicus Wheeler

Formica sibylla: Cook (in part), p. 393.

Labelled sibylla: 2 9 from Pasadena, CALIF.

In each of the above the specimens cited are the only examples
from the cited localities with the identification listed. They are,
therefore, the presumed basis for the record in Cook’s book.

ACKNOWLEDGMENT

This note is a by-product of grant no. A6501 of the National
Research Council of Canada.

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 309:1-7.

Snelling: Ant Genus Formica

LITERATURE CITED

Arnaud, PH, Jr. and Wale, Mary M. 1963. Thomas Wrentmore
Cook (1884-1962). Pan-Pacific Entomol. 49:177-181.
Brown, W.L. and Wilson, E.O. 1953. The Ants of California by
Thomas W. Cook (Review) Entomol. News 64:163-164.
Cook, T. W. 1953. The Ants of California, Pacific Books. Palo Alto,
Calif. 464 pp.

Francoeur, A. 1973. Revision taxonomique des especes
nearctiques du groupe fusca, genre Formica (Formicidae,
Hymenoptera). Soc. Entomol. Quebec, Memoire. no. 3,316 pp.
Snelling, Roy R. 1976. A revision of the honey ants, genus
Myrmecocystus. (Hymenoptera: Formicidae). Los Angeles Co.
Mus. Nat., Sci. Bull. 24:1-163.

Accepted for publication November 8, 1977.

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 309:1-7.

SERIAL PUBLICATIONS OF THE
NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY

Prior to November 30. 1 973, publications of the Natural History Museum have appeared under various titles — Leaflet
Series, Museum Graphic, Education Series. History Series , Science Series, Study Guides, Contributions in History, History
Bulletin, Science Bulletin, unnumbered catalogs of exhibitions, and other miscellaneous publications.

Contributions in Science
Education Series
History Series
Science Series

the world.

Acceptance of manuscripts will be determined by the significance of new information. Priority will be given to
manuscripts by staff members. AH manuscripts must be recommended by the curator in charge of each discipline or by the
Editorial Board. Peer judgment will be accomplished through the various divisions of the Museum and by outside

reviewers.

Number 310
March 16, 1979

CONTRIBUTIONS IN SCIENCE

NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY

CONTENTS

Abstract 3

Methods and Materials 4

Osteology 5

Systematics : Family Caulophrvnidae Regan 12

Robia new genus 12

Key to the Genera and Species of the Caulophrynidae 12

Robia legula new species 14

Caulophryne Goode and Bean 1896 14

Caulophryne pelagica (Brauer 1902) 14

Caulophryne jordani Goode and Bean 1896 15

Caulophryne polynema Regan 1930 16

Caulophryne species 18

Caulophryne setosus Goode and Bean 1896 nomen nudum 20

Caulophryne racemosa Monod 1960 nomen nudum 20

Discussion 21

Acknowledgments 24

Literature Cited 24

ISSN: 0459-8113

Suggested Citation: Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 310:1-25.

Contributions in Science are articles in the earth and life sciences, presenting results of original research in Natural
History. Science Bulletin (Numbers 1-30; 28 June 1965 to 10 April 1978) and Contributions in Science (Numbers 1-300;
23 January 1957 to 16 June 1978) were merged into a single imperial octavo serial beginning with Number 301.

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 310:1-25.

SYSTEMATICS AND DISTRIBUTION OF CERATIOID ANGLERFISHES OF THE
FAMILY CAULOPHRYNIDAE WITH THE DESCRIPTION OF A NEW GENUS AND

SPECIES FROM THE BANDA SEA1

By Theodore W. Pietsch2

Abstract: The ceratioid anglerfish family Caulophrynidae is revised on the basis of all known material. The
genus Caulophryne is represented by 26 female specimens collected from all three major oceans of the world.
Of the six nominal species, three are recognized: C. pelagica, with C. ramulosa and C. acinosa as synonyms;
C. jordani, with C. regard as a synonym; and C. polynema. A new caulophrynid genus and species is
described from a single female recently collected from the Banda Sea. It differs from Caulophryne in having a
considerably longer illicium, shorter jaw teeth, and a smaller number of relatively short median-fin rays. The
tentative distribution of each species is plotted, evolutionary relationships are discussed, and a key to the
genera and species of the family is provided.

This study supports a classification of the Ceratioidei in which the Caulophrynidae is the primitive sister-
group of the Melanocetidae, Himantolophidae, Diceratiidae, Centrophrynidae, Oneirodidae, and Thaumatich-
thyidae. The position of the closely related families Linophrynidae, Gigantactinidae, and Neoceratiidae (and
perhaps the Ceratiidae) is unclear. This assemblage is apparently derived either from a caulophrynid-like ances-
tor, forming a part of a sister-group that is primitive to all other ceratioids, or it is derived from some
oneiroidid-like ancestor. In either case, it is concluded that sexual parasitism has evolved independently in at
least two separate ceratioid lineages.

The ceratioid family Caulophrynidae includes globose, bathy-
pelagic anglerfishes, easily distinguished from members of allied
families by the absence of an escal bulb, the presence of two
pectoral radials, extremely long dorsal and anal rays, and neuro-
masts of the acoustico-lateralis system located at the tips of elon-
gate filaments. The only currently recognized genus of the family
was established by Goode and Bean (1896) with the description
of Caulophryne jordani based on a single female specimen with a
parasitically attached male collected off Long Island, New York,
by the Fisheries Steamer ALBATROSS . Since this original publica-
tion, five additional forms have been described, each from a sin-
gle adolescent or adult female: C. pelagica (Brauer 1902); C.
polynema Regan 1930; C. ramulosa and C. acinosa Regan and
Trewavas 1932; and C. regard (Roule and Angel 1932). At the
time of Bertelsen’s (1951) monograph on the Ceratioidei the
family was represented by only nine female specimens, one para-
sitic male, a male in metamorphosis, and 16 larvae. Not finding
sufficient reason for subdividing this material into several
species, Bertelsen (1951:37) provisionally recognized three sub-
species: C. jordani jordani, representing a western North Atlantic
subspecies; C. jordani pelagica, with C. ramulosa and C.
acinosa as synonyms; and C. jordani polynema, including C.
regani as a synonym.

The available material of Caulophryne represents three species:
C. pelagica, represented by six female specimens collected from
the eastern and Indo-west Pacific Ocean; C. jordani, by 11

females from all three major oceans of the world; and C. poly-
nema, known from eight females and one attached male from the
eastern Pacific and eastern Atlantic. The separation of these
species is based on differences in dorsal and anal fin ray counts,
tooth counts, illicium length, and illicial and escal morphology.

Despite a nearly three-fold increase in the number of known
metamorphosed female specimens of Caulophryne since Bertel-
sen’s (1951) revision, males are represented by only three indi-
viduals. Two of these are parasitically attached adults; the third is
an unattached metamorphosis stage.

A new caulophrynid genus is also described in this paper and
compared with Caulophryne. The single known female specimen
of this new form is part of the extensive midwater collections
made by Bruce H. Robison of the Marine Science Institute, Uni-
versity of California, Santa Barbara, during the 1975 ALPHA
HELIX Southeast Asian Bioluminescence Expedition.

'Review Committee for this Contribution:

Gerhard Krefft
Robert J. Lavenberg
Norman B. Marshall

department of Biology, California State University, Long Beach, Cali-
fornia 90840; and Research Association in Ichthyology, Natural History
Museum of Los Angeles County, Los Angeles, California 90007. Pres-
ent address: College of Fisheries, University of Washington, Seattle,
Washington 98195.

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 310:1-25.

Pietsch: Caulophrynid anglerfishes

METHODS AND MATERIALS

Standard lengths (SL) are used throughout. Measurements
were taken from the left side whenever possible and rounded to
the nearest 0.5 mm. To insure accurate fin-ray counts, skin was
removed from the pectoral fins and incisions were made to reveal
the rays of the dorsal and anal fins. Sockets, indicating missing
teeth in the jaws and on the vomer, were included in total tooth
counts. Jaw-tooth counts are the sum of both left and right sides.
Head length is the distance from the anterior tip of the upper jaw
to the posteriormost margin of the preoperculum. Head width is
the distance between the tips of the sphenotic spines. Head depth
is the distance from the tip of the sphenotic spine to the base of
the quadrate spine. Lower-jaw length is the distance from the
symphysial spine to the posteriormost margin of the articular.
Illicium length is the distance from the articulation of the pte-
rygiophore of the illicium and the illicial bone to the distal sur-
face of the esca, excluding escal appendages. Lengths given for

dorsal and anal rays are estimates of actual values since their
distal portions are nearly always broken off.

Terminology used in describing the various parts of the angling
apparatus follows Bradbury (1967). Definitions of terms used for
the different stages of development follow Bertelsen (1951).

Locality data are presented for all specimens examined. Coor-
dinates for the starting position only of each trawl are included.
Most collections were made with an open, 3-meter Isaacs-Kidd
midwater trawl (IKMT). Material deposited at the Institut fur
Seefischerei, Hamburg, was collected with a Combined Midwater
Bottom Trawl, having a rectangular mouth and a 1600 mesh cir-
cumference (CMBT-1600). This and similar gear are described
more fully by SchMe (1966, 1969). Material from the Institute
of Oceanographic Sciences, Surrey, England, was collected with
a Rectangular Midwater Trawl, mouth area eight square meters,
mesh size five millimeters (RMT 8/5), equipped with an opening
and closing device (Clark 1969).

Material used for the osteological investigation (two female

Figure 1. Dorsal view of cranium of Caulophryne jordani, IOS, 68.0 mm.

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 310:1-25.

Pietsch: Caulophrynid anglerfishes

PTEROTIC

Figure 2. Lateral view of cranium of Caulophryne jordani, IOS, 68.0 mm.

Caulophryne jordani, 41.0-68.0 mm) was cleared and stained
with alizarin red S following the trypsin digestion technique
(Taylor 1967). In many cases dissections were made of uncleared
specimens to confirm observations made on cleared and stained
specimens and to determine ontogenetic changes. Bone terminol-
ogy follows Pietsch (1972a) and Nybelin (1963). In osteological
drawings cartilage is stippled, and where necessary for clarity,
open spaces are rendered in solid black.

Unless otherwise indicated, all diagnoses and descriptions are
based on females. For males and larvae see Bertelsen
(1951:31-38). Material is deposited in the following museums
and Institutions:

BMNH: British Museum (Natural History), London.

IOAN: Institute of Oceanography, Academy of Sciences of the

USSR, Moscow.

IOS: Institute of Oceanographic Sciences, Surrey, England

(formerly the National Institute of Oceanography).

ISH: Institut fur Seefischerei, Hamburg.

LACM: Natural History Museum of Los Angeles County.

MCZ: Museum of Comparative Zoology, Harvard University.

MOM: Musee Oceanographique de Monaco.

ROM: Royal Ontario Museum, Toronto.

USNM: National Museum of Natural History, Washington.
ZMB: Zoologisches Museum der Humboldt-Universitat, Ber-

lin.

ZMUC: Zoological Museum, University of Copenhagen.

OSTEOLOGY

Figures 1-13

The osteology of Caulophryne was partially described by

Regan (1926:16, fig. 7), Regan and Trewavas (1932:42, figs.
56-58), and Bertelsen (1951:31, fig. 8). In the following
account, only those comparative aspects that need amending or
that have not previously appeared in the literature are discussed.

Cranium (Figs. 1-6): the ethmoid cartilage of Caulophryne
broadly covers the dorsal surface of the vomer, extending beyond
the margins of this bone on all sides. Anteriorly and laterally the
head of the vomer turns upward to cup the ethmoid cartilage. The
ventral surface of the vomer may bear as many as five, recurved
and depressible teeth arranged in a transverse row.

In oneirodids (Pietsch 1974a:8, figs. 1, 4) the symphysial carti-
lage of the upper jaw butts up against a nearly vertical supraeth-
moid bone. In contrast, backward movement of the upper jaw of
Caulophryne is limited by the lateral extensions of the vomer; an
elongate, narrow groove is formed between the parallel lateral
ethmoids and the obliquely placed supraethmoid providing space
within which the small symphysial cartilage (see below) can
slide. The anterior part of the pterygiophore (basal bone) of the
illicium does not lie in this narrow groove as stated by Regan
(1926:16).

The frontals are considerably shorter than those of oneirodids
(Pietsch 1974a, figs. 1, 2). They meet on the midline for most of
their length. Anteriorly, they contribute to the extremely narrow
orbital region, overlapping the posterior ends of the lateral eth-
moids; posteriorly they become greatly expanded laterally.

The parietals of Caulophryne occupy a more anterior position
than in oneirodids, and are far removed from the posttemporals.
Each is expanded laterally in an oblique position, overlapping
four other bones: the sphenotic laterally, the pterotic and epiotic
posteriorly, and the supraoccipital medially. Anteromedially,
each parietal is slightly overlapped by the respective frontal bone.

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 310:1-25.

Pietsch: Caulophrynid anglerfishes

■SPHENOTIC

Figure 3. Ventral view of cranium of Caulophryne jordani, IOS, 68.0 mm.

Figure 4. Anterior view of anterior half of cranium of Caulophryne
jordani, IOS, 68.0 mm.

The pterosphenoid is absent (Pietsch 1972a:29).

Each sphenotic is a large, anterolaterally directed, cone-shaped
element, the apex of which forms an elongate, blunt projection.
On its ventral margin, each frontal, with the respective prootic
bone, forms an oval-shaped cartilaginous area within which the
anterior head of the double-headed hyomandibular articulates.
The pterotic is also cone-shaped, but posterolaterally directed
with a much larger base. A facet for the articulation of the poste-
rior head of the hyomandibular bone is located ventraliy at the
apex of each pterotic.

In oneirodids (Pietsch 1974a:7, figs. 1-3) the prootics are sep-
arated anteriorly by the anteroventral process of the supraoccipi-
tal. In Caulophryne the supraoccipital maintains a dorsal posi-
tion, largely overlapped anteriorly by the frontals, so that the
prootics meet on the midline contributing to the formation of the
posterior surface of the orbit.

Mandibular arch (Figs. 7, 8): Each premaxillary has a short
articular process lying slightly behind and below an ascending
process of similar size. Lying on the midline at the posterior tips
of the ascending and articular processes of the premaxillae is a
small, rounded symphysial cartilage loosely held by short liga-
ments and connective tissue. A postmaxillary process of the pre-
maxillary is absent. The curved elongate portion of each premax-
illary bears 12 recurved, depressible teeth of mixed sizes.

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 310:1-25.

Pietsch: Caulophrynid anglerfishes

SUPRA OCCIPITAL

SPHENOTIC

Figure 5. Anterior view of posterior half of cranium of Caulophryne jordani, IOS, 68.0 mm

PTEROTIC

CUT SURFACE EPIOTIC

Figure 6. Posterior view of cranium of Caulophryne jordani, IOS, 68.0 mm.

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 310:1-25.

Pietsch: Caulophrynid anglerfishes

ASCENDING PROCESS

Figure 7. Upper jaw of Caulophryne jordani: A. IOS, 68.0 mm, lateral
view of maxillary and premaxillary, right side; B. LACM 6844-3, 41.0
mm, lateral view of symphysis of premaxillaries showing symphysial car-
tilage.

The articular head of the maxillary consists of a slightly con-
cave dorsal process which receives the anterior head of the
double- headed palatine bone (see below), and a ventromedial
projection which passes beneath, and provides support for, the
premaxillary.

On each side, the posterior ends of the premaxillary and maxil-
lary are united by a short ligament that passes posteriorly to
attach to the ascending process of the articular. The anterior-max-
illomandibular ligament (labial cartilage of Le Danois 1964;
Pietsch 1972a:31), so well developed in oneirodids (Pietsch
1974a:9), is absent.

Anteriorly the dentaries curve to meet on the midline where
they form a strong symphysial spine. The anterior half of each
dentary bears nine to ten recurved, depressible teeth of mixed
sizes.

Palatine arch (Fig. 8): The palatine arch is elongate and slender
throughout. The anterior end of the palatine is T-shaped, consist-
ing of a posterior head which is attached to the ethmoid cartilage,
and an anterior head, which is loosely held within a shallow cav-
ity of the dorsal process of the respective maxillary bone by con-
nective tissue and by the palatomaxillary ligament. A narrow
mesopterygoid is present but makes no contact with the metap-
terygoid.

Hyoid Arch (Figs. 8, 9): The suspensorium of Caulophryne is
unusually narrow and elongate, and strongly directed obliquely
backwards (compare with a typical oneirodid; Pietsch 1974a, fig.
8). The posterior head of the double-headed hyomandibular is the
larger and longer of the two heads, forming a broad articulation
with the pterotic. The anterior head of the hyomandibular articu-

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 310:1-25.

lates within a cartilaginous area formed between the sphenotic
and prootic bones (the pterotic does not border on the articular
area of the anterior head of the hyomandibular as it does in onei-
rodids; Pietsch 1974a; 10, figs. 2, 3).

The greater length of the suspensorium, relative to other cera-
tioids (see above), is due largely to the extremely narrow, elon-
gate quadrate. A large cleft is formed between the anterior and
posterior portions of the quadrate within which lies the narrow
ventral end of the symplectic. The interhyal is large compared to
that of oneirodids (Pietsch 1974a, fig. 8).

The epihyal and especially the ceratohyal are narrow and elon-
gate compared to those of oneirodids (Pietsch 1974a, fig. 9). In
all but the most specialized lophiiforms (e.g., Gigantactis, Neo-
ceratias) examined by me (Pietsch 1974a, Appendix B), the cera-
tohyal is notched anterodorsally to receive the posterior extension
of the elongate, dorsal hypohyal (Rosen and Patterson 1969:439,
fig. 57; Pietsch 1972a, fig. 14; 1974a, figs. 9, 10). In Caulo-
phryne the anterodorsal notch is absent; the dorsal hypohyal is
short and does not extend back over the dorsal surface of the
ceratohyal. At midlength each ceratohyal forms a dorsal process
that is securely attached to the anterolateral surface of ceratobra-
chial I by ligamentous connective tissue. A narrow ligament also
extends anteriorly from the dorsal process of the ceratohyal to
insert on the cartilage of the hypohyals.

The left ceratohyal of the 68.0 mm specimen (Fig. 9A) appears
to be abnormally developed in having a deeply excavated pos-
teroventral margin; together with the anteroventral comer of the
epihyal, it supports only five branchiostegal rays: one anterior
element (without an expanded head) on the inner side of the ante-
rior, depressed portion of the ceratohyal, and four posterior ele-
ments (with expanded heads) on the outer surface of the elevated
portion of the ceratohyal. The right ceratohyal (Fig. 9B) appears
to be normally developed bearing six branchiostegals. Only one
of these, however, is located anteromedially; five (all with
expanded heads) are in a posterolateral position. Examination (by
dissection) of all additional available material of Caulophryne
showed the normal complement of six branchiostegal rays, two
anteromedial elements, and four posterolateral elements. Previ-
ous to this, no lophiiform examined by me has had more than
four posterior, laterally articulated branchiostegals; this number is
considered diagnostic for the Paracanthopterygii as well as the
Acanthopterygii (Greenwood, et al. 1966:353; Rosen and Patter-
son 1969:439).

Opercular bones (Fig. 8): The reduced operculum is bifurcated
proximally forming two, extremely slender, and nearly parallel
forks. The lower fork, more than three times the length of the
upper, supports the suboperculum distally. The suboperculum is
small, weakly-ossified (thought to be absent by Regan and
Trewavas 1932:43, fig. 56), and deeply notched on its posterior
margin. The large, posteriorly- curved preoperculum strengthens
the entire length of the long hyoid arch extending dorsally along
the posterior margin of the hyomandibular well past the point of
the articulation of the operculum.

Branchial arches (Fig. 10): Pharyngobranchials I and IV are
absent; those of the second and third arches are extremely well
developed, bearing four to six recurved depressible fangs. Epi-
branchial I and ceratobranchial V are reduced, lying free in the
connective tissue matrix. No ossified or cartilaginous remnants of
hypobranchials or basibranchials could be found.

Vertebrae and caudal skeleton (Fig. 11): In the two cleared and
stained specimens examined there are 20 vertebral centra (includ-
ing the half-centrum to which is fused the hypural plate; Pietsch

Pietsch: Caulophrynid anglerfishes

Figure 8. Lateral view of left side of lower jaw and suspensorium of Caulophryne jordani, IOS, 68.0 mm, with opercular bones in place.

DORSAL PROCESS

Figure 9. Lateral views of hyoid apparatus of Caulophryne jordani, IOS,
68.0 mm: A. Left side; B. Right side. Interhyal not shown.

1972a: 38) of which ten or 11 are caudal vertebrae (those centra
bearing complete haemal arches). There is a single, reduced
epural (Pietsch 1972a:38). The hypural plate, deeply notched
posteriorly in the larger cleared and stained specimen, but entire
in the smaller specimen, bears the overlapping bases of eight
principal caudal rays, the uppermost of which is exceptionally
large. The central four caudal rays are bifurcated distally.

Median fins and illicial apparatus (Figs. 11, 12): The rays of
the median fins are unusually long compared to other ceratioids,
measuring, in extreme cases, more than twice the standard
length. These rays are all biserial, segmented and unbranched,
supported by elongate radials. In the 68.0 mm specimen there are
19 dorsal rays supported by 14 radials. The anteriormost radial
supports the first dorsal ray and lies above the neural spine of the
12th pre-ural centrum. The posteriormost radial (which appears
to be the product of two fused radials) supports the 14th and 15th
dorsal rays and lies above the neural spine of the sixth pre-ural
centrum. In the anal fin there are 17 rays supported by 14 closely
associated radials. The anteriormost radial supports the first anal
ray and lies below the haemal spine of the eighth pre-ural
centrum. The posteriormost radial supports the 14th anal ray and
lies below the haemal spine of the fourth pre-ural centrum.

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 310:1-25.

Pietsch: Caulophrynid anglerfishes

Figure 10. Branchial arches of Caulophryne jordani, IOS, 68.0 mm. The
ventral part of the branchial basket is shown in dorsal view, the dorsal
part (epibranchials and pharyngobranchials) is folded back and shown in
ventral view.

The pterygiophore of the illicium, excluding its downtumed,
cartilaginous posterior portion, is approximately 16 percent of
standard length. The remnant of the second cephalic ray is a
minute, toothlike ossification that lies on the pterygiophore just
behind the articulation of the illicial bone. The length of the illic-
ial bone varies slightly among Caulophryne species and becomes
longer proportionately with growth (Fig. 16).

Pectoral girdle, pectoral fin, and pelvic bones (Fig. 13): The
posttemporals of Caulophryne are small compared to those of
other ceratioids. In oneirodids each posttemporal meets the
respective parietal and overlaps the pterotic, epiotic and exoccip-
ital (Pietsch 1974a: 14, figs. 1-3, 5, 6). The reduced posttem-
poral of Caulophryne overlaps only the pterotic and epiotic being
far removed from both the parietal (see above) and the exoccipi-
tal.

The supracleithrum of Caulophryne is unusually large covering
most of the lateral surface of the upper portion of the cleithrum.
The lower portion of the cleithrum is reduced to a slender, thread
of bone.

The scapula consists of a rounded ossification bounded on
three sides by cartilage. The posterior margin of the ossified part
of the scapula supports the uppermost pectoral radial. The cora-
coid consists of an expanded dorsal portion which meets the carti-
lage of the scapula, an anteroventral process which is connected
by a ligament to the reduced lower portion of the cleithrum, and
an extremely large posteroventral process which meets the slen-

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 310:1-25.

Pietsch: Caulophrynid anglerfishes

der postcleithrum distally.

There are two separate, well-ossified radials in fully metamor-
phosed specimens of Caulophryne, a small upper radial that bears
no pectoral fin rays, and a much larger, distally expanded, lower
radial that bears all the rays of the pectoral fin. The lower radial,
however, is probably the product of two or more fused radials. A
cleared and stained metamorphosing female (10.0 mm) clearly
shows a deep, posterior cleft in the lowermost radial indicating at
least three centers of ossification in the pectoral lobe (Fig. 13B).
Fusion of pectoral radials appears to be common among cerati-
oids (Pietsch 1972a:41, fig. 23; Bertelsen and Struhsaker 1977).

Caulophryne is unique among ceratioids in the retention of
pelvic fins in larval stages. Metamorphosis involves the loss of
the pelvic fin rays but a pelvic bone is retained. The slender
pelvic bones of Caulophryne lie free in the connective tissue ma-
trix, far removed from the cleithrum. Unlike other ceratioids,
these elements do not have cartilaginous distal tips.

Skin spines: Thorough microscopic examination of the skin of
cleared and stained specimens failed to show the presence of em-
bedded dermal spines.

Figure 12. Illicial apparatus of Caulophryne jordani, IOS, 68.0 mm.

Figure 13. Lateral view of pectoral girdle, pectoral fin, and pelvic bone of Caulophryne: A. C. jordani, IOS, 68.0 mm; B. C. sp., IOAN, 10.0 mm.

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 310:1-25.

Pietsch: Caulophrynid anglerfishes

SYSTEMATICS
Family Caulophrymidae Regan

Type genus Caulophryne Goode and Bean 1896.

Caulophrynidae Regan 1912:285, 288. Regan 1926:4, 16, 22. Regan
1930:195. Regan and Trewavas 1932:6, 14, 24, 25, 26, 42, 100. Roule
and Angel 1933:55. Fowler 1936:1145, 1346. Belloc 1949:958. Bertelsen
1951:29, 31. Grey 1956:232. Monod 1960:684. Le Danois 1964:13, 14.
Kobayashi, et al. 1968:8. Fitch and Lavenberg 1968:125, Fig. 69. Ueno
1971:102. Pietsch 1972a:18, 43, 45. Maul 1973:666. Pietsch 1976:783,
788, 791, 792.

DIAGNOSIS: The metamorphosed females of the family
Caulophrynidae are distinguished from those of all other ceratioid
families by having two pectoral radials, extremely elongate dor-
sal and anal rays, eight caudal rays, neuromasts of the acoustico-
lateralis system located at the tips of extremely elongate fila-
ments, larval pelvic fins, and the absence of an escal bulb. In
addition, female caulophrynids share the following combination
of characters: jaws equal anteriorly; supraethmoid present; pari-
etals present, displaced anteriorly; pterosphenoid absent; ante-
rior-maxillomandibular ligament absent (labial cartilage of Le
Danois 1964; Pietsch 1972a:31); hyomandibular with a double
head; 2 hypohyals; 6 (rarely 5) branchiostegal rays; operculum
bifurcate, both forks reduced; suboperculum notched posteriorly,
anterior spine or projection absent; pharyngobranchial I absent;
epibranchial and ceratobranchial teeth absent; hypobranchials and
basibranchials absent; a single, reduced epural; posterior margin
of hypural plate notched in some specimens; only an ossified
remnant of second cephalic ray present; posttemporal reduced;
lower portion of cleithrum reduced; posteroventral process of
coracoid well-developed; pelvic fins present in larvae; pelvic
bones of metamorphosed females rod-like, not expanded distally;
esca without denticles; skin spines absent.

DESCRIPTION: Body relatively short, globular (Figs. 14, 20);
oral valve well-developed, lining inside of both upper and lower
jaws; two nostrils on each side at the end of a single short tube;
eye small and subcutaneous, appearing through a circular, trans-
lucent area of integument; gill opening oval in shape, situated
just posteroventral to pectoral lobe; all four epibranchials closely
bound together by connective tissue; posterior third of cerato-

branchial I bound to ceratobranchial II; no opening behind fourth
arch; gill filaments present on anterior-most tip of ceratobranchial
I and full length of ceratobranchials II through IV; pseudobranch
absent; pattern of placement of acoustico- lateralis system as
described for other ceratioids (Pietsch 1969, 1972b, 1974a, b);
neuromasts situated at the tips of elongate filaments; ovaries
paired; two, short pyloric caeca.

Illicium becoming proportionately longer with growth (Fig.
16); anterior end of pterygiophore of illicium exposed, emerging
on snout, its posterior end concealed under skin; esca unpig-
mented and translucent, simple, without elaborate appendages
(Robia), or consisting of several branched appendages or numer-
ous filaments ( Caulophryne ); escal bulb and central lumen
absent; entire length of illicium covered with translucent fila-
ments in some forms of Caulophryne.

Teeth slender, recurved and depressible, those in lower jaw
less numerous, but slightly longer than those in upper jaw; number
of teeth in lower jaw 14-34, in upper jaw 20-45; total number
of teeth on vomer 1-5; pharyngobranchials II and III heavily
toothed.

Color in preservative dark brown to black over entire surface
of body (except for esca, and in some forms, the illicium) and
oral cavity; viscera unpigmented.

D. 6-22; A. 5-19; P. 15-18 (Table 1); pelvics with 3-4 rays
in larvae and metamorphosis stages, absent in adolescents and
adults; C. 8 (2 unbranched - 4 branched - 2 unbranched).

The following measurements, expressed in percent of SL, are
summarized for the females (20.0- 142.0 mm) of all species:
head length 27.2-41.7; head depth 32.7-45.4; head width
21.9-30.6; lower jaw 31.3-48.1; premaxillary 28.9-44.4;
longest tooth in lower jaw 5. 1-9.5; longest dorsal ray
63.4-155.3; longest anal ray 39.0—15 1.1; illicium 16.0-268.3.

Robia new genus

Type species: Robia legula new species.

DIAGNOSIS: The genus Robia is distinguished from Caulo-
phryne (the only other member of its family) in having a consid-
erably longer illicium (268.3 percent of SL), shorter jaw teeth,

Key to the Genera and Species of the Caulophrynidae

The following key will differentiate adolescent and adult female specimens only. Characters that allow specific identification of males
(known from only three specimens, two of which are parasitically attached to females) and larvae (16 specimens; Bertelsen 1951:35, figs.
8, 1 1) are unknown.

IA. Illicium long, 268 percent of SL in a 41 mm specimen; D.6, A. 5 Robia legula new genus and species

(single known specimen, 41.0 mm).

IB. Illicium short, less than SL; D. 14-22, A. 12-19 Caulophryne 2

2A. Illicium with translucent filaments along entire length (Figs. 19, 21); length of illicium less than 35 percent SL (Fig. 16); D. 16-22,

A. 14-19 (Table 1) 3

2B. Illicium without filaments (Fig. 17); length of illicium greater than 35 percent of SL (Fig. 16); D. 14-17, A. 12-16 (Table 1)

C. pelagica (Brauer 1902).

3A. Upper jaw with more than 35 teeth in specimens larger than 40 mm, 30 or more in specimens larger than'20 mm (Fig. 18); illicium
with more than 50 filaments along entire length in specimens larger than 32 mm (Fig. 21); D. 20-22 (rarely 19), A. 18-19 (rarely

17) (Table 1) C. polynema Regan 1930.

3B. Upper jaw usually with less than 35 teeth in specimens larger than 40 mm, less than 30 in specimens larger than 20 mm (Fig. 18);

illicium with less than 15 filaments along length (Fig. 19); D. 16-18 (rarely 19), A. 14-17 (rarely 18) (Table 1)

C. jordani Goode and Bean 1896.

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 310:1-25.

Pietsch: Caulophrynid anglerfishes

TABLE 1

Frequencies of fin-ray counts for species of Caulophrynidae

Species

Dorsal

Anal

Pectoral

14 15

19 20

5 12 13

Robia legula

Caulophryne pelagica

3 1

Caulophryne jordani

Caulophryne polynema

1 1

Total

3 1

2 1

1 3 1

Figure 14. Robia legula, Holotype, LACM 36024-1, 41.0 mm. Drawn by Caryl Maloof.

and a smaller number of relatively short median fin rays (D.6,
less than 65 percent of SL; A. 5, less than 40 percent of SL).

ETYMOLOGY: The name Robia , to be treated as a noun of
feminine gender, is given in honor of Bruce H. Robison (better
known to his friends as “Robie”) in recognition of his contribu-
tions to our knowledge of midwater fishes.

Robia legula new species

Figures 14, 15, 24

HOLOTYPE: LACM 36024-1, 41.0 mm; ALPHA HELIX
Southeast Asian Bioluminescence Expedition Station 81, Banda

Sea, 4° 56.5'S, 129° 59.5'E, RMT-8 with closing device,
1000- 1500 m, 0416 - 0616 hr, 28 April 1975.

DESCRIPTION: Illicium without filaments, darkly pigmented
along entire length except near esca; esca translucent, with two
short lateral appendages and a slightly more opaque distal tip
bearing three short appendages (Fig. 15).

Total number of teeth in lower jaw 31, in upper jaw 33;
vomerine teeth 4; D. 6; A. 5; P. 17 (Table 1).

Measurements in percent of SL: head length 39.0; head depth
39.0; head width 23.4; lower jaw 47.6; premaxillary 37.8; long-
est tooth in lower jaw 5.1; longest dorsal ray 63.4; longest anal
ray 39.0; illicium 268.3.

Contrib. Sci. Natur. Hist. Mils. Los Angeles County. 1979. 310:1-25.

Pietsch: Caulophrynid anglerfishes

Figure 15. Esca of Robia legula, Holotype, LACM 36024-1, 41.0 mm.

Rest of characters as for genus and family.

DISTRIBUTION: Robia legula is known from a single speci-
men collected from the Banda Sea (4° 56. 5 'S, 129° 59. 5 'E) with
an opening-closing trawl between 1000 and 1500 m.

ETYMOLOGY: The name legula is derived from the Latin
legula , meaning collector or gatherer, alluding to the extraor-
dinary collecting abilities of this anglerfish and its collector.

Caulophryne Goode and Bean 1896

Caulophryne Goode and Bean 1896:496, fig. 409 (type species Caulo-
phryne jordani Goode and Bean 1896, by original designation and
monotypy).

Melanocetus Brauer 1902:295 (in part; type species Melanocetus johnsoni
Gunther 1864, by monotypy). Brauer 1906:321, pi. 15, fig. 5 (after
Brauer 1902).

Ceratocaulophryne Roule and Angel 1932:500 (type species Ceratocau-
lophryne regani Roule and Angel 1932, by monotypy). Roule and
Angel 1933:55 pi. 3, figs. 26, 26A (after Roule and Angel 1932;
expanded description).

DIAGNOSIS: The genus Caulophryne is distinguished from
Robia (the only other member of its family) in having a consider-
ably shorter illicium (less than 50 percent of SL), longer jaw
teeth, and a greater number of relatively long median fin rays (D.
14-22, greater than 70 percent of SL; A. 12- 19, greater than 60
percent of SL).

Caulophryne pelagica (Brauer 1902)

Figures 16-18, 24

Melanocetus pelagicus Brauer 1902:295 (original description, single
specimen). Brauer 1906:321, plate 15, fig. 5 (after Brauer 1902).
Regan 1926:22 (“probably a synonym of C. jordani”).
Caulophryne pelagicus Regan 1912:288 (new combination; listed).
Caulophryne jordani Regan 1926:22 (in part; specimen later described as
C. ramulosa sp. n. by Regan and Trewavas 1932). Kobayashi, et
al. 1968:8-12, figs. 1, 2, tables 1, 2 (additional specimen; descrip-
tion).

Caulophryne ramulosa Regan and Trewavas 1932:100, 101, pi. 7, fig.
160 (original description; single specimen, previously recorded by
Regan 1926, as C. jordani; in key) Bertelsen 1951:36, 37, tables 2,
3 (description; a synonym of C. jordani pelagica ).

Caulophryne acinosa Regan and Trewavas 1932:100, 101, pi. 8, fig. 161
(original description; single specimen; in key). Bertelsen 1951:36,
37, tables 2, 3 (description; a synonym of C. jordani pelagica).
Caulophryne pelagica Regan and Trewavas 1932:100, 102, fig. 162 (new
orthography; in key; description after Brauer 1902, 1906, esca fig-
ured). Bertelsen 1951:36, 37, fig. 9, table 3 (figure after Brauer
1906; a subspecies of C. jordani).

Caulophryne jordani pelagica Bertelsen 1951:33, 37, fig. 9, table 3 (in
key; description, comparison with all known material; includes C.
ramulosa and C. acinosa). Grey 1956:233 (synonymy; after Bertel-
sen 1951). Kobayashi, et al. 1968:9 (new specimen most similar
to).

MATERIAL: Six known females, 11.0- 92.0 mm.

Holotype of Melanocetus pelagicus: ZMB 17711, 11.0 mm;
VALDIVIA Station 228, Indian Ocean, 2° 38'S, 65° 59'E,
0-2500 m, bottom depth 3460 m.

Holotype of Caulophryne ramulosa: ZMUC P9245, 57.0 mm;
DANA Station 1209(1), Gulf of Panama, 7° 15'N, 78° 54'W,
3500 m wire, 1845 hr, 17 January 1922.

Holotype of Caulophryne acinosa: ZMUC P9244, 19.0 mm;
DANA Station 3920(2), Indian Ocean, 1° 06'S, 62° 25'E, 3500 m
wire, bottom depth 4630 m, 1830 hr, 9 December 1929.

NONTYPE MATERIAL: LACM 30619-6, 65.0 mm;
VELERO IV Station 9903, 29° 30'N, 118° 54'W, 3-m IKMT,
0-834 m, 2315-0515 hr, 7 August 1964. LACM 36023-1,
13.0 mm; ALPHA HELIX Southeast Asian Bioluminescence
Expedition Station 143, 0° 14.5'S, 128° 46.7'E, RMT-8,
1250- 1500 m, 1715- 1930 hr, 20 May 1975.

Fisheries Museum, Faculty of Fisheries, Hokkaido University
F-1639, 92.0 mm; No. 3 Shichisei Mam, 42° 40'N, 144° 40'E,
0- 500 m, 0800- 1200 hr, 26 April 1967.

DIAGNOSIS: In addition to differences in escal morphology
(Figs. 17, 19, 21), C. pelagica may be distinguished from C.
jordani and C. polynema in having a slightly longer illicium (Fig.
16), fewer dorsal fin rays (Table 1), and by the absence of illicial
filaments (Figs. 17, 19, 21). It is further separated from C. jor-
dani by having a greater number of upper jaw teeth (Fig. 18).
Caulophryne pelagica probably also has longer median fin rays
than its congeners, but this character is difficult to quantify accu-
rately because the distal ends of the rays are nearly always
broken off.

DESCRIPTION: Illicium without filaments, darkly pigmented
along entire length except near esca; esca consisting of five or six
elongate, filamentous appendages: two, more lateral in position,
bearing none to several side branches, and three or four, more
distal, bearing numerous side branches; opaque areas absent
(Regan and Trewavas 1932:101, 102, figs. 160-162; Fig. 17).

Total number of teeth in lower jaw 24-34, in upper jaw
28-40 (Fig. 18); vomerine teeth 2-3; D. 14-17; A. 12-16, P.
15-18 (Fig. 18, Table 1).

Measurements in percent of SL; longest tooth in lower jaw
5. 8-6. 8; longest dorsal ray 105-228; longest anal ray 132-174;
illicium 31.6-43.9 (Fig. 16).

Rest of characters as for genus and family.

DISTRIBUTION: Caulophryne pelagica is represented by two
specimens collected in the Indian Ocean, a third record from the
Halmahera Sea, East Indies (13.0 mm, collected with a closing
trawl between 1250-1500 m), a fourth from off Guadalupe
Island, Mexico, a fifth off Japan, and a sixth specimen from the

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 310:1-25.

0 10 20 30 40 50 60 70 80 90 100 110 120 130 140

Figure 16. Illicial length versus standard length for Caulophryne species.

Standard length in mm

Gulf of Panama (Fig. 24).

COMMENTS: The results of this investigation confirm Bertel-
sen’s (1951:36) opinion that C. ramulosa and C. acinosa are
junior synonyms of C. pelagica. The illicia of the holotypes of
all three are “well-preserved and constructed in a very similar
fashion.” Further, all three nominal forms agree in having a
slightly longer illicium and fewer median fin rays than other
species.

Caulophryne jordani Goode and Bean 1896

Figures 1-13, 16, 18, 19, 24

Caulophryne jordani Goode and Bean 1896:26*, 496, 541, fig. 409
(original description; single specimen; erroneous name in list of
plates, index, figure caption). Jordan and Evermann 1898:2735
(description after Goode and Bean 1896). Jordan and Evermann
1900, fig. 957 (figure after Goode and Bean 1896). Gill 1909:586,
fig. 24 (brief description and figure after Goode and Bean 1896).
Regan 1912:288 (description after Goode and Bean 1896). Beebe
1926:422 (additional specimen). Regan 1926:4, 22, figs. 7, 16
(cranial osteology; expanded description). Beebe 1929:19 (addi-
tional specimen). Regan 1930:191, 193 (comparison with C. poly-
nerna). Regan and Trewavas 1932:6, 100, 101, fig. 159 (in key;
brief description after Goode and Bean 1896; illicium figured).

^ — Figure 17. Esca of Caulophryne pelagica, LACM 30619-3, 65.0 mm.

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 310:1-25

Pietsch: Caulophrynid anglerfishes

Roule and Angel 1933:57 (comparison with C. regani). Bertelsen
1951:33-38, figs. 8-11, tables 2, 3 (description, comparison with
all known material; three subspecies). Lavenberg and Ebeling
1967:195; fig. 5 (vertical distribution). Kobayashi, et al.
1968:8- 12, figs. 1, 2, tables 1, 2 (additional specimen here
referred to C. pelagica; description). Fitch and Lavenberg
1968:125-127, fig. 69 (distinguishing characters; natural history).
Ueno 1971:102 (listed). Pietsch 1972a:29, 35, 36, 38, 41-43, 45,
fig. 24(3) (osteology; otolith described, figured). Pietsch 1976:783
(reproduction).

Ceratocaulophryne regani Roule and Angel 1932:500 (original descrip-
tion; single specimen). Roule and Angel 1933:55-57, pi. 3, figs.
26, 26a (expanded description: figures of holotype, lateral, dorsal
views). Belloc 1949:17 (listed). Bertelsen 1951:36, table 3 (a syn-
onym of C. jordani polynerna). Kobayashi, et al. 1968:8 (after Ber-
telsen 1951).

Caulophryne regani Fowler 1936:1347 (new combination; in key;

description after Roule and Angel 1932, 1933).

Caulophryne jordani polynerna Bertelsen 1951:33, 37 table 3 (in part;
includes C. regani here referred to C. jordani). Grey 1956:233
(after Bertelsen 1951). Maul 1973:666 (after Bertelsen 1951).
Caulophryne jordani jordani Bertelsen 1951:33, 37, table 3 (in key; one
of three subspecies; description, comparison with all known mate-
rial). Grey 1956:232 (synonymy; after Bertelsen 1951).

MATERIAL: Eleven known females, 23:0-109.0 mm.

Holotype of Caulophryne jordani: USNM 39265, 26.0 mm;
ALBATROSS, off Long Island, New York, 39° 27'N, 71° 15'W,
0-2335 m, 19 September 1887.

Holotype of Ceratocaulophryne regani: MOM Galerie: Level
3, Window 55, 38.0 mm; HIRONDELLE II Station 3279, west of
the Azores, 38° 55'N, 34° 07'W, 0-3000 m, 23 August 1912.

NONTYPE MATERIAL: LACM 11317-1, 54.0 mm; USNS
ELTANIN Station 1825, 39° 58'S, 160° 34'E, 3-m IKMT,
0-1625 m, 0218-0613 hr, 5 December 1966. LACM 33924-1,

54.0 mm; VELERO IV Station 17668, 32° 42'N, 118° 13'W; 3-m
IKMT, 0-750 m, 0947-1505 hr, 28 June 1972. LACM 6844-3,

41.0 mm; VELERO IV Station 7325, 33° 36'N, 118° 24' W, 3-m
IKMT, 0-640 m, 1129-1420 hr, 9 March 1961 (cleared and
stained).

IOS uncatalogued, 68.0 mm; DISCOVERY II Station 7480, 39°
55.2'N, 20° 07.4'W, RMT, 1250-1510 m, 0921- 1321 hr, 7
November 1970 (cleared and stained). IOS uncatalogued,

36.0 mm; DISCOVERY II Station 8281-43, 31° 44'N, 63°
58'W, RMT-8, 1235-1260 m, 1358-1658 hr, 20 March 1973.

USNM 148469, ca. 75 mm; ALBATROSS Station 2530, 40°
53'N, 66° 24'W, 14 June 1885. USNM 216461, 54.0 mm;
OCEAN ACRE Cruise 10, Station 21N, 31° 27'N, 64°49'W, 3-m
IKMT, 0-880, 1955-0135 hr, 6 June 1970.

ROM 27250, 109 mm; BRAN DAL Tow 3, 50° 30'N, 49°
30'W, Engel Trawl, 0-1070 m, 0710-1000 hr, 12 July 1968.

MCZ 51279, 23.0 mm; ANTON BROUN Cruise 6, Station
339B, APB 7231, 04° 14'S, 65° 02'E, 3-m IKMT, 0-275 m,
bottom depth 3900 m, 0715-1610 hr, 30 May 1964.

DIAGNOSIS: In addition to differences in escal morphology
(Pigs. 17, 19, 21), C. jordani is distinguished from C. pelagica
and C. polynerna in having fewer upper jaw teeth (Fig. 18). The
dorsal fin of C. jordani has a greater number of rays than that of
C. pelagica, and fewer rays than that of C. polynerna (Table 1).
It is further distinguished from C. pelagica in having a slightly
shorter, filamentous illicium; illicial filaments are absent in C.
pelagica , but considerably more numerous in C. polynerna (Figs.
17, 19, 21).

DESCRIPTION: Illicium pigmented except near esca, with 5
to 14 elongate (up to 87 percent of illicium length), translucent

filaments along entire length, all but distal-most arising from pos-
terior margin; esca with a posterolateral appendage bearing
numerous short filaments and having a palmate (usually bilobed),
opaque, distal tip; two distal appendages: the posterior- most
slender, bearing numerous filaments, the anterior-most short and
stout, opaque along posterior margin, and bearing numerous
filaments (separation into two distal appendages not as evident in
specimens less than 40.0 mm); and an elongate, filamentous,
anterolateral appendage (illicial and escal filaments lost in holo-
type. see Regan and Trewavas 1932:101, fig. 159; Fig. 19).

Total number of teeth in lower jaw 16-26, in upper jaw
20-39; vomerine teeth 2-5; D. 16-19, A. 14-18, P. 16-18
(Fig. 18, Table 1).

Measurements in percent of SL: longest tooth in lower jaw
6. 1-9.3; longest dorsal ray 78-115; longest anal ray 82-110;
illicium 17.6-33.9 (Fig. 16).

Rest of characters as for genus and family.

DISTRIBUTION: Caulophryne jordani is known from all
three major oceans of the world: seven records from the North
Atlantic, one from the Indian Ocean, one from the Southern
Ocean at approximately 40°S, 160°E, and two from off southern
California. The two IOS specimens from the north Atlantic (36.0
and 68.0 mm) were collected with a closing trawl between 1235
and 1260 m, and 1250 and 1510 m, respectively (Fig. 24).

COMMENTS: The illicial apparatus of the holotype of C. jor-
dani is somewhat damaged as indicated by a drawing provided by
Regan and Trewavas (1932, fig. 159), and as noted by Bertelsen
(1951:36). Filaments normally found along the length of the illic-
ium and arising from the esca are absent. The posterolateral escal
appendage and two distal appendages, however, remain intact. In
these and all other characters the additional material (with a
minor exception, see below) here assigned to C. jordani com-
pares very well with the holotype.

Bertelsen (1951:37) believed that Ceratocaulophryne regani
and Caulophryne polynerna might be synonyms on the basis of
similarity in illicial morphology, “position of the fins, and rela-
tive dimensions.” This conclusion, however, is not supported by
data provided by Jprgen Nielsen of the Zoological Museum,
University of Copenhagen. In illicial and escal morphology, jaw
tooth counts, and in dorsal and anal fin ray counts (16 and 16,
not 15 and 14, respectively, as given by Roule and Angel
1933), the holotype of C. regani compares well with the material
here recognized as C. jordani. Ceratocaulophryne regani is
removed from the synonymy of C. polynerna and placed within
that of C. jordani.

The 54.0 mm specimen of C. jordani from the Southern Ocean
(LACM 11317-1) differs from other individuals of this species
in having a higher number of jaw teeth (Fig. 18).

Caulophryne polynerna Regan 1930

Figures 16, 18, 20, 21, 24

Caulophryne polynerna Regan 1930:191-195, figs. 1-3 (original
description; single female specimen with parasitically attached
male). Parr 1930:131, fig. 4 (figure after Regan 1930). Regan and
Trewavas 1932:100, 101 (in key; description after Regan 1930).
Fowler 1936:1347 (in key; description after Regan 1930). Bertelsen
1951:36, 37, fig. 12, table 3 (figure after Regan 1930; a subspecies
of C. jordani). Pietsch 1976:789 (reproduction).

Caulophryne jordani polynerna Bertelsen 1951:33, 37, fig. 12, table 3 (in
key; description, comparison with all known material; includes

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 310:1-25.

Pietsch: Caulophrynid anglerfishes

36
34
32
30
28
26
24
22
20
18
16
14
12

10 20 30 40 50 60 70 80 90 100 110 120 130 140 150

i i i r

• L ACM 11317 1

i 1

A C. pelagica
• C. jordami
° C. polynema
0 C. sp.

_J L_

Standard length in mm

Figure 18. Number of teeth in upper and lower jaws versus standard length for Caulophryne species.

Ceratocaulophryne regani Roule and Angel 1932). Grey 1956:233
(synonymy; after Bertelsen 1951).

MATERIAL; Nine known females, 14.0- 142.0 mm.

Holotype of Caulophryne polynema: BMNH 1930.2.7.1, 142
mm; “long line, in deep water,” off Funchal Bay, Madeira; 1
February 1929.

NONTYPE MATERIAL: LACM 33923- 1, 103 mm;
VELERO IV Station 14962, 33° 10'N, 118° 31'W, 3-m I KMT,
1000 m wire, 0455-0943 hr, 10 February 1971. LACM

30619-3, 32.0 mm; VELERO IV Station 9903, 29° 30'N, 118°
54'W, 3-m IKMT, 0-834 m, 2315-0515 hr, 7 August 1964.
LACM 32772-1, 24.5 mm; TERITU Cruise Vanilla Fudge, Sta-
tion 69.11.4, 22° OO'N, 158° 00' W, 3-m IKMT, 0-1150 m,
1225-1635 hr, 12 November 1969. LACM 35897-1, 14.0 mm;
TERITU Station 74.5.7, 21° 20-30'N, 158° 20-30'W, 4-m
IKMT, 0- 1000 m, 1405- 1650 hr, 25 May 1974.

IOS uncatalogued, 1 10 mm; DISCOVERY II Station 8275, 32°
01.4'N, 50° 33.5'W, RMT, 0-1000 m, 2339-0150 hr, 5 March

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 310:1-25.

Pietsch: Caulophrynid anglerfishes

Figure 19. Esca of Caulophryne jordani, LACM 33924-1, 54.0 mm.

1973. IOS uncatalogued, 100 mm; DISCOVERY II Station
7856-50, 30° 03.7'N, 23° 00.2'W, RMT, 1000-1250 m,
0917-1317 hr, 5 April 1972. IOS uncatalogued, 11.5 mm; DIS-
COVERY II Station 7089-8, 17° 45'N, 25° 17'W, RMT,
900-1010 m; 0320-0659 hr; 13 November 1969.

ISH 3000/71, 44.0 mm; WALTHER HERWIG Station 510/71;
27° 18'N, 19° 44' W; CMBT-1600, 0-2000 m; 1752-2151 hr;
20 April 1971.

DIAGNOSIS: In addition to differences in escal morphology
(Figs. 17, 19, 21), C. polynema is distinguished from C. pelag-
ica and C. jordani in having a greater number of dorsal fin rays

(Table 1) and a highly filamentous illicium (Figs. 17, 19, 21). It
is further separated from C. pelagica in having a slightly shorter
illicium (Fig. 16), and from C. jordani in having more upper jaw
teeth (Fig. 18).

DESCRIPTION: Illicium lightly pigmented along anterior
margin (fully pigmented in 44.0 mm specimen), and bearing
numerous, elongate (up to 71 percent of illicium length), translu-
cent filaments, most of which arise from posterior margin; more
than 50 filaments in specimens larger than 32 mm, reduced to
approximately 10 in smaller specimens; esca consisting of a
posterolateral appendage with a palmate (usually bilobed),
opaque, distal tip, and a tapering terminal appendage with an
opaque distal tip; escal appendages highly filamentous in speci-
mens larger than 44 mm, number and length of filaments reduced
in smaller specimens (most escal filaments lost in holotype, see
Regan, 1930:193, fig. 2; Fig. 21).

Total number of teeth in lower jaw 19-32, in upper jaw
30-45; vomerine teeth 1-3; D. 19-22, A. 17-19, P. 15-18
(Fig. 18, Table 1).

Measurements in percent of SL: longest tooth in lower jaw
5. 6-9. 5; longest dorsal ray 70-155; longest anal ray 60-151;
illicium 16.0-32.1 (Fig. 16).

Rest of characters as for genus and family.

DISTRIBUTION: Five specimens of C. polynema have been
collected from the eastern north Atlantic Ocean as far west as
approximately 50° W. Four additional records are known from
the eastern north Pacific: two from Hawaii and two from off
southern California. The 100 mm IOS specimen from the eastern
north Atlantic was captured with a closing trawl between 1000
and 1250 m (Fig. 24).

COMMENTS: Caulophryne polynema was separated by Regan
and Trewavas (1932:100) from the other species because of its
higher number of dorsal and anal-fin rays. Bertelsen (1951:36)
however, noted that the high number of fin rays in caulophrynids
(compared to other ceratioids) has such a large individual varia-
tion (D. 14 to 22, A. 12 to 19) that “we can scarcely attach
much weight to this character.” The greater amount of material
available now indicates that despite this variation within the
genus, fin- ray counts are of significant taxonomic importance.
Caulophryne polynema can nearly always be separated from its
congeners by its greater number of dorsal and anal-fin rays (see
Key, Table 1).

Caulophryne species

Figures 16, 18, 22, 23

LACM 36025-1, female, 98.0 mm with parasitic male, 12.0
mm; ALPHA HELIX Southeast Asian Bioluminescence Expedi-
tion Station 37, Midwater Trawl 22, 4° 56'S, 129° 26'E, RMT-8,
0- 2000 m, 0320-0530 hr, 12 April 1975.

DESCRIPTION OF FEMALE: Illicium unpigmented with
approximately 30 elongate (up to 25 percent of illicium length),
translucent filaments along entire length, those on proximal half
of illicium arising from posterior margin; esca, a tapering
appendage bearing numerous simple filaments and four highly
branched lateral appendages: three, more proximal in position,
one, more distal; opaque areas absent (Fig. 22).

Gonads well developed (right ovary, 22 mm long or 22.4 per-
cent of SL, containing numerous eggs approximately 0.3 mm in
diameter) .

Total number of teeth in lower jaw 14, in upper jaw 22;

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 310:1-25.

Figure 20. Caulophryne polynema, LACM 33923-1, 103 mm. Drawn by Elizabeth Anne Hoxie.

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979 . 310:1-25.

Pietsch: Caulophrynid anglerfishes

Figure 21. Escae of Caulophryne polynema: A. IOS, 100 mm; B. IOS, 110 mm. Courtesy of G Brovard and E. Bertelsen.

vomerine teeth 1; D. 17, A. 15, P. 16-16 (Fig. 18, Table 1).

Measurements in percent of SL: longest tooth in lower jaw 9.2;
longest dorsal ray 95 + ; longest anal ray 95 +; illicium 27.6 (Fig.
16).

Rest of characters as for genus and family.

DESCRIPTION OF MALE; Upper and lower denticulars (Ber-
telsen 1951:21) embedded in an unpigmented, conical papilla on
belly of female; passageway from outside into pharynx and out
through gill openings retained (Pietsch 1976:790); eyes well
developed; nostrils degenerate; gills developed; testes large (2.3
mm long, 19.2 percent of SL); dorsal, anal and pectoral-fin ray
counts undetermined; pelvics absent; characteristic gelatinous
envelope of larvae absent; entire head and body lightly pigmented
(Fig. 23).

COMMENTS: This female and attached male cannot reason-
ably be placed within the material of any of the three recognized
species of Caulophryne. The female has the elongate, branched
escal appendages characteristic of C. pelagic a but also the illicial
filaments found only in C. jordani and C. polynema (compare
Figs. 17, 19, 21, 22). The illicium appears to be slightly longer
than that of C. jordani and C. polynema, but shorter than that of
C. pelagica (Fig. 16). Jaw tooth counts are at the bottom of the
range of variation for females of all species combined (Fig. 18).
Finally, fin-ray counts compare best with C. jordani (Table 1).

The attached male (Fig. 23) represents the second example of
sexual parasitism in the Caulophrynidae. It does not differ in any

significant way from the previously described specimen (Bertel-
sen 1951:37, fig. 12).

Caulophryne setosus Goode and Bean 1896

NOMEN NUDUM

Caulophryne setosus Goode and Bean 1896:26*, 541, fig. 409.

COMMENTS: This name appears in a figure caption, and is
entered in the list of plates and index of Goode and Bean’s
(1896) original description of Caulophryne jordani, without ap-
plication to a description or type. This error was caught by Jor-
dan and Evermann (1898:2735): “plate named C. setosus, by slip
in proof reading.” The name setosus also appears in the cata-
log of fishes of the National Museum of Natural History (S.J.
Kamella, personal communication).

Caulophryne racemosa Monod 1960

NOMEN NUDUM

Caulophryne racemosa Monod 1960:687, fig. 80.

COMMENTS: This name appears in a figure illustrating stages
of specialization of lophiiform pectoral radials. There is no appli-

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 310:1-25.

Pietsch: Caulophrynid anglerfishes

cation to a description or type. No doubt, Monod (1960) meant to
refer to C. ramulosa. His figure was taken from Regan and Tre-
wavas’ (1932, fig. 58) illustration of the pectoral radials of the
holotype of this nominal species.

DISCUSSION

The discovery of yet another new ceratioid genus brings the
total number of genera based on females to 34. Although new
forms continue to be described, it is unlikely that the total number
of ceratioid species will increase significantly (Bertelsen
1951:28). In the last ten years 19 forms have been resurrected
from synonymy or described as new (Pietsch 1969, 1972b, 1973,
1974a, b, 1975, 1978; Bertelsen 1973; Nolan and Rosenblatt
1975). During this same time, 28 nominal forms have fallen into
synonymy, thus actually decreasing the number of recognized
forms.

More so than any other ceratioid group, the caulophrynids
exhibit a confusing mosaic of primitive and derived character
states. Utilizing the variation found within the entire order Lophi-
iformes as representing the ancestral character pool for the
Caulophrynidae, the following character states that describe
Caulophryne are considered primitive;

1. frontal bones short, posterior in position, meeting on
midline in front of supraoccipital (this is the condition of
the frontals in lophioids and antennarioids; and, in addi-
tion to caulophrynids among ceratioids, the Neocerati-
idae).

2. illicial trough absent (a narrow depression, usually run-
ning the full length of the cranium, is present in most
ceratioids to receive the pterygiophore of the illicium; it
is absent in caulophrynids and neoceratiids as well as in
lophioids and antennarioids).

3. dorsal ceratohyal process present (this process is present
in lophioids and antennarioids, but absent in all other
ceratioids).

4. single epural present (no trace of an epural is found in
any other ceratioid; lophioids and antennarioids, except
for brachionichthyids, have one epural).

5. high number of median-fin rays (although Caulophryne
exhibits a slight increase over the already high number of
median-fin rays characteristic of lophioids and antennari-
oids, this probably represents a primitive condition over
the greatly reduced median- fin ray counts of most other
ceratioids; exceptions include some gigantactinids, neo-
ceratiids, and melanocetids).

6. median-fin rays exceptionally long (like those of larval
and young lophioids and antennarioids, but unlike any
other ceratioid).

7. sexual dimorphism in the illicial apparatus of larval
stages possibly absent (the 16 known caulophrynid larvae
all have a well developed outer illicial rudiment, a struc-
ture normally found only in female ceratioids; the sex of
these larvae cannot be determined from examining the
gonads, but since it seems unlikely that only one sex is
represented, Bertelsen (1951:31) suggested that the sex-
ual dimorphism characteristic of other larval ceratioids,
but absent in lophioids and antennarioids, might be
absent in caulophrynids as well).

8. escal bulb absent (the esca of caulophrynids is not a
bulbous, bacteria-containing organ as in other ceratioids

but a tuft of filaments like that of many lophioids and
antennarioids; in addition, Bertelsen (1951:31) stated that
the illicium of caulophrynid larvae “is not globular or
club-shaped as in most other female ceratioid larvae, but
more finger- like as in certain antennarioid larvae.”

9. posteroventral coracoid process present (this process is
also found in antennarioids; among ceratioids it is present
in ceratiids, in the oneirodid genera Spiniphryne and
Oneirodes, and present, but cartilaginous in gigantacti-
nids).

10. pectoral fins exceptionally large, especially in larvae and
young (like those of larval and young lophioids and
antennarioids, as well as larval gigantactinids among
ceratioids; Bertelsen 1951:35, 148, figs. 11, 99, 104).

11. pelvic fins retained in larval stages (pelvic fins, well
developed in all lophioids and antennarioids, are not
present in larvae or adults of any other ceratioid).

Caulophrynids are considered derived in having the following
character states:

1. parietals anterior in position, far removed from posttem-
porals and expanded laterally.

2. pterosphenoid absent (also absent in linophrynids and
gigantactinids).

3. anterior maxillomandibular ligament absent (also absent
in linophrynids, gigantactinids and neoceratiids).

4. dorsal hypohyal reduced (absent in Gigantactis).

5. hyoid apparatus narrow and elongate (similar in lino-
phrynids, gigantactinids, and neoceratiids).

Figure 22. Esca of Caulophryne sp., LACM 36025-1, 98.0 mm.

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 310:1-25.

Pietsch: Caulophrynid anglerfishes

Figure 23. Caulophryne sp., parasitic male, LACM 36025-1, 12.0 mm.

6. opercular bones reduced (as are those of linophrynids,
gigantactinids, and neoceratiids).

7. hypobranchials and basibranchials absent (all hypobran-
chials are absent in linophrynids and gigantactinids;
hypobranchial II is absent, but a small hypobranchial III
is present in neoceratiids; all basibranchials are absent in
ceratiids, linophrynids, gigantactinids and neoceratiids).

8. caudal rays 8 (ceratiids also show a reduction in the
number of caudal rays to 8; all other ceratioids have 9;
neoceratiids occasionally have 10, Bertelsen 1951:159,
table 36).

9. posttemporals reduced, overlapping pterotics and
epiotics, but far removed from parietals and exoccipitals
(similarly reduced in gigantactinids).

10. lower portion of cleithrum reduced to a slender thread of
bone (similarly reduced in Gigantactis) .

1 1 . pectoral radial number reduced by fusion to 2 (all other
adult ceratioids have 3 or more radials).

12. neuromasts of acoustico-lateralis system located at the
tips of elongate filaments (this specialization is unique to

caulophrynids; Regan and Trewavas 1932:23, 24, fig.
16C).

13. illicial filaments present in some species (also present in
some species of Gigantactis).

14. males known to become sexually parasitic (Pietsch
1976:788; reproductive mode shared by ceratiids, lino-
phrynids, neoceratiids, and the oneirodid genus Lep-
tacanthichthys).

Due to the presence of a number of the primitive character
states listed above (7, 8, 11), Bertelsen (1951:28) placed the
Caulophrynidae “first in the suborder.” At the same time, he
argued that the Linophrynidae showed “such a highly specialized
condition that they must be placed last.” Since members of both
these families are characterized by having parasitic males, Bertel-
sen’s classification requires that this mode of reproduction evolved
more than once. For unstated reasons, Greenwood, et al.
(1966:397) implied a much closer relationship between the
caulophrynids and linophrynids. Likewise, at the 1975 Annual
Meeting of the American Society of Ichthyologists and Herpe-
tologists in Williamsburg, Virginia, I reported finding numerous,
derived character states shared by these two families as well as
with the Gigantactinidae and Neoceratiidae. Ignoring all primi-
tive character states at that time, I presented a phylogeny that
argued for a monophyletic origin of sexual parasitism within a
lineage derived from some oneirodid- like ancestor (Fig. 25; see
also Pietsch 1976:791). This more detailed study of the Caulo-
phrynidae has revealed a considerable number of additional
resemblances (albeit, primitive states) to less derived lophiiforms
(lophioids and antennarioids) that can hardly be ignored. That
these primitive character states suddenly reappeared in a lineage
that arose from an ancestor derived in all, is highly improbable;
the importance of recognizing and utilizing primitive character
states in phylogenetic analysis is apparent.

A return to Bertelsen’ s (1951) classification is proposed in

Figure 24. Known distribution of species of Caulophrynidae. A single symbol may represent more than one capture.

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 310: 1—25 .

Pietsch: Caulophrynid anglerfishes

Figure 25. Phylogenetic relationships of families of Ceratioidei based on shared derived character states. Starred families contain species known to have
parasitic males.

Caulophrynidae* Ceratiidae* Gigantactinidae

Neoceratiidae* Linophrynidae* Oneirodidae* Thaumatichthyidae Centrophrynidae

Diceratiidae Himantolophidae

Melanocetidae

Figure 26. Phylogenetic relationships of families of Ceratioidei based on shared primitive as well as derived character states. Starred families contain
species known to have parasitic males.

which the Caulophrynidae forms the primitive sister-group of the
Melanocetidae, Himantolophidae, Diceratiidae, Centrophrynidae,
Gneirodidae and Thaumatichthyidae (Fig. 26). The phylogenetic
position of the closely related families Linophrynidae, Gigantac-
tinidae and Neoceratiidae (and perhaps the Ceratiidae) is
unclear. All share numerous derived character states among
themselves and with the Caulophrynidae; but, most of these are
either reduction states or loss of parts, conditions that are found

in deepsea groups in general that could well be convergent in
origin. This seems to leave two alternatives: either this assem-
blage is derived from a caulophrynid-like ancestor, thus forming
part of a sister-group that is primitive to all other ceratioids , or it
is derived from some oneirodid-like ancestor. Clarification of
these problems must await further comparative studies; but, regard-
less of the phylogenetic position of the Linophrynidae and its
allies, this interpretation, while recognizing the basal position of

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 310:1-25.

Pietsch: Caulophrynid anglerfishes

the Caulophrynidae, indicates a return to Bertelsen’s (1951) view
that sexual parasitism has evolved independently in at least two
separate ceratioid lineages.

Three characters are available for interpreting intrafamiliar
relationships:

1. median-fin ray counts. Within Caulophryne there is a
trend toward an increase in the number of dorsal and
anal-fin rays over the already high number of rays charac-
teristic of lophioids and antennarioids. Robia, on the other
hand, has a greatly reduced number of median-fin rays.
The extremes of these opposite trends are both considered
derived character states.

2. illicium length. Elongation of the illicium is a trend found
in several other ceratioid groups (Pietsch 1972b, 1974a, b,
1975). The short illicia of Caulophryne species is a primi-
tive condition; the extremely elongate illicium of Robia is
derived.

3. illicial filaments. Within other ceratioid groups there is a
trend toward an increase in morphological complexity of
the illicial apparatus (Pietsch 1972b, 1974a, 1975). Fila-
ments present along the length of the illicium of C. jor-
dani and C. polynema (absent in all other ceratioids except
for some species of Gigantactis) is considered a derived
state.

This character analysis indicates that Robia, with its greatly
reduced median-fin ray counts and extremely elongate illicium, is
the more derived of the two caulophrynid genera. Caulophryne
pelagica has a longer illicium than its congeners, but is consid-
ered the least derived member of the genus, having the lowest
median-fin ray counts and lacking illicial filaments. Caulophryne
jordani and C. polynema appear to be more closely related to
each other than either is to C. pelagica; both exhibit an increase
in the number of median-fin rays and share the presence of illicial
filaments. Caulophryne polynema is the most derived member of
the group having the highest median-fin ray counts and the most
complex illicial structure.

ACKNOWLEDGMENTS

I thank E. Bertelsen for critically reading the manuscript. The
following people and their institutions provided material: A.
Wheeler (BMNH); N. Merrett and J. Badcock (IOS); G. Krefft
(ISH); R. J. Lavenberg and J.W. Neumann (LACM); K.F. Liem
and K. Hartel (MCZ); W.B. Scott and A. Emery (ROM); R.H.
Gibbs and S. Kamella (USNM); E. Bertelsen and J. Nielsen
(ZMUC); T.A. Clarke (Hawaii Institute of Marine Science); and
B.H. Robison (Marine Science Institute, University of California
Santa Barbara). Additional thanks go to J. Nielsen and M.L.
Bauchot for providing data on the holotype of Ceratocaulophryne
regani. Figure 14 was made by Caryl Maloof and figure 20 by
Elizabeth Anne Hoxie. Geert Brovad (ZMUC) provided photo-
graphs used in figure 2 1 .

The work was supported by National Science Foundation
Grants GB-40700 and DEB 76-82279. Partial assistance from the
Johannes Schmidt Fund of the University of Copenhagen and the
Office of Graduate Studies and Research, California State Uni-
versity, Long Beach, is also gratefully acknowledged.

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Lavenberg, R.J. and A.W. Ebeling. 1967. Distribution of
midwater fishes among deep water basins of the southern
California shelf. Proc. Symp. Biol. Calif. Islands, R.N.
Philbrick, Ed., Santa Barbara Botanical Garden, Inc., pp.
185-201.

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 310:1-25.

Pietsch: Caulophrynid anglerfishes

Le Danois, Y. 1964. Etude anatomique et systematique des
antennaires, de l’ordre des Pediculates. Mem. Mus. Nat.
Hist. Nat., Ser. A, Zool. 31:1-162.

Maul, G.E. 1973. In Hureau, J.C. and T. Monod, eds. Check-
list of the Fishes of the North-eastern Atlantic and of the
Mediterranean, UNESCO, 1:666.

Monod, T. 1960. A propos du pseudobrachium des Antennarius
(Pisces: Lophiiformes). Bull, de l’lnst. Frangais d’Afrique
Noire 22, Ser. A, 2:620-698.

Nolan, R.S. and R.H. Rosenblatt. 1975. A review of the
deep-sea angler fish genus Lasiognathus (Pisces: Thauma-
tichthyidae). Copeia 1975:60-66.

Nybelin, O. 1963. Zur Morphologie and Terminologie, des
Schwanzskelettes der Actinopterygier. Ark. Zool. 15(35):
485-516.

Parr, A.E. 1930. On the probable identity, life-history and
anatomy of the free-living and attached males of the
ceratioid fishes. Copeia 1930(4): 129- 135.

Pietsch, T.W. 1969. A remarkable new genus and species of
deep-sea anglerfish (family Oneirodidae) from off Guada-
lupe Island, Mexico. Copeia 1969:365-369.

. 1972a. A review of the monotypic deep-sea anglerfish

family Centrophrynidae: taxonomy, distribution and oste-
ology. Copeia 1972:17-47.

1972b. Ergebnisse der Forschungsreisen des FFS

“Walther Herwig” nach Siidamerika. XIX. Systematics and
distribution of ceratioid fishes of the genus Dolopichthys
(family Oneirodidae) with the description of a new species.
Arch. FischWiss. 23(1 ): 1— 28 .

. 1973. A new genus and species of deep-sea anglerfish

(family Oneirodidae) from the northern Pacific Ocean.
Copeia 1973:193-199.

. 1974a. Osteology and relationships of ceratioid

anglerfishes of the family Oneirodidae, with a review of the
genus Oneirodes Liitken. Nat. Hist. Mus. Los Angeles Co.,
Sci. Bull. 18:1-113.

. 1974b. Systematics and distribution of ceratioid

anglerfishes of the genus Lophodolos (family Oneirodidae).
Breviora 425:1- 19.

. 1975. Systematics and distribution of ceratioid angler-
fishes of the genus Chaenophryne (family Oneirodidae).
Bull. Mus. Comp. Zool. 147(2):75- 100.

1976. Dimorphism, parasitism, and sex: reproductive

strategies among deep-sea ceratioid anglerfishes. Copeia
1976(4):78 1-793.

. 1978. A new genus and species of deep-sea anglerfish

from the eastern north Pacific Ocean with a review of the
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Regan, C.T. 1912. The classification of the teleostean fishes of
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. 1930. A ceratioid fish ( Caulophryne polynema, sp.

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(Ceratioidea). Dana Rept. 2., 113 pp.

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Accepted for publication January 5, 1978.

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 310:1-25.

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CONTRIBUTIONS IN SCIENCE

NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY

CONTENTS

Abstract

Paltostoma Schiner

Paltostoma costaricensis new species

Paltostoma delectata Alexander

Paltostoma diriageni new species ....

Paltostoma exserta new species

Paltostoma zwicki new species

Acknowledgments

Resumen

Literature Cited

Figures 1-40

... 3
... 3
... 4
... 5
... 6
.... 7
.... 8
.... 8
.... 9
.... 9
10 ff

ISSN: 0459-8113

Suggested Citation: Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 311:1-22.

Contributions in Science are articles in the earth and life sciences, presenting results of original research in Natural
History. Science Bulletin (Numbers 1-30; 28 June 1965 to 10 April 1978) and Contributions in Science (Numbers 1-300;
23 January 1957 to 16 June 1978) were merged into a single imperial octavo serial beginning with Number 301.

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 311:1-22.

THE FAMILY BLEPHARICERIDAE IN COSTA RICA

(D1PTERA)1

By Charles L. Hogue2

ABSTRACT: Five species of net-winged midges of the family Blephariceridae are recorded for Costa Rica.
All belong to the genus Paltostoma; one was previously known ( delectata Alexander) but the remaining are
described here as new species ( costaricensis , diriageni, exserta and zwicki). The paper includes also a
diagnosis and biologic notes on the genus.

The Blephariceridae or “net-winged” midges are notable in
the extreme adaptations they exhibit for life in torrential water.
Six mid-ventral suckers permit the larvae to hold fast and
travel from place to place on smooth rock surfaces in the
fastest currents, in waterfalls and cataracts; the pupae are
attached permanently to the substratum by ventrolateral adhesive
pads. The adults are capable of emergence while totally sub-
mersed. This takes place through an explosive egress of the body
from the pupal case and unfolding of the appendages, followed
by immediate flight. Such rapid escape is made possible by prior
development and expansion of the wings and legs within the
pupal case. The process requires compression and folding of the
complete wing and causes numerous creases to form in its mem-
brane, giving it the reticulate appearance from which the family
receives its common name.

Although occurring widely in the Neotropical Region, the
Blephariceridae are seldom collected. Consequently, our knowl-
edge of species distributions is fragmentary and the family’s
regional taxonomy is in a rudimentary state (Hogue 1971). Five
genera are presently recognized (including the very primitive
Edwardsina, with species also in Australia), all apparently hav-
ing austral origins. Of these, evidently only Paltostoma has
extended northward through Central America, penetrating Mex-
ico along the Sierra Madre Occidental to latitude 21° N (unpub-
lished record). The genus also occurs on several Caribbean
islands.

The Costa Rican fauna includes 5 species. The first known was
delectata, described by Alexander in 1953 from 2 males, and
which, until my recent collecting, remained the sole regional rep-
resentatives of the family. This paper adds 4 additional species
and provides descriptions and illustrations by which all may be
recognized. Because little collecting has been accomplished and
speciation seems to have occurred between populations that are
only narrowly separated geographically, there remain strong pos-
sibilities for discovery of still other forms.

The descriptions of the adults are based largely on pharate
specimens dissected from their pupal cases. Therefore, measure-
ments of legs and wings, as well as color pattern characters, are
not determinable. Quantitative data are given in millimeters and

represent means calculated from small samples, hence ranges are
omitted. Terminology follows that of Hogue 1973 and 1978.

Paltostoma Schiner

Paltostoma presently contains 13 named species, including
those newly described in this paper. Material of several more
exists in collections and, surely, more will be discovered when
streams are sampled in the vast Andean Mountain system. The
genus is characterized as follows:

DESCRIPTIONS

Adult

COLORATION: Generally well sclerotized; velvety brown
with sharply defined, silvery (“frosty”) or opalescent reflections
which vary in position depending on the angle of viewing.

SIZE: Small to medium-sized Blephariceridae; wing length
4.5-9 mm.

HEAD: Small. Normal type. Suprafrons very broad, flat;
frontal carina absent. Parietal sclerite very narrow. Ocelli sessile,
separate, on slightly elevated boss. Eyes disjunct; undifferen-
tiated into upper and lower divisions; ommatidia all of approxi-
mately equal size.

MOUTHPARTS: Male. — Proboscis very long (PL/HW 2.5).
Labium jointed near base. Labrum rigid, slender; apex attenuate.
Margin of hypopharynx entire, a few spicules apically. Maxillary
lobes very long, approximately half the length of labium. Label-
lar lobes small, length slightly greater than width of labium.
Palpus very short, segments 1-3 fused, 4-5 absent; sensory pit

'Review Committee for this Contribution:

Charles P. Alexander
Julian P. Donahue
Roy R. Snelling
Peter Zwick

2Senior Curator, Section of Entomology, Natural History Museum of Los
Angeles County, 900 Exposition Boulevard, Los Angeles, California
90007.

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 311:1-22.

Hogue: Costa Rican Blephariceridae

of segment 3 terminal. Mandibles absent. Female. — Proboscis
moderately long (PL/HW 1.4). Labium jointed at about mid-
length. Labrum rigid, broad, apex acute. Lateral margin of hypo-
pharynx serrate, apex with lateral hair- like projections. Maxillary
lobes short, slightly less than half the length of labrum. Labellar
lobes very short, length less than width of labium. Palpus very
short, segments 1-3 distinct, 4-5 absent; sensory pit of segment
3 terminal. Mandibles present.

ANTENNA: Length 1.5- 1.8 head width, that of male slightly
longer proportionately (1.8- 1.9); 15-segmented. Scape globular,
pedicel canopiform3, flagellomeres cylindriform, length of latter
about 2.0 width; flagellar segments 1-5 each with a small group
of placoid sensilla dorsally.

WING: Venation (Fig. 14) reduced: R2, R3, M3, and basal
sector of M3+4 absent; 1A reduced; R2_5 long; branch of R4 and
R5 near apex of wing. Dorsal macrotrichia on vein Rs only, a
few also ventrally at apex of this vein. Wing shape broad (L/W
2.9), anal lobe moderately developed.

LEGS: Tibial spur formula 0-0-1 in both sexes, hind tibial
spur large, minutely spiculate on inner surface. Tarsal segment 5
longer than 4, a group of spiniform setae ventrobasally. Tarsal
claws each with a single, large sub-basal tooth.

ABDOMEN: Short, strongly tapered posteriorly.

MALE GENITALIA: Segments VIII and IX unmodified. IXth
tergite lobes triangular, a strong ventral armature produced from
the lateral margin of each lobe, its arms converging anteriorly.
Basistyle elongate, longer than dististyle. Outer dististyle broad,
rectangular, variously incised and setose. Inner dististyle undi-
vided, a long, curved, irregular finger. Phallosome vesica trian-
gular; apodeme longer than vesica, a vertical, oval flange. Para-
meres undeveloped as separate, distinct tubular structures, their
morphological position assumed by a broad, apically reflexed
plate (“ventral plate”4 of Stuckenberg 1958:101, see Fig. 9).
Penis filaments 3, all simple, very long, straight rods, without
special apical modifications. Tegmen narrow, anchor- shaped;
base small, forming a shallow subanal pouch; apex broadly spat-
ulate; margin recurved. Lateral portion of ventral bridge (Fig. 8)
forming a complex curved plate weakly continuous with the ven-
tral bridge proper.

FEMALE GENITALIA: VUIth stemite lobe elongate, median
fold deep, wide, Oviscapt elongate, apical lobes extended latero-
posteriorly, a ventral, transverse fl-shaped apodeme present in
inner piece. Spermathecae ovoid; ducts very long, exit from body
of spermatheca laterally, their proximal sections coiled 1 to sev-
eral times around body before leading to gonotreme. Sensilla of
cercus all chaetiform, apical tubular sensilla absent; a short, sub-
apical, ventral finger-like lobe extending posteriorly from the
cercus.

CHAETOTAXY: Body without long hairs. Setal groups mini-
mal and inconspicuous. Occipital bristles of head short, scattered
and not differentiated into upper and lower series. Postgenal
bristles very few (1-3).

Larva

Intercalary convexities undeveloped. Head capsule with deep
lateral incisions. Antenna short, 2- segmented. Dorsal pseudopods
and sclerotized major plates and processes absent. Dorsal integ-
ument densely striate. Ventral gill filaments stellate in arrange-
ment, 9-10 (usually 10) in number. Primary trunk sensilla
recognizable as follows: ic (on anterior margin of division, rather
than at level of intercalary portion), short, chaetiform; very small
it (in normal positions on divisions II- VI); moderately long

chaetiform sensilla st (P, I-V), psu (M-T), tp (II- V), ss (P-T,
in line) and pd (I- VI). Secondary sensilla in two fields on dor-
sum: small, scattered capitate to conoid types generally and simi-
larly shaped but much larger types in linear series antero- and
posterolateral Iy on each division, the latter often flattened and
tending to close interlobular spaces dorsally. Terminal setae
reduced in size and number to 1-2.

Pupa

Elliptical to oval in outline, cross-section hemi-oval, flat. Lat-
eral margins underfolded. Dorsal cuticle of all sclerites except
cephalic and alar densely set with long spines; those middorsally
flanked basally with 2 or more very short spinelets (spines often
lacking in specimens suffering from rough handling or stream
abrasion). Respiratory processes erect, disjunct at base; individ-
ual plates (lamellae) rigid, triangular, with acute apices; inner
plates smaller and weaker than outer. Antennal cases short,
extending to a level less than apex of the mouthpart cases.
Ventral adhesive pads four in number.

DISTRIBUTION: Restricted to Neotropical Region between
latitudes 25° S and 21° N, Pacific Cordillera and Caribbean Is-
lands. See the map (Fig. 40) for the distribution of species in
Costa Rica.

Paltostoma costaricensis new species

Figures 1-2, 5-9, 25, 27-29, 37, 40

DIAGNOSIS

This species may be distinguished by the following combina-
tion of characteristics of the male genitalia: long penis filaments
with gradually widened bases. Broadly rounded, improminent
IXth tergite lobe; arms of ventral armatures of the lobes contigu-
ous and fused anteriorly. Outer dististyle incised apicoventrally,
the dissected portion forming a thumblike projection.

It is also decidedly smaller than any other Costa Rican Palto-
stoma. Although comparable measurements for most structures
are not available because of the lack of fully developed speci-
mens of both sexes of all the species, the fact is readily apparent
from general inspection. Head widths and lengths of wing vein
Rs, characters which could be compared among all species, indi-
cate that costaricensis is approximately 20% smaller than either
zwicki or delectata and 40% smaller than diriageni and exserta.
DESCRIPTIONS

Male (Figs. 1, 5-9)

A composite description, based on four pharate specimens dis-
sected from their pupal cases:

COLORATION: Not determinable, probably similar to that of
female (see below).

SIZE: A small Paltostoma. Measurements as follows (from 3
pharate specimens): Wing and leg dimensions not determinable.
Head width 0.66. Clypeus-labrum length combined 1.76. Palpal
segments 1-3 combined 0.20.

HEAD (Fig. 1): Palpal segment proportions indeterminable,
fusion of segments 1-3 complete, no perceptible constrictions or
joints. Antennal segment proportions 1.3, 1.6, 1.00, 1.0, 1.1,

3Shape like a typical Egyptian canopic jar.

4This homology is questionable, however. Comparative anatomical
studies are needed; the term is used here for convenience only.

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 311:1-22.

Hogue: Costa Rican Blephariceridae

1.1, 1.1, 1.1, 1.1, 1.1, 1.1, 1.1, 1.1, 1.1, 1.4. Ultimate segment
1.3 length of penultimate.

LEGS: Segment proportions not available.

WING: Venation and trichia typical for genus.

GENITALIA (Figs. 5-9): IXth tergite lobe improminent, apex
broadly rounded; arms of ventral armature contiguous anteriorly.
Outer dististyle incised apicoventrally, the dissected ventro-
median portion forming a thumb-like projection. Inner dististyle
simple, smoothly curved and tapered, slightly capitate. Tegmen
elongate, narrow, with two parallel straplike sclerotizations; basal
portion narrow, articulating with the contiguous arms of the ven-
tral armature of the IXth tergite lobes . Phallosome totally lacking
parameres; penis filaments evenly tapered, only slightly widened
basad.

Female (Figs. 2, 25)

Based primarily on specimen from Golfito; head structures
from Rincon series:

COLORATION: Head. — All sclerites medium-brown, darker
towards vertex and posteriorly, with extensive velvety texture
and silvery opalescent reflections; ocellar triangle dark-brown.
Mouthparts and antenna dull medium-brown, the latter darker.
Thorax. — Generally medium-brown. Scutum, scutellum, anterior
pronotum and ventral portions of epistemum and meron darker;
extensive silvery opalescent reflections as on head; remaining
sclerites and membranous portions dull yellow-brown. Disc of
scutum with dark-brown, parallel, median vittae separated from a
more posterior, broad, V-shaped, dark-brown vitta bordering the
transverse suture. Legs. — Generally dull medium-brown, lighter
basad, darker abruptly distad (before apices of tibiae), including
tarsi and claws. Wing. — Membrane hyaline; veins brownish-
black. Haltere. — Stem light-brown basad, darkening distad;
knob velvety gray-brown. Abdomen. — Color characteristics not
available.

SIZE: Measurements as follows (from 5 pharate and one
mature specimen): Body length not determinable. Wing length
6.8, width 2.3. Head width 0.84. Clypeus length 0.39. Labrum
length 0.67. Palpal segments 2-3: 0.16, 0.22. Leg segment
lengths as below (from Golfito specimen):

fore

mid

hind

femur

4.4

5.7

7.1

tibia

5.4

5.5

6.3

tarsus 1

1.89

2.00

2.06

0.79

0.86

0.43

0.41

0.44

0.24

0.26

0.28

0.21

0.28

0.26

0.25

HEAD (Fig. 2): Palpus very short, segments 1-3 distinct,
proportions 2-3: 1.0, 1.4. Antennal segment proportions: 1.0,
1.5, 1.00, 1.1, 1.1, 1.1, 1.1, 1.1, 1.1, 1.1, 1.1, 1.1, 1.0, 1.1,
1.8; ultimate segment 1.6 length of penultimate.

LEGS: Progressive segment proportions as below:

foreleg

1.0

1.2

0.4

0.5

0.6

1.1

midleg

1.0

0.4

0.5

0.6

0.9

hindleg

1.0

0.9

0.3

0.2

0.6

0.9

1.2

WING: Venation and trichia typical for genus.

GENITALIA (Fig. 25): Oviscapt broad, ratio length/width =
1.3. Setae of VHlth sternite lobe short, 10- 14 in number.

Pupa (Fig. 37)

SIZE: Small. Measurements (N=10): male, length 2.9, width
1.7; female, length 4.3, width 2.5.

STRUCTURE: Outline elongate-oval, L/W 1.7, strongly con-
vex, cross section hemispherical, peripheral margin rounded.
Cephalic sclerite strongly underfolded (to level of tips of palpus
case). Cuticular spines proportionately long, longest lateral
spines about as long or slightly longer than mid-dorsal length of
scutellar sclerite; numerous spines in mesoposterior region of
scutum; parabasal spinelets, when present, one on each side of
the major spine and mostly of equal length. Lamellae of respira-
tory processes divergent, no. 1 much so, projecting anteriorly
beyond body margin; subequal in length; outer pair heavier than
inner and triangular; inner pair 0.6- 0.7 width of outer at their
bases, no. 2 the smallest; apex of no. 3 slightly attenuate and
tortuate, curving around inner margin of no. 2.

Larva (Figs. 27-29)

SIZE: Small. Measurements (N=10): Body length 5.0; head
capsule width 1.26.

CHAETOTAXY (trunk segments only): Secondary dorsal sen-
silla all small and coniform; those of antero- and posterolateral
linear series larger and coniform to clavate, the latter more poorly
developed dorsomesad.

STRUCTURE: Pseudopods short, L/W about 2.3, arising from
ventrolateral depression. Posterolateral lobe of anal division
poorly developed, apex directed caudad. Lateral margin of anal
division entire, a cluster of secondary coniform sensilla with
large alveoli situated on a slight boss anterior to posterolateral
lobe. Posterior extreme of anal segment not excessively pig-
mented nor sclerotized, the margin a smoothly rounded arc con-
tinuous with lateral margin. Ventral integument mesal to base of
pseudopod strigose over only a small area immediately mesal to
insertion of pseudopod.

MATERIAL AND TYPE LOCALITY:

Types. COSTA RICA, Puntarenas Province, 1.8 miles W Rincon, Lin-
guita Creek, 11 March 1971, C.L. Hogue and J.R Donahue: HOLOTYPE 6
(pharate, dissected from and mounted entire on slide series No. CLH 73-
16a-e: LACM); ALLOTYPE 9 (pharate, dissected from and mounted
entire on slide series No. CLH 73-156a-e: LACM); 66 , 5 9 PARATYPES
(pharate adults mounted entire on slide series).

Additional. COSTA RICA, Alajuela Province, 0.7 miles W Grecia, Rio
Vigia, 18 June 1972, C.L. Hogue (1 larva: LACM). Puntarenas Province,
Golfito, 15 August 1957, A.S. Menke (19: LACM). Las Cruces, (rock
quarry) 6 miles S San Vito de Jaba, Rio Jaba, 22 March 1967, R.W.
McDiarmid, RWM 21 (2 pupae, 14 larvae: LACM). 27 June 1972, C.L.
Hogue (2 pupae, 21 larvae: LACM). 1.8 miles W Rincon, Linguita Creek, 3
July 1963, C.L. Hogue (2 larvae: LACM). II March 1971, C.L. Hogue and
J.P. Donahue (32 pupae and pupal skins, 24 larvae: LACM).

ETYMOLOGY: The species is named for Costa Rica, its coun-
try of origin.

Paltostoma delectata Alexander

Figures 12-14, 30-33, 40

Paltostoma delectata Alexander 1958:47. Type locality: Costa Rica, San
Jose Province, Rio Buena Vista, Rivas. Deposited in private collec-
tion of C.P. Alexander, Amherst, Massachusetts.

Paltostoma delectata, Hogue 1971:8.7.

DESCRIPTIONS

Male (Figs. 12- 14)

From the original description and the single paratopotype:

COLORATION: Head. — Front dull-orange, silvery pruinose,
mouthparts pale-brown; remainder of head chestnut-brown,
variegated with silvery on the anterior vertex. Antenna. — Not
determinable. Thorax. — Scutum chestnut-brown, including four
broad confluent prescutal stripes, leaving broad silvery areas on

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 311:1-22.

Hogue: Costa Rican Blephariceridae

the posterior pronotum and at the prolongations of the [parapsi-
dal?] “suture.” Lateral sclerites yellow. Legs. — ”... coxae
and trochanters yellow; forefemora chiefly black, more bright-
ened basally and with a very obscure yellow to scarcely evident
subterminal yellow ring; remaining femora yellow, the extreme
tips brownish black; foretibiae and tarsi black, the remaining tarsi
obscure brownish yellow. ...” Wing. — Membrane hyaline;
vein brownish black to black. Haltere. — Stem yellow, knob
blackened. Abdomen. — First tergite yellow, the remainder
brown, their extreme bases paler and more or less pruinose. Ster-
nites more extensively yellow, silvery pruinose beneath, the sides
infuscated. Genitalia. — Dark-brown.

SIZE: Measurements as follows (from paratopotype): Body
length 3.5. Wing length 4.9, width 1.9. Head width 0.80.
Clypeus-labrum length combined 1.55. Palpal segments 2-3,
lengths not available. Leg segment lengths as below:

fore

mid

hind

femur

2.9

3.8

4.9

tibia

3.5

3.7

4.4

tarsus 1

1.41

1.58

n.a.

0.57

0.63

n.a.

0.27

0.30

n.a.

0.16

n.a.

0.25

0.24

n.a.

HEAD: Exact palpal segment proportions not determinable
from material, but the palpus is very short and similar to that of
the other species with two segments (morphologically, segments
2-3 fused). The indistinct intersegmental articulation 2-3,
common to this organ in male Paltostoma, probably led to Alex-
ander’s remark in the original description that the palpus is
apparently 1 -segmented. Antennal segment proportions: 1.0, 1.1,
LOO, 1.0, 1.0, 1.1, 1.0, 1.0, 1.0, 1.0, 1.0, 1.0, 1.0, 1.0, 1.5.
Ultimate segment 1.5 length of penultimate.

LEGS: Progressive segment proportions as below:

foreleg

1.0

1.2

0.4

0.5

0.6

1.6

midleg

1.0

0.4

0.5

1.5

hindleg

1.0

0.9

n.a.

WING (Fig. 14): Venation and trichia typical for genus.

GENITALIA (Figs. 12-13): IXth tergite lobe prominent,
symmetrically triangular, apex broadly acute; arms of ventral
armature contiguous cephalad. Margin of outer dististyle entire,
obtusely rectangular. Inner dististyle simple, apex slightly spatu-
late. Tegmen elongate, narrow; apex truncate; basal portion nar-
row, articulating with the contiguous arms of ventral armature of
IXth tergite lobes. Phallosome lacking parameres, longitudinal
striae evident in ventral plate; penis filaments short, evenly
tapered, only slightly widened basad, basal fourth alveolate.

Female

Unknown.

Pupa (Not figured)

SIZE: Medium. Measurements (N = 5): all females (?), length
5.0, width 3.6.

STRUCTURE: Outline elliptical, L/W 1.4- 1.5, compressed,
cross section hemilenticular, peripheral margin acute. Cephalic
sclerite underfolded only a little more than half the distance to
level of tips of palpus cases. Anterior margin of scutellar sclerite
shallowly U-shaped. Cuticular spines proportionately long, longest
lateral spines slightly longer than middorsal length of scutellar
sclerite; only two spines in mesoposterior corner of scutum; para-
basal spinelets commonly multiple (2-4) and of unequal length.

Lamellae of respiratory processes parallel, erect, subequal in
length; outer pair heavier than inner and triangular, inner pair
about 0.5 width of outer at their bases; no. 3 the smallest and
narrower than no. 2 and set mesad of same; apices of both inner
lamellae straight; apices of lamellae no. 4 of each side conver-
gent.

Larva (Figs. 30-33)

SIZE: Medium. Measurements (N=10): Body length 6.1; head
capsule width 1.41 .

CHAETOTAXY (trunk segments only): General secondary
dorsal sensilla mostly small and coniform, grading to larger and
spiniform laterad; a triplet (short series of 3, rarely 2) of con-
spicuously larger spiniform sensilla in the anterolateral and pos-
terolateral tergal regions (latter triplet lateral to primary subtergal
sensillum, tp,). Sensilla of antero- and posterolateral linear series
large and spiniform; lateral and ventral members of both series
with apices acuminate and sometimes multiple.

STRUCTURE: Pseudopods elongate, L/W about 3.5; arising
from near apex of pleuron. Posterolateral lobe of anal division
well developed, about half the size of the pseudopods, its apex
directed posterolaterad. Lateral margin of anal division entire, set
with a few larger spiniform secondary sensilla. Posterior extreme
of anal division densely pigmented and heavily sclerotized and
separated from posterolateral lobes by a deep depression. Ventral
integument mesal to insertion of pseudopod mammillate over a
broad area, the mammillae spinose laterad, grading to papillae
mesad.

MATERIAL:

COSTA RICA. San Jose Province. 2 miles W Grecia, Rio Trojas, 18 June
1972, C.L. Hogue (7 pupae, 24 larvae: LACM). Pan American Highway and
Rio Cajon, 22 June 1972, C.L. Hogue (11 larvae: LACM). Pan American
Highway and Rio Union, 22 June 1972, C.L. Hogue (1 larva: LACM). Rivas,
January 1934. D. Rounds (Holotype 6 , 1 paratype 6 : Alexander Collection).
10 miles E San Isidro del General, Rio General, 22 June 1972, C.L. Hogue (1
larva: LACM). 1.8 miles S San Isidro del General, Rio Pedregoso, 28 June
1972, C.L. Hogue (24 larvae: LACM).

REMARKS

Unfortunately, delectata is imperfectly known. The larva and
pupa described here, although slightly large for the size of the
adult, I judge to belong to this species since they were taken in
the same drainage system (Rio General) from whence the holo-
type comes (see Fig. 40). However, there are no collections of
early stages associated with adults to confirm this.

Paltostoma diriageni new species

Figures 3-4, 15-19, 26, 34-36, 38, 40
DIAGNOSIS

Male genitalic features as follows set this species apart from its
relatives: long penis filaments, gradually widening basally; bases
of these structures non-alveolate internally. IXth tergite lobe
asymmetrically triangular (similar to that of zwicki but the two
lobes more divergent); arms of ventral armature widely separated
anteriorly. Margin of outer dististyle entire.

DESCRIPTIONS

Male (Figs. 3, 15- 19)

Based on 3 pharate specimens dissected from their pupal cases:

COLORATION: Not determinable.

SIZE: Measurements as follows: wing and leg dimensions not
determinable. Head width 1. 16. Clypeus-labrum length combined
2.88. Palpal segments 2-3 combined 0.35.

HEAD (Fig. 3): Palpus very short, segments 2-3 fused
although limit of former marked by enlarged setae; segment 1

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 311:1-22.

Hogue: Costa Rican Blephariceridae

distinct, 2-3 constricted at presumed joint; segment proportions
1:(1:0.5). Antenna! segment proportions: 1.0, 1.5, 1.00, 1.2,
1.3, 1.3, 1.3, 1.2, 1.2, 1.2, 1.3, 1.3, 1.3, 1.4, 1.6; ultimate
segment 1.2 length of penultimate.

LEGS: Segment proportions not determinable.

'WING: Venation and trichia typical for genus.

GENITALIA {Figs. 15-19): IXth tergite lobe prominent,
asymmetrically triangular, apex broadly acute; arms of ventral
armature widely disjunct anteriorly. Margin of outer dististyle
entire, roughly pentagonal. Inner dististyle simple, apex con-
stricted, a ventromesal lobe present subapically; apicoventral lobe
digitate, pronounced. Tegmen with narrow base, apex spatulate,
basal portion broad, articulating laterally with the disjunct arms
of the ventral armature of the IXth tergite lobes. Phallosome lack-
ing parameres, complex striae evident in ventral plate; penis
filaments long, evenly tapered, only slightly widened basad,
extreme basal portions non-alveolate internally.

Female (Figs. 4, 26)

Based on 4 pharate specimens dissected from their pupal cases:

COLORATION: Not determinable.

SIZE: Measurements as follows: Wing and leg dimensions not
determinable. Head width 1.19. Clypeus length 0.57. Labrum
length 1.08. Palpal segments 2-3: 0.27, 0.44.

HEAD: Palpus very short, segments 1-3 distinct, proportions
2-3: 1.0, 1.6. Antennal segment proportions: 1.2, 1.2, LOO,
1.2, 1.3, 1.2, 1.1, 1.1, 1.0, 1.0, 1.1, 1.0, 1.1, 1.3; ultimate
segment 1.2 length of penultimate.

LEGS: Segment proportions not determinable.

WING: Venation and trichia typical for genus.

GENITALIA (Fig. 26): Oviscapt elongate, ratio L/W=1.8.
Setae of VUIth stemite lobe long, 22-31 in number.

Pupa (Fig. 38)

SIZE: Large. Measurements (N= 10): male length 6.7, width
4.4; female length 7.2, width 5.0.

STRUCTURE: Outline elliptical, L/W 1.4- 1.5, compressed,
cross section hemilenticular, peripheral margin acute. Cephalic
sclerite underfolded only slightly more than half the distance to
level of tips of palpus case. Anterior margin of scutellar sclerite
shallowly V-shaped. Cuticular spines proportionately short, long-
est lateral spines distinctly less (about 0.75) than mid-dorsal
length of scutellar sclerite; numerous spines in mesoposterior
region of scutum, parabasal spinelets commonly multiple (2-4)
and of unequal length. Lamellae of respiratory processes parallel,
erect, subequal in length; outer pair heavier than inner and tri-
angular, inner pair 0.6-0. 7 width of outer at their bases; no. 3
the smallest and narrower than no. 2 and set mesad of same;
apices of both inner lamellae slightly recurved but neither bent
tortuously .

Larva (Figs. 34-36)

SIZE: Large. Measurement (N = 10): body length 6.3; head
capsule width 1.69.

CHAETOTAXY (trunk segments only): General secondary
dorsal sensilla all small and coniform; those of antero- and pos-
terolateral linear series much larger than general dorsals; cylindri-
form mesally, becoming coniform, flattened and sometimes
ridged, laterally.

STRUCTURES: Pseudopods short, L/W about 2.3; arising
from ventrolateral depression. Posterolateral lobe of anal division
well developed, only slightly smaller than pseudopods, apex
directed posterolaterad. Lateral margin of anal division with a
conspicuous lobe anterior to posterolateral lobe, set with an arc
of large, ovate sensilla. Posterior extreme of anal division

densely pigmented and heavily sclerotized and separated from
posterolateral lobes by a deep incision. Ventral integument mesal
to base of pseudopod mammillate over a broad area, mammillae
spinose laterad, grading to papillae mesad.

MATERIAL AND TYPE LOCALITY:

Types. COSTA RICA, Alajuela Province. Rio Angel below El Angel
Falls, 13 April 1975. C.L. Hogue: HOLOTYPE 8 (pharate, dissected from
and associated with pupal case; in alcohol: genitalia on slide No. CLH
75-219; head on slide No. CLH 75-233: LACM); ALLOTYPE 9 (as 8:
genitalia on slide No. CLH 75-221; head on slide No. CLH 76-158:
LACM); 3 <3, 1 9 PARATYPES (pharate adults in alcohol, dissected from
pupal cases with genitalia and heads on slides: LACM).

Additional. COSTA RICA, Alajuela Province, Rio Angel below El Angel
Falls, 13 April 1975, C.L. Hogue (19 <3,319 pupae and sexed whole pupal
skins; 15 fragmentary pupal skins; 2 larvae: LACM). Puntarenas Province,
Las Cruces, (rock quarry) 6 miles S San Vito de Jaba, Rio Jaba, 27 June
1972, C.L. Hogue (1 pupa. I larva: LACM). 1.8 mi. W Rincon, Linguita
Creek, 3 July 1963, C.L. Hogue (1 9 , pharate, dissected from pupal case; 1
pupa, 3 partial pupal skins, 10 larvae: LACM). 11 March 1971, C.L. Hogue
and J.P. Donahue (2 pupae, 2 larvae: LACM). San Jose Province, 2 miles
W Grecia, Rio Trojas, 18 June 1972, C.L. Hogue (1 pupa: LACM). 12.5 miles
N San Isidro del General, 15 June 1963, C.L. Hogue (7 larvae: LACM).

ETYMOLOGY: The name was chosen to commemorate the
name of one of Costa Rica’s most famous and clever caciques. In
1522, Diriagen is reputed to have nearly defeated a force led by
Gonzalez Davila in an ambush into which the latter’s greed for
Indian treasure led him.

Paltostoma exserta new species

Figures 10-11, 39, 40

DIAGNOSIS

This species is distinct from all other known Paltostoma in the
form of the pupal lamellae. Normally the four plates of the bran-
chia are more or less separate from one another and about equal
in length (although the inner two may be narrower than the
outer), but in exserta the inner pairs are appressed by the outer
and extend about 20% of their own length beyond them.

Only small differences exist in other features, notably in the
male genitalia, between exserta and the very similar diriageni.
This applies principally to the shape of the inner dististyle which
is considerably broader in the former than in the latter species.
This structure also has a less pronounced apicoventral lobe in
exserta than in diriageni. A further difference may be found in
the outer dististyle lobe which is smoothly rounded in shape in
exserta while it is asymmetrically hexagonal in diriageni.

The larva of exserta is unknown.

DESCRIPTIONS

Adults

The great similarity of this species to diriageni (see above)
makes a full description of the adults superfluous. From the small
amount of material available, which consists entirely of pharate
individuals dissected from their pupal cases, nothing can be
detected to distinguish the females. The same is true of the males
except for a few characteristics in the genitalia. The latter are
described as follows from the single fairly well sclerotized, but
pharate, specimen:

MALE GENITALIA (Figs. 10-11): IXth tergite lobe prom-
inent, asymmetrically triangular, apex broadly acute; arms of
ventral armature widely disjunct anteriorly. Margin of outer disti-
style entire, oval; dorsal portion concave. Inner dististyle simple,
broad and angular, a ventromesal lobe present subapically;
apicoventral lobe acute and moderately pronounced. Tegmen
with narrow base, apex spatulate; basal portion broad, articulat-

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 311:1-22.

Hogue: Costa Rican Blephariceridae

ing laterally with the disjunct arms of the ventral armature of the
IX th tergite lobes. Phallosome lacking parameres, complex striae
evident in ventral plate; penis filaments long, evenly tapered,
only slightly widened basad; extreme basal portions of filaments
non-alveolate internally.

Pupa (Fig. 39)

SIZE: Large. Measurements: male (N=3) length 5.9, width
4.4; female (N = 7) length 7.0, width 5.0.

STRUCTURE: Outline elliptical, L/W 1.4. Cuticular spines
proportionately short, longest lateral spines distinctly less (about
0.75) than mid-dorsal length of scutellar sclerite; condition of
parabasal spinelets not determinable due to abraded condition of
available material. Branchial lamellae compressed, outer pair
much heavier than inner and triangular, appressed to inner pair
which are sagittate in form and extend considerably beyond the
apices of the outer pair (by about 20% of their total length); outer
lamellae covered by a dense, white, flocculent material of
unknown origin.

Larva

Unknown.

MATERIAL AND TYPE LOCALITY:

Types. COSTA RICA, Alajuela Province. Rio Angel below El Angel
Falls, 13 April 1975, C.L. Hogue: HOLOTYPE 3 (pharate, dissected from
and associated with pupal case; in alcohol: genitalia on slide No. CLH 77-1;
head on slide No. CLH 77-10: LACM); ALLOTYPE 9 (as 3: genitalia on
slide No. CLH 77-3: LACM); 3 9 PARATYPES (pharate adults in alcohol,
dissected from pupal cases with genitalia and heads on slides: LACM).

Additional. COSTA RICA, Alajuela Province, Rio Angel below El Angel
Falls, 13 April 1975. C.L. Hogue (2 3 , 8 9 pupae: LACM).

ETYMOLOGY: Exserta is Latin, meaning “extended be-
yond” and is in reference to the protruding middle branchial
lamellae of the pupa.

Paltostoma zwicki new species

Figures 20-24, 40

DIAGNOSIS

The following characteristics of the male genitalia distinguish
this new species from its congeners: short penis filaments with
basal third abruptly widened and alveolate internally. IX th tergite
lobe asymmetrically triangular with an acute apex, arms of ven-
tral armature contiguous anteriorly. Outer dististyle incised
apicoventrally.

DESCRIPTIONS

Male (Figs. 20-24)

Based on the holotype and a single male paratopotype:

COLORATION: Head. — All sclerites medium-brown, darker
towards vertex and caudad, with extensive velvety texture and
silvery opalescent reflections; ocellar triangle dark-brown.
Mouthparts and antenna dull brown, the latter darker. Thorax. —
Generally medium-brown. Scutum, scutellum, epistemum and
meron darker and with reflections as on head; remaining sclerites
dull yellow-brown. Legs. — Generally dull gray-brown, lighter
basad, becoming darker distad; tarsi, including claws, almost
black. Apices of femora darkened. Wing. — Membrane hyaline,
veins brownish-black. Halt ere. — Stem yellow basad, darkening
distad; knob velvety brown. Abdomen. — Basal half of first
tergite yellow, the remainder of this and other tergites velvety-
brown with silvery reflections caudad, a distinct reflective black
band basally. Sternites light-brown, also reflective. Genitalia. —
Dark brown.

SIZE: Measurements as follows (from combined material.

maximum N=2): Body length not determinable. Wing length
5.0, width 1.7. Head width 0.82. Clypeus length 0.23. Labrum
length 1.66. Palpal segments 1-3 combined 0.21, 2-3 combined
0.15, 2 alone 0.06. Leg segment lengths as below (from Pedre-
gosa specimen):

fore

mid

hind

femur

3.3

4.2

5.0

tibia

3.9

4.1

4.8

tarsus 1

1.38

1.43

1.56

0.59

0.68

0.46

0.34

0.38

0.24

0.23

0.24

0.20

0.28

0.29

0.26

HEAD: Palpus very short, segments 2-3

fused, segment 1 dis-

tinct, 2-3 constricted at presumed joint; segment 2 short; propor-
tions: 1 :( 1 : 1 .4) . Antennal segment proportions (from Pedregosa
specimen, segments 9-14 damaged): 1.7; 1.7, 1.00, 1.3, 1.3,
1.4, 1.3, 1.3, 1.3, 1.4, 1.1, 1.4, 1.3, 1.3, 1.9; ultimate segment
1.5 length of penultimate.

LEGS: Progressive segment proportions as below:

foreleg

1.0

1.2

0.4

0.6

0.7

1.2

midleg

1.0

0.4

0.5

0.6

1.2

hindleg

1.0

0.3

0.5

0.8

1.3

WING: Venation and trichia typical for genus.

GENITALIA (Figs. 20-24): IXth tergite lobe prominent,
asymmetrically triangular, apex broadly acute; arms of ventral
armature contiguous anteriorly. Outer dististyle incised apicoven-
trally, ventromedian margin not markedly produced. Inner disti-
style slightly expanded apicad, apex abruptly narrowed and
attenuate. Tegmen broad, apex rounded with a median, narrow
sclerotized shaft; basal portion narrow, articulating with the con-
tiguous arms of the ventral armature of IXth tergite lobes. Phal-
losome with slightly sclerotized straps in center of ventral plate,
suggesting rudimentary parameres; penis filaments short, basal
third abruptly widened, this portion alveolate internally.

Female, pupa and larva

Unknown.

MATERIAL AND TYPE LOCALITY:

Types. COSTA RICA, San Jose Province, Ttirrialba, 5-15 March 1954,
C.H. Batchelder: HOLOTYPE 3 (genitalia on slide No. CLH 76-161;
USNM); PARATYPE 3 (genitalia on slide No. CLH 76-162: USNM).
March 1954. C.H. Batchelder (light trap).

Additional. COSTA RICA, Province Unknown, Pedregosa, no date, D.L.
Rounds (1 3: USNM).

ETYMOLOGY: The species is named in honor of Peter
Zwick, distinguished student of the family Blephariceridae.
REMARKS

I have not been able to locate the locality “Pedregosa” of
Rounds’ collection; it may be Pedregoso, 1 km W San Isidro del
General.

ACKNOWLEDGMENTS

I owe gratitude to the following persons for making available
the material upon which this report is based, either by providing
funds and special help in the field, or by loaning specimens:
Charles P. Alexander, John E.S. Dockweiler, Julian P. Donahue,
Alan Landsburg, and Peter Zwick.

I would like to acknowledge also the invaluable assistance of
Betty Birdsall and Carol Madle with preliminary drawings and to

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 311:1-22.

Hogue: Costa Rican Blephariceridae

thank particularly Joy Roan for her fine rendering of figures of
the larvae (Figs. 27-36).

RESUMEN

Se hace referencia a cinco especies de mosquitos de alas reticu-
ladas de la familia Blephariceridae, procedentes de Costa Rica.
Las cinco especies pertenecen al genero Paltostoma; con excep-
cion de P. delectaia Alexander, ya descrita, las cuatro restantes
(costaricensis, diriageni, exserta y zwicki) se describen como
nuevas especies. La publicacion incluye ademas un diagnostico,
acompahado de notas biologicas, referentes al genero.

LITERATURE CITED

Hogue, C.L. 1971. Family Blephariceridae, number 8, pages
8.1-8.12 in Papavero, N. ed. A catalogue of the Diptera of
the Americas south of the United States. Museu de Zool-
ogia, Universidade de Sao Paulo, Sao Paulo.

1973. The net-winged midges or Blephariceridae of

California. California Insect Survey, B. 15:1-83.

. 1978. The net- winged midges of eastern North Amer-
ica, with notes on new taxonomic characters in the family
Blephariceridae. Natural History Museum of Los Angeles
County, Contrib. Sci. 291:1-41.

Stuckenberg, B.R. 1958. Taxonomic and morphological studies
on the genus Paulianina Alexander (Diptera: Blephariceri-
dae). Institut Scientifique de Madagascar, Memoires, Serie
E 10:97-198.

Alexander, C.P. 1953. Undescribed species of nematocerous

Diptera. Part II. Brooklyn Entomol. Soc., B. 48:21-49. Accepted for publication April 10, 1978.

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 311:1-22.

Hogue: Costa Rican Blephariceridae

heads in frontal aspect. 1, male. 2, female.

Ccmtrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 311:1-22

Hogue: Costa Rican Blephariceridae

Figures 3-4. Paltostoma diriageni, heads in frontal aspect. 3, male. 4, female.

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 311:1-22.

Hogue: Costa Rican Blephariceridae

Figures 5-9. Paltostoma costaricensis, male genitalia. 5, basistyle, dististyle and associated structures, dorsal aspect. 6, tegmen and ventral bridge. 7,
right dististyle, inner aspect. 8, IXth tergite lobes and ventral armature. 9, phallosome and ventral plate.

Conrrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 311:1-22.

Hogue: Costa Rican Blephariceridae

Figures 10-14. Paltostoma, male genitalia and wing. 10—11, P. exserta. 10, right dististyle, inner aspect. 11, phallosome and ventral plate. 12-14, P .
delectata. 12, right dististyle, inner aspect. 13, IXth tergite lobes and ventral armature. 14, wing.

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 311:1-22.

Hogue: Costa Rican Blephariceridae

Figures 15- 19. Paltosroma diriageni, male genitalia. 15, basistyle, dististyle and associated structures, dorsal aspect. 16, tegmen and ventral bridge. 17,
right dististyle, inner aspect. 18, IXth tergite lobes and ventral armature. 19, phallosome and ventral plate.

Contrib. Sci. Nalur. Hist. Mus. Los Angeles County. 1979. 311:1-22.

Hogue: Costa Rican Blephariceridae

Figures 20-24. Paltostoma zwicki, male genitalia. 20, basistyle, dististy le and associated structures, dorsal aspect. 21, tegmen and ventral bridge. 22,
right dististyle, inner aspect. 23, IXth tergite lobes and ventral armature. 24, phallosome and ventral plate.

Contrib. Sci. Natur. Hist. Mus. Los Angeles Count}’. 1979. 311:1-22.

Hogue: Costa Rican Blephariceridae

Figures 25-26. Paltostoma, female genitalia, ventral aspect. 25, P. costaricensis. 26, P diriageni.

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 311:1-22.

Hogue: Costa Rican Blephariceridae

Figures 27-29. Paltostoma costaricensis, larva. 27, anal division, outline. 28, trunk division II, dorsal aspect. 29, same, ventral aspect.

Contrib. Set. Natur. Hist. Mus. Los Angeles County. 1979. 311:1-22.

Figure 30. Paltostoma delectata, larva, dorsal aspect, left; ventral aspect, right.

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 311:1-22.

Hogue: Costa Rican Blephariceridae

Figures 31-33. Paltostoma delectata, larva. 31, anal division, outline. 32, trunk division II, dorsal aspect. 33, same, ventral aspect.

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 311:1-22.

Hogue: Costa Rican Blephariceridae

Figures 34-36. Paltostoma diriageni, larva. 34, anal division, outline. 35, trunk division 0, dorsal aspect. 36, same, ventral aspect.

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 311:1-22.

Hogue: Costa Rican Blephariceridae

Figures 37-39. Paltostoma, pupae, anterior aspect. 37, P costaricensis. 38, P. diriageni. 39, P. exserta.

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 311:1-22.

Hogue: Costa Rican Blephariceridae

Figure 40. Map of Costa Rica with collection localities of Paltostoma shown by symbols. Heavy lines outline major drainage systems, heaviest indicating
continental divide.

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 311:1-22.

PROMYLAGAULUS, PROGRESSIVE A PLODONTOI D RODENTS

OF THE EARLY MIOCENE

lil

By John M. Rensberger

»5

fPlT

3""' 'I pilli 111 ill'illiiliii tti

Published by the NATURAL. HISTORY MUSEUM OF LOS ANGELES COUNTY - 300 EXPOSITION BOULEVARD * LOS ANGELES, CALIFORNIA $000?

Prior to November 30, 1 973, publications of the Natural History Museum have appeared under various titles — Leaflet
Series, Museum Graphic Education Series, History Series, Science Series, Study Guides, Contributions in History \ History
Bulletin, Science Bulletin, unnumbered catalogs of exhibitions, and other miscellaneous publications.

The publication program of the Natural History' Museum continues to publish a series of reports that give accounts of
new discoveries in history and science as well as new advances in museum knowledge. The theme of the Natural History
Museum serials will emphasize basic research through the continuing active series:

Contributions in Science

Science Series

the world..

Acceptance of manuscripts will be determined by the significance of new information. Priority will be given to
manuscripts by staff members. All manuscripts must be recommended by the curator in charge of each discipline or by the
Editorial Board, Peer judgment will be accomplished through the various divisions of the Museum and by outside

reviewers.

Exposition Boulevard, Los Angeles, California 90007.

Edward Ostermeyer
Editor

Number 312
March 16, 1979

CONTRIBUTIONS IN SCIENCE

NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY

PROMYLAGAULUS, PROGRESSIVE APLODONTOID RODENTS

OF THE EARLY MIOCENE

By John M. Rensberger

Published by the NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY • 900 EXPOSITION BOULEVARD • LOS ANGELES, CALIFORNIA 90007

CONTENTS

Abstract 3

Abbreviations of Institutions 3

Dental Nomenclature 3

Methods 4

Morphologic Relationships 4

Systematic Paleontology: Order RODENTIA,

Superfamily Aplodontoidea Matthew 1910, Family Mylagaulidae Cope 1881

Promylagaulus McGrew 1941 13

Promylagaulus riggsi McGrew 1941 14

Promylagaulus lemhiensis Nichols 1976 14

Promylagaulus ovatus new species 15

Promylagaulus montanensis new species 15

Promylagaulus sp 16

Phyletic Relationships of the Species 16

Acknowledgments 17

Literature Cited 17

ISSN: 0459-8113

Suggested Citation: Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 312:1-18.

Contributions in Science are articles in the earth and life sciences, presenting results of original research in Natural
History. Science Bulletin (Numbers 1-30; 28 June 1965 to 10 April 1978) and Contributions in Science (Numbers 1-300;
23 January 1957 to 16 June 1978) were merged into a single imperial octavo serial beginning with Number 301.

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 312:1-18.

PROMYLAGAULUS, PROGRESSIVE APLODONTOID RODENTS

OF THE EARLY MIOCENE1

By John M. Rensberger2

ABSTRACT: Promylagaulus, until recently represented by only a single described species of uncertain age, P .
riggsi, is a diverse and rather widely distributed genus, most members of which seem to occur in a restricted
interval of the early Miocene. Of the two new species described here, P. ovatus is from an upper part of the
Monroe Creek Formation in South Dakota, and P . montanensis occurs in a restricted interval in the Deep River
Formation of Montana. A form from northern Wyoming, when better known, may represent another species.
Each of these three occurrences is within the Entoptychus-Gregormymys Concurrent-range Zone of Fisher and
Rensberger (1972). P. lemhiensis, from northern Idaho, is apparently from an only slightly older interval.

These species differ in hypsodonty, size and lophodonty. No trend toward characteristics definitive of the
Mylagaulidae is evident among these species and a number of features indicate that Promylagaulus was not
ancestral to the later mylagaulids. Although similar in a number of characteristics to an earlier and more
primitive aplodontoid, Meniscomys. Promylagaulus frequently retained at least one primitive feature that was
absent in the earliest known members of Meniscomys.

A few years ago, a biostratigraphic study of the Deep River
and Fort Logan formations of Montana was undertaken in order
to more clearly correlate Oligo-Miocene sequences of vertebrates
of the far west with faunas within and east of the Rocky Moun-
tain region. One of the taxa recovered in the course of this study
is an aplodontoid rodent resembling specimens referred to Promy-
lagaulus riggsi McGrew, described from the Monroe Creek
Formation of South Dakota by J.R. Macdonald (1963, 1970) and
L. Macdonald (1972), and P. lemhiensis Nichols (1976) from the
early Miocene of Idaho.

The problem of assessing the relationships of small samples of
Promylagaulus is difficult because the cheek teeth in this genus
are high crowned and complex, with numerous enamel bordered
lakes of different depths and which vary in size and shape with
wear. The dentition in the type specimen of P. riggsi (McGrew
1941) has few characteristic features that are not made ambigu-
ous by the advanced stage of wear. The more recently described
specimens of Promylagaulus were reported upon in papers
devoted to entire faunas and the comparative morphologies have
not as yet been presented in detail. This paper compares the new
material of Promylagaulus from the Deep River Formation with
that from South Dakota and Idaho, and presents additional evi-
dence concerning the relationship of these forms.

Abbreviations of Institutions

AMNH American Museum of Natural History
FMNH Field Museum of Natural History
ISU The Museum, Idaho State University
LACM Los Angeles County Museum of Natural History
SDSM South Dakota School of Mines and Technology
UM Department of Geology, University of Montana
UWBM University of Washington, Burke Memorial Washing-
ton State Museum

Dental Nomenclature

The principal cusps of the cheek teeth in Promylagaulus (Fig.
1) correspond to those of Haplomys (Rensberger 1975: Fig. 1).
Whereas the major cusps are homologous to those of other groups
of rodents, the enamel inflections, fossettes and fossettids are
frequently not homologous. Different authors have applied the
terminology used by Stirton (1935) for the castorids to the myla-
gaulids, but in conflicting ways. Because of fundamental differ-
ences in the origins of the inflections and lakes in the castorids
and aplodontoids, there is no obvious and natural way to transfer
a number of the names from the former to the latter. Rather than
introduce new terms for these structures in the aplodontoids at
this time, and to avoid confusion with the several usages of the
castorid nomenclature, I have simply used descriptive terms
based upon the position of the structure with reference to the
center of the tooth (Fig. 1). Thus the structures in any of the four
quadrants are prefixed by anterolingual, posterolingual, etc., and
those falling in the midsection of a side by labial, lingual, ante-
rior or posterior. An advantage of this method is that when
applied to fossettes and fossettids, it lends itself to situations in
which a structure originates in different ways. For example, with-
in a single species of Promylagaulus there may be both antero-
lingual and posterolingual fossettids on P4, or there may be only
a lingual fossettid in those positions, depending upon whether or
not the mesostylid is joined to the mesoconid by a crest. This

'Review Committee for this Contribution:

Craig Black
Mary Dawson
David Whistler

2Burke Memorial Washington State Museum and Department
of Geological Sciences, University of Washington, Seattle 98195.

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 312:1-16.

Rensberger: Early Miocene Promylagaulus

PAS PA MSS ME

Figure 1. Nomenclature of occlusal structures in Promylagaulus. u-left P4; b- left M1 or M2; c — little worn right P4; d-well worn right P4; e-right DP4;
/-right Mi or M;. Cross hatching = dentine; labial side toward top, anterior toward left. ABF = anterolabial fossette (-id); ABI = anterolabial inflection;
AC = anterocone; AF = anterior fossettid; AI = anterior inflection; ALF = anterolingual fossette (-id); ALI = anterolingual inflection; BI = labial
inflection; CF = central fossette (-id); END = entoconid; HLD = hypolophid; HYLD = hypoconulid; HYD = hypoconid; LF = lingual fossettid; LI =
lingual inflection; MCD = mesoconid; ME = metacone; MED = metaconid; MEL = metaconule; ML = metaloph; MLDII = metalophulid II; MSD =
mesostylid; MS DC = mesostylid crest; MSS = mesostyle; MTDC = metastylid crest; PA = paracone; PAS = parastyle; PBF = posterolabial fossette;
PBI = posterolabial inflection; PF = posterior fossettid; PL = protoloph; PLD = posterolophid; PLF = posterolingual fossette; PLI = posterolingual
inflection; PR = protocone; PRD = protoconid; PRL = protoconule (paraconule) .

method somewhat simplifies the terminology in a group in which
the occlusal structure can be very complex.

METHODS

The most objective information bearing on the relationship of
the fossil jaws, teeth and skull representing Promylagaulus is the
stratigraphic- geographic occurrence of the elements. Specimens
from a single geographic and stratigraphic position and among
which there is no evidence of a significant discontinuity in mor-
phology were probably members of a single population or very
closely related ones. This information, together with the complex
combinations of morphologic similarities and dissimilarities pre-
sented in the following sections show that at least four distinct
taxonomic groups are represented.

The distinct geographic-stratigraphic associations are:

Deep River Fm., central Montana, (P . montanensis)

Lemhi Valley, western Montana, (P. lemhiensis)

Anthills locality, uppermost Monroe Creek Fm., S. Dakota,
(P . ovatus)

Specimen SDSM 6277 , Monroe Creek Fm., S. Dakota, (P.
ovatus )

P. riggsi, unknown locality, S. Dakota
Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 312:1-18.

MORPHOLOGIC RELATIONSHIPS

The material from the Deep River Formation consists of two
right mandibles: UWBM 38075, with P4-M3; and UWBM
38103, with a lower incisor fragment and DP4-M2. The speci-
mens are from approximately the same stratigraphic position in a
5 to 7 m thick massive sandstone at a single exposure. Although
the relatively little worn and moderately worn occlusal patterns of
the molars in the two specimens differ greatly, analogous differ-
ences are also present among the isolated teeth reported from an
anthill locality in the Monroe Creek Formation of South Dakota
and referred to Promylagaulus riggsi by L. Macdonald (1972).
The specimens from the Deep River Formation probably repre-
sent a single population, or at least closely related ones.

Lower Incisor

The enamel face of the incisor in the Deep River taxon is
broad and only slightly convex transversely. It closely resembles
that of Meniscomys hippodus Cope, of the John Day Formation,
except that the width is somewhat greater, consistent with the
overall larger size of the mandible. The incisor is wider and
slightly flatter than that of SDSM 6277, cf. Promylagaulus riggsi

Rensberger: Early Miocene Promylagaulus

(Macdonald 1970:33) from South Dakota, a specimen of smaller
overall size.

Deciduous Lower Premolar

The DP 4 of the Deep River form (UWBM 38103) is well worn
but still retains several fossettids (Fig. 2a). A central anterior
fossettid is triangular with one angle in a posterior position and
the remaining angles in anterolabial and anterolingual positions.
The anterolingual corner of the fossettid is not quite closed at the
existing stage of wear. The anterior fossettid in SDSM 6277 also
opens anterolingually, although the fossettid is more worn and
less triangular than in the specimen from Montana. In a specimen
of Meniscomys hippodus (UWBM 29160), the anterior fossettid
is oval, narrower transversely than that in SDSM 6277 or the
Deep River form, and opens directly craniad. Three unworn to
lightly worn deciduous lower premolars, 117384, 117436 from
the anthill locality (LACM 1871 = SDSM 6229) were not
described by L. Macdonald (1972), although they are question-
ably assigned on the specimen record to Promylagaulus riggsi.
Although pronounced morphologic differences exist among these
specimens, they probably belong to the taxon represented at that
locality by permanent teeth, as will be shown below. The anterior
fossettid or inflection in the specimens of DPa from the anthill
collections (Fig. 3b) is triangular, as in the Deep River form.

Figure 2. Promylagaulus montanensis. a-occlusal view right DP4-M2,
UWBM 38103; £>-occlusal view right P4-M3, UWBM 38075. Both
specimens from level 1 of locality Spring Creek 1 (UWBM A5867),
Deep River Formation, Montana. Labial side toward top of figure. Scale
= 1 mm. Drawing by Mark Orsen.

Figure 3. Promylagaulus ovatus, lower cheek teeth, a-occlusal view little worn right P4, LACM 117372; £>-occlusal view left DP4, LACM 117436;
c-occlusal view. worn left P4, LACM 117368; <7-occlusal view worn right P4, LACM 117370; e-occlusal view left Mi, LACM 1 17435; /-same,
posterior view. All except/ stereo pairs. Labial side toward top of figure. Scale = 1 mm.

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 312:1-18.

Rensberger: Early Miocene Promylagaulus

TABLE 1

Length, Width of P-i; Depth of Labial Enamel on P4 (mm)

DLEP

MUSEUM NUMBER

LOCALITY

WTR

WTA

DLEP

UWBM 38075

Deep River Fm (UWA 5867)

4.1

2.2

2.7

2.8

0.68

ISU 18594

Lemhi Valley (ISU 59003)

4.7

1.9

2.7

2.0

0.43

ISU 18601

Lemhi Valley (ISU 59003)

5.0

2.2

3.0

2.2

0.44

AMNH 56333

Darton's Bluff*

3.6

1.8

2.2

—

LACM 9340

Wolff Ranch (LACM 20051)

3.8

2.0

2.4

—

LACM 9342

Wolff Ranch (LACM 20051)

3.5

1.8

2.2

1.9

0.54

LACM 23530

Anthill (LACM 1871)

3.4

1.5

1.9

—

LACM 1 17367

Anthill (LACM 1871)

3.7

1.9

2.0

1.8

0.49

LACM 23532

Anthill (LACM 1871)

3.5

1.7

1.9

1.8

0.51

LACM 117437

Anthill (LACM 1871)

3.5

1.6

2.2

1.8

0.51

WTR

WTA

DLEP

anteroposterior length near base of crown,

transverse width of trigonid measured perpendicular to vertical axis of tooth,
transverse width of talonid, measured perpendicular to vertical axis of tooth,
depth of labial enamel measured ventrad from apex of posterior dentinal tract
measurements taken from published illustration (McKenna and Love 1972).

although the sequence of closure of the two anterior comers is
variable.

An arcuate anterolabial inflection in UWBM 38103 lies in a
transverse orientation near the anteroposterior center of DP4 and
anterior to the mesoconid. Opposite the inflection and near the
lingual margin of the tooth is an anteroposteriorly aligned fosset-
tid that is represented in Meniscomys hippodus by a shallow val-
ley which doesn't form a fossettid. In both UWBM 38103 and
M. hippodus the fossettid or valley is bounded lingually by an
elongate metastylid crest at the margin of the tooth. Both the
arcuate anterolabial fossettid and the elongate anterolingual fos-
settid are present in the teeth of the anthill locality (Fig. 3b).
However, in one specimen of the latter the mesoconid lacks the
labial spur that isolates the anterolabial fossettid in the other
specimens.

The base of the posterolingual inflection is barely preserved in
the specimen from the Deep River Formation. It lies near the

1.5 2.0 2.5 3.0

DLEP

ANTHILL
WOLFF RH.
DEEP R.
LEMHI

Figure 4. Bivariate plot of depth of labial enamel below highest point of
posterior dentinal tract (DLEP) and anteroposterior length (LP) of P4 in
specimens of Promylagaulus from Anthill (LACM 1871), Wolff Ranch
(LACM 2005), Deep River (Spring Creek 1, level 1; UWBM A5867) and
Lemhi Valley (UM V-7303 = ISU 59003) localities.

lingual margin of the tooth between the areas of the entoconid
and the mesostylid. The margin of the tooth is extended as a small,
lingual protuberance slightly anterior to the opening of the inflec-
tion. These structures are similar to conditions present in the
specimens of both Meniscomys hippodus and the anthill collec-
tion. The teeth in the anthill collection still retain the posterolin-
gual fossettid, which is more posteriorly situated with respect to
the posterolabial fossettid than is that in M. hippodus. The hypo-
conid of the specimens in the anthill collection is more anteriorly
situated and less expansive posterolabially than in M. hippodus
(Fig. 3b). It is this feature which relates the deciduous premolars
most clearly to the molars in the anthill collection.

Lower Premolar

The worn permanent premolar from the Deep River Formation
is larger than those from the anthill collection, but its size, espe-
cially length, is exceeded by those from Lemhi Valley (Table 1).
The anterior fossettid in the worn P4 of UWBM 38075 from the
Deep River Formation (Fig. 2b) is anteroposteriorly elongate and
narrow, but slightly expanded anteriorly. In the Lemhi collection
the fossettid is also elongate, but in SDSM 6277 and the numer-
ous isolated teeth of the anthill collection (Fig. 3c, d) the fosset-
tid is wider and shorter than that in the Deep River form.

The labial inflection in UWBM 38075 is directed posterolin-
guad, toward the center of the tooth (Fig. 2b). It ends internally
at the flat surface of the mesoconid and lacks an anterior branch,
but a narrow posterior portion representing the inflection poste-
rior to the mesoconid in Haplomys (Rensberger 1975:6), the pos-
terolabial inflection, extends to within 0.1 mm of the posterior
fossettid. The hypoconid is posterolabially compressed and ante-
riorly expanded so as to contribute to the posterolingual trend of
the labial inflection. In the Lemhi collection, SDSM 6277 and
the unworn and little worn teeth in the anthill collection (Fig.
3a- d), the hypocone is similarly compressed and anteriorly
expanded, the labial inflection abuts a normally flat mesoconid
with little if any tendency to extend anteriad, and a narrow
branch of the labial inflection extends behind the mesoconid
toward the posterior fossettid, as in the Deep River form. There

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 312:1-18.

Rensberger: Early Miocene Promylagaulus

TABLE 2

Ratios of Size of Posterior and Lingual Fossettids to Width of P4

WPF4 WLIF4 WC

MUSEUM NUMBER

LOCALITY

WPF4

WL1F4

ISU 18601

Lemhi Valley (ISU 59003)

2.7

1.1

0.41

0.8

0.30

1.8

0.67

LACM 1 17370

Anthill collection
(LACM 1871)

2.2

0.7

0.32

0.6

0.27

1.5

0.68

LACM 117352

Anthill collection
(LACM 1871)

2.2

0.8

0.36

0.6

0.27

1.4

0.64

LACM 117366

Anthill collection
(LACM 1871)

2.2

0.7

0.32

0.6

0.27

1.3

0.59

UWBM 38075

Deep River Fm (UWA 5867)

2.7

0.8

0.30

0.6

0.22

1.5

0.55

WPF4

WLIF4

transverse width of P4 across talonid.

transverse width of posterior fossettid on P4, measured from outer margin of enamel,
transverse width of lingual fossettid on P4, measured from outer margin of enamel,
transverse width of P4 in center, from posterior end of posterolabial inflection.

is usually no mesoconid process on the hypoconid of P. lemhi-
ensis or most of the specimens from the Monroe Creek Forma-
tion; however in several specimens from the anthill site a crest
from the mesoconid joins the latter to the anterior process of the
hypoconid to form a small posterolabial fossettid.

In Meniscomys hippodus, in contrast to many of the specimens
from Montana, Idaho and South Dakota, the labial process of the
mesoconid is persistent and joins at depth with a process from the
hypoconid to enclose the posterolabial inflection as a fossettid.
This tendency is present in the stratigraphically lowest of the
known specimens of Meniscomys. As a consequence of this con-
dition, the labial inflection in worn teeth of M. hippodus is con-
fined to a transversely aligned slot that internally represents only
the anterolabial inflection.

The posterior fossettid is still present in the moderately
advanced stage of wear of UWBM 38075 (Fig. 2b). The lake is
oval, with the long axis trending anterolabiad. The posterior fos-
settid in the teeth in the anthill collection is larger, relative to the
width of the tooth (Table 2), than in the Deep River form (Fig.
3d). The fossettid is largest relative to tooth width in the Lemhi
collection. In SDSM 6277 and P. lemhiensis, the fossettid is
bounded in an early stage of wear by a distinct cusp joining the
entoconid and mesoconid and forming the hypolophid, but in the
anthill collection this cusp is much reduced.

A posteriorly elongate but slightly triangular lingual fossettid is
present in the P4 of the UWBM 38075. This fossettid is trans-
versely wider relative to tooth width and more oval in the Lemhi
and anthill collections (Table 2). The lake is bounded lingually
by the surface of the metastylid crest, anterolabially by the sur-
face of the mesoconid and posterolabially by the hypolophid
(Fig. 2b). In all forms the posterolingual inflection extends as a
shallow groove well below the stage of wear at which the fosset-
tid closes. A considerable amount of variation in the structures
bounding the fossettid is present among the little worn anthill
specimens and SDSM 6277. Usually a strong metastylid crest
(Fig. 3a) extends caudad to join either the hypolophid or a meso-
stylid cusp (or crest). There may be no mesostylid. When a
mesostylid is present, it may be joined to both the metastylid
crest and the mesoconid, so that the fossettid is restricted to an
anterolingual position, at least in the upper part of the crown. In
individuals in which the metastylid crest is undeveloped (Fig.
3c), the mesostylid tends to be flanked by anterolingual and pos-

terolingual inflections which may not form fossettids at all, or
only a small one. Specimens in the anthill collection in which the
metastylid crest dominates and connects to the hypolophid (Fig. 3a,
d) resemble those of the Lemhi collection and probably exhibit the
structure that would be shown by UWBM 38075 in an earlier stage of
wear.

In the primitive species of Meniscomys from the John Day
Formation (Rensberger, in manuscript) individuals both with and
without a metastylid crest are present. Those with the metastylid
crest may lack a mesostylid, or if the latter is present, have an
incipient lingual fossettid communicating (in the absence of a
union between mesostylid and mesoconid) with the posterolin-
gual fossettid. Individuals of later populations in the John Day
Formation consistently have a mesostylid bounded by anterolin-
gual and posterolingual inflections which do not close with wear
to form fossettids.

The hypsodonty of the premolar in the Deep River form, as
indicated by the depth of the labial enamel plate beneath the top
of the posterior dentinal tract, exceeds that in the other collec-
tions (Fig. 4 and Table 1). Although the absolute length of the
plate in the Lemhi collection slightly exceeds measurements in
the anthill collection, the Lemhi specimens are the more brachy-
odont, as is shown by Figure 4 and the ratio of enamel length to
tooth length (DLEP/LP in Table 1).

A locality (LACM 2005) northeast of Wolff Ranch in the
Wounded Knee region of South Dakota yielded three lower cheek
teeth, two of which are premolars which are not distinguishable
in occlusal pattern or hypsodonty (Table 1 ) from the teeth in the
anthill collection. The museum records indicate the specimens
were derived from the Sharps Formation, but the color of the
enamel is dark brown, like that of most of the teeth from the
anthill collection in the Monroe Creek Formation.

A pair of mandibular rami with heavily worn teeth having a
pattern similar to that of UWBM 38075 were found at Darton’s
Bluff, Big Horn Mountains, northern central Wyoming (McKen-
na and Love 1972:23). The wear is such that only two enamel
lakes, the anterior and lingual fossettids, remain on the P4
(McKenna and Love 1972: Figs. 15, 16), which makes difficult
determination of the relationships of the form. The anteroposte-
rior and transverse dimensions are smaller than those of UWBM
38075 and the Lemhi specimens but compare favorably with
those from the anthill collection (Table 1).

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 312:1-18.

Rensberger: Early Miocene Promylagaulus

TABLE 3

Length and Width of Mi, M2, M3 (mm)

MUSEUM

NUMBER

LOCALITY

uLl

uL2

uL3

uWl

uW2

uW3

wLl

wL2

wL3

ISU 18594

Lemhi Valley (ISU 59003)

2.8

2.6

2.0

2.1

1.6

UM 5193

Lemhi Valley (UMV 7303)

—

2.4

1.9

UWBM 38103

Deep River Fm. (UWA 5867)

2.7

2.3

—

2.0

—

UWBM 38075

Deep River Fm. (UWA 5867)

—

1.9

2.2

1.8

AMNH 56333

Darton’s Bluff (AMNH)
left teetht

2.0

2.3

1.7

SDSM 6277

Monroe Creek Fm.
(SDSM V-6210)

2.4

2.2

1.8

1.7

LACM 23534

Anthill (LACM 1871)

2.8*

—

2.1

—

2.3

—

LACM 117441

Anthill (LACM 1871)

2.4

—

1.9

—

1.7

—

LACM 117435

Anthill (LACM 1871)

—

2.2

—

1.8

1.7

—

LACM 117374

Anthill (LACM 1871)

—

2.4

—

2.1

—

2.3

—

LACM 117373

Anthill (LACM 1871)

—

2.2

—

1.9

—

2.2

—

LACM 117415

Anthill (LACM 1871)

1.8

LACM 117404

Anthill (LACM 1871)

1.7

LACM 117406

Anthill (LACM 1871)

1 6

UCMP 75554

Meniscomys hippodus
(UCMP V-6351)

2.0

1.9

1.8

1.5

UCMP 105101

Meniscomys hippodus
(UCMP V-66104)

1.7

2.0

2.1

UCMP 105104

Meniscomys hippodus
(UCMP V-66104)

—

1.8

2.1

u = unworn or only moderately worn occlusal dimension,
w = well worn or dimension taken near base of crown.

L = anteroposterior length.

W = transverse width.

1, 2, 3, = Mi, M2, Ms.

* by far the largest molar in the anthill collection,
t measurements taken from published illustration.

Lower First and Second Molars

The lower molars of the Deep River form are larger than those
of the anthill collection but slightly smaller than those of the
Lemhi collection (Table 3). In Mu2 of UWBM 38103 (Fig. 2a),
an anterior fossettid is bounded posterolingually by the meta-
lophulid II, which extends to the apex of the metaconid. The
anterior fossettid in SDSM 6277 and the anthill and Lemhi col-
lections is similarly formed.

In Meniscomys from the John Day Formation, only the labial
arm of the metalophulid II is present. A slight expansion in the
metalophulid II is present at the center of the tooth in the primi-
tive species but no crest extends to the metaconid. In some indi-
viduals the expansion is stronger and restricts the anterior fosset-
tid, but in others the fossettid is unrestricted. Occasionally the
metaconid in M. hippodus is expanded and contacts the meta-
lophulid II to create a small anterior fossettid in late stages of
wear. In M. hippodus, the metalophulid II is continuous with a
crest from the mesostylid; in UWBM 38103, the Lemhi collec-
tion, the anthill collection and SDSM 6277 the mesostylid crest
appears to be an accessory to the metalophulid II, for it joins the
latter at an angle.

The metastylid crest is stoutly developed in the Deep River
form (Fig. 2a). It is slightly expanded opposite the middle of the
tooth, in the position occupied by the mesostylid in primitive
aplodontoids. In Mi the crest extends 0.4 mm posterior to a

transverse line through the center of the mesostylid expansion,
thereby partially closing the lingual inflection. The metastylid
crest of SDSM 6277 and the anthill collection (Fig. 3e) is similar
but more lightly developed and seldom extends beyond the meso-
stylid. Though worn, the crest in ISU 18594 appears to have
been less prominent than that in the Deep River form. The meta-
stylid crest is absent in Meniscomys hippodus. However, in more
primitive individuals of Meniscomys from the John Day Forma-
tion, the metastylid crest may be present as a very rudimentary
structure.

An anterolingual fossettid is present in the Deep River form,
the Lemhi collection and SDSM 6277, and is variably present in
the anthill collection. It is bounded by the metalophulid II, the
diagonally directed mesostylid crest, and the metastylid crest.
When the anterolingual fossettid is not complete in the anthill
collection, this is usually the result of weak development of the
mesostylid crest. In Mi of the Deep River specimen, the meso-
stylid crest from the metalophulid II is stout but barely makes
occlusal contact with the mesostylid. In M. hippodus an antero-
lingual fossettid is seldom formed because the metalophulid II
does not extend toward the metaconid.

In the specimen from the Deep River Formation, the Lemhi
collection, SDSM 6277, the anthill collection, and Meniscomys
hippodus, the central fossettid is the deepest of the four or five
fossettids. In the Mi of a primitive form of Meniscomys from the
John Day Formation, a lingual inflection is present but there is

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 312:1-18.

Rensberger: Early Miocene Promylagaulus

little or no indication of a crest tending to enclose a central fos-
settid. In more advanced forms a central fossettid is formed in
Mi. In the M2 of Meniscomys the central fossettid is usually
closed.

The lingual inflection in Mi and M2 of the Deep River form is
closed at the lingual extremity by a low crest forming a basin or
shelf at the base of the inflection, well below the closure of the
central fossettid (Fig. 2a). A small, shallow fossettid is formed at
the base of this inflection on Mi, but is not quite formed on M2.
In SDSM 6277, ISU 18594 and most of the anthill collection, the
shelf is absent and the inflection usually fades out at depth. A
shelf may or may not be present in Meniscomys hippodus; in
primitive specimens of Meniscomys from the John Day Forma-
tion a shelf tends to be present but does not enclose a fossettid.

The mesoconid in the Deep River specimen is a triangular
cusp, with a heavy labial spur that joins the anterior margin of
the hypoconid to enclose a very small, shallow posterolabial fos-
settid (Fig. 2a). The configurations in SDSM 6277 and the anthill
collection are similar except that the mesoconid in Mi is smaller
and less distinct (Fig. 3e). The mesoconid spur is directed antero-
labiad in the Deep River specimen, whereas it runs posterolabiad
in SDSM 6277 and strongly so in the anthill specimens (Fig. 3e).
In Meniscomys hippodus the mesoconid is a distinct cusp, but the
structure connecting it with the hypoconid is a thinner crest than
that in Promylagaulus .

The hypoconid in the Deep River form, the Lemhi collection,
SDSM 6277, and the anthill collection is posterolabialiy com-
pressed and, as in Pa, expanded anteriorly (Figs. 2a, 3e). In the
Deep River specimen the anteroposterior expanse of the labial
inflection is less than in the other specimens and more deeply
incised medially, especially in Mi. The posterolophid is poste-
riorly expanded and convex in the specimens from Montana,
Idaho and South Dakota, but the convexity is most pronounced,
especially at the transverse midpoint of the crest, in the Deep
River form. The hypoconid of Meniscomys hippodus is expanded
rather than compressed posterolabialiy, and the posterior surface
of the posterolophid is relatively straight.

The posterior fossettid in the specimen from Montana is poly-
gonal in shape, owing to prominence of the hypoconulid and
entoconid, with the long diameter aligned anterolabially. In
SDSM 6277 and the anthill collection, the fossettid has the same
orientation but is more smoothly oval in outline. In Meniscomys
hippodus the posterior fossettid is more compressed than in any
of the other forms and the long axis is aligned anteroposteriorly.

Mi in the lightly worn dentition from the Deep River Forma-
tion, UWBM 38103, as in the intermediately worn specimen
from the Lemhi Valley (ISU 18594), appears somewhat larger
than M2 (Fig. 2a). In the heavily worn dentition from Montana,
UWBM 28075, Mi is slightly smaller in occlusal dimensions
than M2 (Fig. 2b). Mi in the little worn dentition of SDSM 6277
is slightly larger at the occlusal surface than M2. The isolated
lower molars of the anthill collection vary greatly in size, prob-
ably in part due to such positional variation. In little worn denti-
tions of Meniscomys from Oregon, Mi and M2 are almost equal
in size. However, in well worn dentitions Mi becomes relatively
smaller, especially in anteroposterior length, than in UWBM
38075.

At the late stage of wear represented in the Deep River form
(Fig. 2b), the labial inflection has disappeared and the occlusal
edge of the labial enamel forms a thick crest extending from the
anterolabial corner of Mi or M2 across the labial face and onto
the posterior face. The remaining three-fifths or more of the

occlusal margin in Mi lacks enamel. An isolated bit of lingual
enamel persists in M2. The only fossettid remaining in these teeth
is the central one in M2. The molars at this stage are reduced in
size and rather oval in shape.

In the teeth from the anthill collection, the basal enamel of the
hypoconid region extends more prominently posterolabiad than in
UWBM 38075, giving the teeth an angular outline that persists
with wear. Furthermore, the posterior surface at depth is invag-
inated, marking a division between the vestigial roots which is
not indicated in UWBM 38075 nor ISU 18594. In Meniscomys
hippodus, the labial inflection persists into late stages of wear,
and the teeth retain an angular outline.

Full measurement of the degree of hypsodonty in the molars
from the Deep River Formation is impossible because the enamel
is either partially hidden by alveolar bone, or much of it has been
worn away. Judging from the amount of enamel remaining on M2
of UWBM 38075, the ratio of enamel length to tooth width was
greater than that in a specimen of the Lemhi collection (Fig. 5
and Table 4). The relative length of enamel is greater in the ant-
hill collection than in the Lemhi collection. An isolated molar
from another locality (SDSM V59) is less hypsodont than that of
the Lemhi collection or those of the anthill collection and may be
more closely related to P. riggsi. The depth of the labial enamel
in Meniscomys hippodus is one-third or less that in the Lemhi or
anthill collections. Although the type of Promylagaulus riggsi
lacks the lower dentition, an indication of the degree of hypso-
donty of this specimen relative to that of the other collections is
obtainable from the upper dentition, described in a later section.

A thin layer of cement extends over the lower part of the
enamel in several of the lower molars of the anthill collection,
and a rather heavy layer extends at least 1.3 mm above the base
of the lingual enamel on UM 5193 of the Lemhi collection.
Cement extends perhaps 0.5 mm up the lingual enamel of M2 in
UWBM 38075.

The first and second molars of AMNH 56333, from Darton's
Bluff, lack fossettids, owing to the advanced stage of wear.
These teeth are oval in occlusal outline — slightly more so than
in the less worn molars of UWBM 38075, and differ in this
respect from the angular pattern in the anthill collection. In con-
trast to UWBM 38075, the lingual and posterior enamel in this
specimen still persists in spite of the deeper stage of wear with

WOLFF RH.

ANTHILL OJ

DEEP R. lUl

LEMHI j EJ

V 59 , 1 I „ . , , . , , , , , , „ . , — ,

0.80 0 84 088 092 096 1,00 1.04 1.08 1.12

Figure 5. Frequency distributions of values of ratio of depth of labial
enamel below highest point of posterior dentinal tract to transverse width
of Mi or Mz in Promylagaulus. Localities are those in Fig. 4, plus SDSM
V59. Base measurements for classes are listed. Arrow indicates incom-
plete measurement.

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 312:1-18.

Rensberger: Early Miocene Promylagaulus

TABLE 4

Depth1 of Labial Enamel on Mi, M2 (mm)

DEI DE2

MUSEUM NUMBER

LOCALITY, TAXON

DEI

DE2

UWBM 38075

Deep River Fm. (UWA 5867)

—

2.0+

—

>0.95

ISU 18594

Lemhi Valley (ISU 59003)

—

2.0

—

0.95

LA CM 117435

Anthill Collection (LACM 1871)

—

2.5

—

1.14

LACM 117441

Anthill Collection (LACM 1871)

2.1

—

1.11

LACM 117373

Anthill Collection (LACM 1871)

—

2.1

—

1.17

LACM 9339

Wolff Ranch (LACM 20051)

1.9

—

1.27

—

SDSM 59158

Monroe Creek Fm. (SDSM V59)

—

2.1

—

0.83

UCMP 105066

Meniscomys hippodus (UCMP V-6506-1)

—

0.7

—

0.37

UCMP 105058

Meniscomys hippodus (UCMP V-6506-2)

—

0.6

—

0.33

'Measured from level of apex of posterior dentinal tract to lowermost margin of labial enamel.

DEI = Depth of labial enamel on Mi .

DE2 = Depth of labial enamel on M2.

W1 = Transverse width of enamel on Mi.

W2 = Transverse width of enamel on M2.

respect to the bases of the lakes. This suggests that the dentinal
tracts may not be as vertically extensive as in UWBM 38075.

Lower Third Molar

This tooth is missing in the little worn dentition of the Deep
River Formation, UWBM 38103, but is just emerging from the
alveolus in SDSM 6277 and is present in ISU 18594. Several
third molars are recognizable among the isolated teeth of the ant-
hill collection. The mesostylid crest in the M3 of SDSM 6277
and specimens of the anthill collection is, unlike the development
in M,_,, absent or incomplete. Metalophulid II extends from the
ectolophid near the protoconid to the metaconid. An anterolin-
gual fossettid is therefore not formed in these specimens. In ISU
18594 the mesostylid crest is complete. In Meniscomys hippodus
the mesostylid crest is complete, as in Mj_2.

The region of the posterior fossettid in the M3 of SDSM 6277,
the anthill collection and ISU 18594 is shallower and anteropos-
teriorly shorter than that of M,_2. This correlates with the size of
the talonid, which is restricted in anteroposterior length. The
posterolophid and ectolophid join at the hypoconid in an acute,
labially prominent V. in correlation with a posterolabial flattening
of the hypoconid. In Meniscomys hippodus the hypoconid is not
flattened on its posterior surface, which is strongly convex, and
the axis of the posterior fossettid is anteroposteriorly aligned. The
hypolophid is frequently weak or absent in the specimens from
South Dakota, but is present and moderately strong in Promyla-
gaulus lemhiensis .

The labial inflection of M3 in ISU 18595, SDSM 6277 and the
anthill collection is more expansive anteroposteriorly than in
M1-2. No posterolabial fossettid is enclosed between hypoconid
and mesoconid. A distinct labial spur is present on the mesoconid
of ISU 18594 but is reduced or absent in the samples from South
Dakota. The labial inflection of M3 in Meniscomys hippodus is
little if any larger than that of M1-2, and a posterolabial fossettid
is present.

The M3 of ISU 18594, SDSM 6277 and the anthill collection

is distinctly smaller in occlusal dimensions than M,_2. In Menis-
comys hippodus the size of M3 more closely approximates that of
the anterior molars.

Upper Incisor

The upper dentition of the Deep River taxon is unknown but in
the Lemhi Valley collection all upper teeth are represented. The
anthill collection contains teeth representing P4, M1, M2, M3, but
not P3 (L. Macdonald 1972:25-26). The dentition in the type of
Promylagaulus riggsi (FMNH P26256) consists of the upper
incisor, P3, and well worn P4-M3.

The upper incisor in the Lemhi Valley collection is gently con-
vex across the anterior face, the medial and lateral margins of
which turn posteriad at approximately equal angles. The tooth
closely resembles that of Promylagaulus riggsi, except that it is
larger. The upper incisor in Meniscomys is similarly proportioned
but appears to be anteroposteriorly relatively shorter in cross sec-
tion.

Upper Third Premolar

The P3 of Promylagaulus riggsi and the Lemhi collection is
more slender and strongly compressed than that of Meniscomys.
The crown lacks a basal swelling which is characteristic of
Meniscomys. The long axis of the occlusal outline trends antero-
labiad (Fig. 6a), whereas that in Meniscomys is aligned anteriorly
or anterolingually. There is no distinct anterior or posterior carina
on the crown, whereas both structures are present in Meniscomys.
The tooth is curved from root to apex of crown, with the concave
side facing anterolinguad, whereas the axis in Meniscomys is
straight. The axial curvature in P. riggsi and UM 5285 places the
apex of the tooth well forward of P4, so that occlusal contact with
P-4 during mastication may have been impossible. The worn sur-
face in P. riggsi is polished and rounded, suggesting that only
food abrasion, not contact wear, has occurred. In the Lemhi
specimens no wear is apparent. A distinct facet, indicating con-
tact wear, is characteristic of the P3 in Meniscomys.

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 312:1-18.

Rensberger: Early Miocene Promylagaulus

Figure 6. Promylagaulus, stereo views of upper cheek teeth. a-P. riggsi, occlusal view left P3-M2, FMNH P26256; b-P ovatus.
occlusal view little worn left P4, LACM 117434, locality LACM 1871; c-P. ovatus, occlusal view worn left P4, LACM 117378,
locality LACM 1871; d-P. riggsi, posterior view left M2, FMNH P26256; e-P . ovatus, occlusal view left M1 or M2. LACM 1 17428.
locality LACM 1871;/-same, posterior view. Scale = 1 mm.

Upper Fourth Premolar

The P4 of the Lemhi collection is larger than that of the anthill
collection. This tooth is smallest in Promylagaulus riggsi (Table
5). The anterocone, the anterolabial cusp characteristic of the P4
of aplodontoids (Fig. 6a-c), is more lingually expanded in the
specimens from the anthill locality and Lemhi Valley than in
Meniscomys, and extends into the area occupied in the latter by a
distinct cusp, the anterostyle. The anterostyle is poorly developed
and sometimes absent in specimens of the anthill collection. The
parastyle and mesostyle (Fig. 6a- c) are thinner and less prom-
inent labially than in Meniscomys, and virtually absent in some
individuals. A distinctive feature, shared by Promylagaulus
riggsi and the specimens of the anthill and Lemhi collections, is a
slight lingual displacement of the labial enamel just anterior to
the mesostyle. The planes of vertical enamel in front of and pos-
terior to the mesostyle more nearly approximate one another in

Meniscomys. The worn occlusal outline of the tooth in the anthill
and Lemhi collections (Fig. 6c) lacks much of the anterolingual
notch between the region of the anterocone and the region of the
protocone in Promylagaulus riggsi (Fig. 6a), making the tooth
appear more oval. In the worn P4 of P. riggsi (Fig. 6a), the plane
of wear in the region of the parastyle and anterocone is more
steeply inclined posteriad than in the teeth of the Lemhi Valley
and anthill collections (Fig. 6c). No appression facet for P3 is
present on P4 in the specimens of the anthill collection, which
suggests that, as in P. riggsi and the Lemhi collection, the crown
of P3 does not occlude with P4 during normal mastication. An
appression facet is normally present in Meniscomys hippodus.

A conspicuous difference in the occlusal pattern from that in
Meniscomys is the presence in all specimens of Promylagaulus of
extreme anteroposterior length in the protocone, and correspond-
ing lingual flatness of the cusp (Fig. 6b). The outline of the worn
tooth in Promylagaulus is normally elongate anteroposteriorly

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 312:1-18.

Rensberger: Early Miocene Promylagaulus

TABLE 5

Length and Width of Upper Cheek Teeth (mm)

MUSEUM

NUMBER

LOCALITY, TAXON

LP4 /

WP4/

LM1/

WM1/

LM2/

WM2/

UM 4038

P. lemhiensis (UMV 7303)

4.0

3.3

2.1(H)

2.9

2.4

2.7

UM 5193

Lemhi Valley (UMV 7303)

—

2.2(H)

2.9

2.2

2.7

UM 5285

Lemhi Valley (UMV 7303)

4.4

3.7

2.5(H)

3.0

2.7

2.8

ISU 18601

Lemhi Valley (ISU 18601)

—

3.9

2.9(H)

3.3

2.4

2.8

LACM 117376

Anthill (LACM 1871)

3.5

2.6

—

LACM 117377

Anthill (LACM 1871)

3.5

2.4

—

LACM 117378

Anthill (LACM 1871)

3.5

2.8

—

LACM 117380

Anthill (LACM 1871)

3.5

2.8

—

LACM 117418

Anthill (LACM 1871)

—

1.9

2.0

—

LACM 117422

Anthill (LACM 1871)

—

2.0

1.9

—

LACM 117425

Anthill (LACM 1871)

—

1.8

1.7

—

LACM 117374

Anthill (LACM 1871)

—

2.0

LACM 117419

Anthill (LACM 1871)

—

2.0

1.9

LACM 1 17424

Anthill (LACM 1871)

—

1.9

1.8

LACM 117417

Anthill (LACM 1871)

—

2.1

2.0

FMNH P26256

P. riggsi

3.3

2.6

1.5

2.0

1.7

2.0

LP4/ =
WP4/ =
LM1/ =
WM 1/ =
LM2/ =
WM2/ =

Anteroposterior length of P4, at occlusal surface.

Transverse width of P4 at level of labial flexure of crown (below basal swelling).
Anteroposterior length of M1 at occlusal surface (wear: H = heavy).

Transverse width of M1 at occlusal surface.

Length of M2 at level of labial flexure of crown.

Transverse width of M2 at level of labial flexure of crown (below basal swelling).

(Fig. 6a- c), whereas the anteroposterior and transverse dimen-
sions are almost equal in Meniscomys. The paracone, proto-
conule, and metaconule are also quite elongate anteroposteriorly
and exhibit flattened labial and lingual surfaces. This trend
toward lophodonty is advanced over that known fox Meniscomys,
although a similarly directed trend has commenced in the latter
(Rensberger. in manuscript).

The crown of the premolar in the anthill and Lemhi collections
and Promylagaulus riggsi is curved along the vertical axis, which
is lingually convex and labially concave (Fig. 6b). Both labial
and lingual surfaces are vertically convex in Meniscomys, so that
the axis appears rather straight. The anterolingual fossette is alto-
gether absent in one unworn specimen of the anthill collection,
frequently disappears in early wear (Fig. 6c), and is absent in
known specimens of the Lemhi collection (Nichols 1976). This
fossette is present in both P. riggsi and Meniscomys . The area
normally occupied by the anterolingual fossette in most aplodon-
toids is partially reached in the specimens of Promylagaulus lack-
ing it by the anteriorly elongate posterolingual fossette and the
lingually expanded anterolabial fossette. The posterolingual fos-
sette is aligned almost anteroposteriorly in the Lemhi and anthill
collections (Fig. 6c), whereas it slants anterolabially in P. riggsi
(Fig. 6a) and Meniscomys.

Upper Molars

The upper molars, like the premolar, are largest in the Lemhi
collection and smallest in Promylagaulus riggsi (Fig. 7). The
molars of P. riggsi, especially M1-2, are rather rectangular in
occlusal outline, and the anteroposterior length is less than the

width (Fig. 6a). In the anthill and Lemhi collections the length
and width are subequal (Fig. 6e). Poor development or lack of
the mesostyle, at least in late wear, and limited lingual convexity
contribute to the rectangular shape. In the little worn molars of
the anthill collection, the anteroposterior length may be greater
than the width, but with wear the width increases to essentially
the length. In the stage of wear in which the labial (vertical)
flexure of the crown has reached the occlusal surface, the length
in the anthill collection is approximately equal to the width,
whereas the width exceeds the length at the same stage in the
Lemhi collection and P. riggsi (Table 5). In all groups including
Meniscomys, the anteroposterior length of M1 shortens more
rapidly with wear than does that of M2. The molars in Menis-
comys bear strong mesostyles and strongly convex lingual mar-

LEMHI I I I I

ANTHILL I I

P. RIGGSI 1 I i i j ■

1.5 2.0 2.5 3.0

Figure 7. Frequency distributions of anteroposterior length of M1 in
specimens of Promylagaulus at localities Lemhi Valley (UM V-7303 =
1SU 59003) and Anthill (LACM 1871), with value for type of P riggsi.
Base measurements for classes are listed.

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 312:1-18.

Rensberger: Early Miocene Promylagaulus

TABLE 6

Depth1 of Lingual Enamel on M1, M2 (mm)

MUSEUM NUMBER

LOCALITY, TAXON

DEI/

DE2/

DEI/

WM1/

DE2/

WM2/

LACM 1 17417

Anthill (LACM 1871)

2.9

1.38

LACM 117419

Anthill (LACM 1871)

—

2.3

—

1.15

LACM 117424

Anthill (LACM 1871)

2.2

—

1.16

—

LACM 117428

Anthill (LACM 1871)

—

2.3

—

1.21

ISU 18601

Lemhi (ISU 59003)

2.0+

1.9+

0.61 +

0.56 +

UM 5285

Lemhi (UMV 7303)

—

2.2

—

0.82

UM 4038

P. lemhiensis (UMV 7303)

—

2.4+

0.89+

FMNH P26256

P riggsi

—

1.4

—

0.70

UWBM 29232

Meniscomys hippodus
(UWA 5183-2)

0.5

0.23

'measured from level of apex of posterior enamel-dentine chevron to uppermost margin of lingual enamel.

WM1/ = Transverse width of M1 near apex of posterior dentinal tract.
DEI/ = Depth of lingual enamel on ML

WM2/ = Transverse width of M2 near apex of posterior dentinal tract.
DE2/ = Depth of lingual enamel on M2.

gins. The width of M’ or M2 in Meniscomys is greater than the
length in all but the earliest stages of wear. The transverse
dimension of the anterior margin of the tooth in the three groups
of Promylagaulus is only slightly if at all less than that of the
posterior margin (Fig. 6e), whereas a very narrow anterior
dimension is characteristic of Meniscomys. With moderate wear,
only the posterolingual fossette remains in Promylagaulus.
whereas several of the smaller fossettes are present until late
stages of wear in Meniscomys.

The unworn M3 in the anthill collection contains anterolabial,
anterolingual, posterolabial and posterolingual fossettes, but no
central fossette. The posterolabial fossette is usually the largest
and deepest, although in several specimens the posterolingual
fossette is dominant. A single, probably posterolabial, fossette is
present in the well worn M3 of Promylagaulus riggsi and the
Lemhi collection. In Meniscomys the central fossette is present
and moderately persistent, and the posterolabial fossette is the
largest and deepest.

ANTHILL

n n

As in P4, the crowns of the molars in Promylagaulus are verti-
cally curved with the concave side labial (Fig. 6f). No compara-
ble curvature is apparent in Meniscomys. Long dentinal tracts
extend down the anterior and posterior sides of the molars in the
anthill collection. A tract is visible on the posterior side of the
left M2 in P. riggsi (Fig. 6d) but is shallower than the corre-
sponding plate in specimens of the anthill collection (Fig. 6f).
The margins of the chevron- shaped enamel edges bordering the
dentinal tracts are more steeply inclined in the anthill specimens
(Fig. 6f) than in P. riggsi (Fig. 6e) and suggest greater hypso-
donty. Furthermore, the lingual enamel plate, measured from the
apex of the chevron (lowest part of the tract), is considerably
longer in the anthill specimens than in P. riggsi (Fig. 8 and Table
6). This difference in enamel length is, for example, approxi-
mately equivalent to the change in length of the enamel of the
upper and lower molars spanning three chronologically succes-
sive taxa of early Miocene pocket gophers, Entoptychus minor,
E. cavifrons, and E. transitorius (Rensberger 1971:79-85). The
depth of the lingual enamel relative to tooth width on M2 is
apparently intermediate in the Lemhi collection between that of
P. riggsi and that in the anthill collection (Fig. 8 and Table 6).
The roots of the upper molars in the anthill collection are greatly
reduced compared to those in the Lemhi collection and P. riggsi.

LEMHI 1 *! I f~L

P. RIGGSI . . . I I

0.6 0.7 0.6 0.9 1.0 1.1 1.2 1.3 1.4

Figure 8. Frequency distributions of ratio of depth of lingual enamel
above lowermost point of posterior dentinal tract to width of M1 or M2 in
Promylagaulus . Localities as listed for Fig. 7. Base measurements for
classes are listed. Arrows indicate incomplete measurements.

SYSTEMATIC PALEONTOLOGY
Order RODENTIA

Superfamily Aplodontoidea Matthew 1910
Family Mylagaulidae Cope 1881
Promylagaulus McGrew 1941

GENOTYPIC SPECIES: Promylagaulus riggsi McGrew 1941.
DEFINITION (REVISED): More hypsodont than Meniscomys.
P3 slender, curving anterolinguad, with long axis of cross section
slanting anterolabiad, lacking occlusal contact. P4 with lingually

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 312:1-18.

Rensberger: Early Miocene Promylagaulus

expanded anterocone, parastyle and mesostyle more reduced than
in Mylagaulodon, ectoloph anterior to mesostyle displaced
linguad, protocone expanded anteroposteriorly and relatively flat
lingually, posterolabial fossette smaller, less persistent than in
other mylagaulids or Meniscomys, anterolingual fossette reduced
or absent, posterolingual fossette extended craniad, anterolabial
fossette expanded linguad.

M'~2 rectangular in occlusal outline, at least in earlier stages of
wear; with mesostyle reduced or absent, tendency for anteropos-
terior elongation of lower part of crown, persistence of postero-
lingual fossette long after others have worn away. P4-M2 with
vertically elongate, curved (labially concave) crowns, anterior
and posterior dentinal tracts.

P4 with long, strong metastylid crest from metaconid to meso-
stylid, enclosing lingual fossettid; mesostylid usually indistinct;
mesoconid flattened labially in mature stages of wear; poor
development of anterior branch of labial inflection, smaller than
in other mylagaulids or Meniscomys, extensive development of
posterior branch; posterolabially compressed, anteriorly expanded
hypoconid, narrow entrance to labial inflection.

Mi with strong central fossettid. M,_2 with strong metastylid
crest; hypoconid posterolabially compressed, anteriorly
expanded; posterolophid posteriorly convex; posterior fossettid
trending anterolabiad; metalophulid II extending to apex of meta-
conid; crest from mesostylid bending craniad to join center of
metalophulid II; anterior and lingual fossettids present; meso-
stylid indistinct, lacking lingual prominence, especially on Mi;
tendency to lose rectangular outline and labial inflection in late
wear.

M3 with hypoconid posteriorly flattened; labial inflection rela-
tively larger anteroposteriorly than in Mu2; absolute occlusal
dimensions much smaller than those of M,_2; no mesostylid crest;
metalophulid II extending from metaconid to protoconid; no pos-
terolabial fossettid.

SPECIES: P. riggsi McGrew, L. Mioc. S. Dak.; P. ovatus (n.
sp.) L. Mioc., S. Dak.; P. lemhiensis Nichols, L. Mioc. Idaho;
P. montanensis (n. sp.) L. Mioc. Montana.

Promylagaulus riggsi McGrew 1941

(Fig. 6a,d)

McGrew, P.O., 1941, Figs. 1, 2c, 3.

TYPE; Skull, lacking anterior extremities of nasals, zygomatic
arches and cranium posterior to orbits; with right and left I,
P3-M2; right M3; FMNH P26256.

TYPE LOCALITY; Four miles south of Porcupine, South Da-
kota.

STRATIGRAPHIC POSITION: “Top of lower Rosebud
beds” (McGrew 1941:6); “Probably Rosebud Formation”
(J.R. Macdonald 1963:179). Monroe Creek Formation or Harri-
son Formation or Rosebud Formation (L. Macdonald 1972:25).

AGE: Arikareean or Hemingfordian, early or middle Miocene,
based on the range of possible stratigraphic positions cited above.
Older than late Arikareean, based on comparative stages of
advancement of other species.

REFERRED SPECIMEN: SDSM 59158, M2, from locality
SDSM V59.

DIAGNOSIS (REVISED): Smaller than other species. Worn
occlusal surface of anterocone on P4 with pronounced inclination
posteriad; anterolingual fossette of P4 present in known speci-

men; posterolingual and anterolabial fossettes more restricted
than in P. ovatus, P. lemhiensis; posterolingual fossette with
strong anterolabial slant; worn occlusal outline with distinct notch
separating anteriorly prominent anterocone from protocone; with
angular posterolingual corner.

Anteroposterior length of M1 and M2 becoming less than trans-
verse width well before wear reaches labial tlexure of crown;
ratio of depth of lingual enamel on M2 to width of tooth 0.70,
less than in other species; sides of posterior dentinal tract of M1-2
less vertical than in other species.

Promylagaulus lemhiensis Nichols 1976

Nichols, R., 1976, Figs. 7, 8b; PI. 1, Figs. 5-10.

TYPE: Fragment of rostrum and palate with root of right I,
worn left P3-M2, UM 4038.

TYPE LOCALITY: Big Wash (Univ. of Montana locality MV
7303, in part). NW'/aNW1/) sec. 17, T. 17 N., R. 25 E., Lemhi
quadrangle, Idaho.

STRATIGRAPHIC POSITION: Specimens from middle part
of 400 foot section, Peterson Creek beds, an informal unit
(Nichols, 1976).

ZONE: Upper Meniscomys Concurrent-range Zone (Nichols,
1976:13).

AGE: Arikareean, early Miocene.

REFERRED SPECIMENS: UM 5193, cranial and mandibular
fragments with worn M^“3; UM 4039, worn P4; UM 4055, little
worn P4; UM 5285, partial skull with I, worn P3-M3; ISU
18594, mandible with little worn P4, moderate to little worn
M,_3; ISU 18601, associated P4 (fragment), M1-3, P4. All from a
series of closely related exposures. University of Montana local-
ity MV 7303 (= ISU 59003); ISU 18594 from stratigraphically
21 m above type specimen.

DIAGNOSIS (REVISED): Overall size greater than that of P.
riggsi, P. ovatus. Occlusal surface of anterocone on P4 nearly
horizontal in mature stages of wear; anterolingual notch, postero-
lingual comer of tooth less pronounced than in P. riggsi, making
occlusal shape more oval; posterolingual fossette aligned more
nearly anteroposteriorly than in P. riggsi; tooth in known speci-
mens lacking distinct protoconule and anterolingual inflection.

Transverse width of M1-2 greater than anteroposterior length at
level of labial flexure of crown, as in P. riggsi, unlike P. ovatus;
ratio of depth of lingual enamel of M2 above apex of posterior
dentinal tract to width of tooth 0.82-0.89+, greater than in P.
riggsi, less than in P. ovatus. Root branches of M1-2 more prom-
inent than in P. ovatus.

Posterior and lingual fossettids of P4 wider relative to tooth
width (ratios 0.41, 0.30, resp.) than in P. montanensis; central
width of tooth greater relative to overall width than in P. mon-
tanensis (ratio 0.67); length (4. 7-5.0 mm) greater than in P.
montanensis, P. ovatus; ratio of depth of labial enamel to length
of tooth (.43-. 44) less than in P. ovatus, P. montanensis .

Ratio of depth of labial enamel on M2 to tooth width (0.95)
less than in P. ovatus. Metastylid crest of M12 with less poste-
rior prominence than in P. montanensis; no posterolingual fosset-
tid, unlike P. montanensis; labial inflection relatively more
expansive anteroposteriorly than in P. montanensis; central pos-
terior convexity less pronounced than in P. montanensis. Heavier
layer of cementum overlapping enamel of cheek teeth than in P.
ovatus, P. montanensis .

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 312:1-18.

Rensberger: Early Miocene Promylagaulus

Promylagaulus ovatus 1 new species

(Figs. 3a-f; 6b,c,e,f)

Macdonald, J.R., 1970, Fig. 14 (Promylagaulus cf. riggsi).

Macdonald, L., 1972:25-26 [Promylagaulus riggsi).

TYPE: Little worn left M2. LACM 117435.

TYPE LOCALITY: Anthills of LACM 1871 (=SDSM V6229)
SW V4, sec. 9, T. 39N., R. 42 W., east of Porcupine Creek,
Sharps Corner quadrangle. South Dakota.

STRATIGRAPHIC POSITION: Upper part of Monroe Creek
Formation (Harksen 1970). Fossils, although recovered from
anthills, apparently derived from two or three 0.6 cm thick bone
bearing layers in nearby vertical cliffs (Harksen, 1970; L. Mac-
donald 1 972:6) . Locality may be at transition between Monroe
Creek and Harrison formations (James E. Martin, pers. comm.).

ZONE: Entoptychus-Gregorymys Concurrent-range Zone
(Fisher and Rensberger 1972).

AGE: Arikareean, early Miocene.

REFERRED SPECIMENS: LACM 117351- 117370, 117372,
117392, 23530, 23532, 23535, 23536 (P4); 117373- 117375,
117403, 117406, 117410-117416, 117438-117441, 23534
(lower molars); 117384, 117436, 23637 (DP4); 117376- 117383,
117385- 117391, 117393- 117396,1 17434, 23538 (P4); 117371,
117397-117400, 117404, 117405, 117407, 117417-117433,
23528, 23529, 23531, 23534 (upper molars); from locality
LACM 1871.

SDSM 6277, right mandible, DP4-M3; from locality V6210.

LACM 9339, right Mi; LACM 9340, right P4; LACM 9342,
right P4; from locality LACM 2005, northeast of Wolff Ranch,
South Dakota.

DIAGNOSIS: Smaller than P. montanensis and P lemhiensis,
slightly larger than P. riggsi. Worn occlusal surface of antero-
cone on P4 less inclined posteriad than in P. riggsi; anterolingual
fossette of P4 only occasionally absent in unworn premolars, but
frequently disappearing with early wear; posterolingual fossette
expanded anteriad, with more nearly anteroposterior alignment
than in P. riggsi; anterolingual notch and posterolingual corner
less pronounced than in P. riggsi. making occlusal shape oval.

Transverse width of M1 or M2 approximately equal to antero-
posterior length at level of vertical flexure of labial surface; ratio
of depth of lingual enamel to width of tooth 1.15-1.21, greater
than in P. riggsi, P. lemhiensis; sides of dentinal tracts more
nearly vertical than in P. riggsi; root branches of less prominence
than in P. lemhiensis .

Posterior and lingual fossettids of P4 wider relative to tooth
width (ratios 0.32-0.36, 0.27) than in P. montanensis, possibly
narrower than in P. lemhiensis; ratio of depth of labial enamel to
length of tooth 0.49-0.54, less than in P. montanensis, greater
than in P. lemhiensis.

Ratio of depth of labial enamel on Mi or M: to tooth width
1.11-1.27, greater than in P. lemhiensis, P. riggsi; metastylid
crest with less posterior prominence than in P. montanensis; no
posterolingual fossettid, unlike P. montanensis; labial inflection
relatively more expansive anteroposteriorly, shallower trans-
versely than in P. montanensis; labial part of posterolophid less
extensive anteriorly than in P. montanensis; deep portion of pos-
terolophid more protuberant posterolabially (less posteriorly) than
in P. montanensis.

DISCUSSION: An isolated cheek tooth from the anthill local-

ity was selected as the type of this species in lieu of the mandible
from SDSM V6210, because of the much larger sample of Pro-
mylagaulus as well as other mammalian taxa at the anthill site.
Furthermore, the cheek teeth are poorly exposed in the young
mandible, leaving doubt as to the stage of advancement in hyp-
sodonty.

It seems likely from this evidence that P. riggsi, for which the
stratigraphic occurrence is uncertain, did not occur in a deposit
which was contemporaneous with those of the anthill locality and
other sites which have yielded P. ovatus. The preservation of the
specimens also suggests this conclusion. The specimens of Pro-
mylagaulus from the anthill locality, that from SDSM V6210,
and those from the Wolff Ranch locality (LACM 2005), all of
which are referable to P. ovatus, are of medium to dark or black-
ish brown color, whereas the teeth of P. riggsi are very light in
color.

Placement of the occurrences of Promylagaulus ovatus in the
Entoptychus-Gregorymys Concurrent-range Zone is based upon
the abundant association of Gregorymys formosus (L. Macdon-
ald 1972:30) and the advanced morphology of Promylagaulus
compared with that of Meniscomys. Meniscomys occurs only
stratigraphically beneath the Entoptychus-Gregorymys Concur-
rent-range Zone in the John Day Formation (Fisher and Rens-
berger 1972).

Promylagaulus montanensis new species

(Fig. 2)

TYPE: Right mandible with DP4, M,_2, incisor fragment;
missing upper part of ascending ramus. UWBM 38103.

TYPE LOCALITY: Spring Creek 1 (UWA 5867-1). SE-NW
trending draw 46°37.4' N, 1 1 1 °3 .6' W; SE14 sec. 14, R. 5 E.,
T. 10 N., Hanson Reservoir quadrangle, Montana.

STRATIGRAPHIC POSITION: Deep River Formation. Spec-
imens from stratigraphically lower of two units. Brownish tan,
massive sandstone.

ZONE: Entoptychus-Gregorymys Concurrent-range Zone.

AGE: Arikareean, early Miocene.

REFERRED SPECIMEN: Right mandible with worn P4-M3;
lacking incisor, ventral margin, condylar process and ascending
ramus; geographic locality and stratigraphic position exactly that
of type specimen. UWBM 38075.

DIAGNOSIS: Larger than P. ovatus and P. riggsi. Anteropos-
terior length of P4 4. 1 mm, less than in P. lemhiensis; ratios of
width of posterior and lingual fossettids to tooth width less than
in P. lemhiensis, P. ovatus; ratio of depth of labial enamel to
length of tooth 0.68, greater than in P. ovatus, P. lemhiensis;
ratio of central width to overall width of tooth 0.55, smaller than
in P. ovatus, P. lemhiensis .

Metastylid crest of M t_2 with greater posterior prominence
than inP. ovatus. P. lemhiensis; small posterolingual fossettid on
M2, incipient posterolingual fossettid on Mi, unlike P. ovatus, P.
lemhiensis; labial inflection anteroposteriorly more compressed,
transversely relatively deeper than in P. ovatus, P. lemhiensis;
central posterior convexity of posterolophid more pronounced
than in P. ovatus, P. lemhiensis.

DISCUSSION: The placement of this form in the Entopty-
chus-Gregorymys Concurrent-range Zone is based upon an asso-
ciated assemblage of taxa, including Gregorymys, to be described
later.

1 ovatus : L., egg shaped, in reference to the occlusal outline of P4.

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 312:1-18

Rensberger: Early Miocene Promylagaulus

Figure 9. A possible phylogeny for the described species of Promylagaulus which would explain the morphological and geographical differences. Arrows
designate chronologic intervals during which increases in size and hypsodonty presumably occurred. The stratigraphic position of P. riggsi is uncertain.

Promylagaulus sp.

McKenna and Love 1972, Figs. 15, 16.

LOCALITY: Darton’s Bluff (AMNH) Big Horn Mountains,
SW'ANE'A, sec. 19, T. 48 N., R. 85 W., Hazelton Peak quad-
rangle, Johnson County, Wyoming. Exposure along bluff near
head of Canyon Creek.

STRATIGRAPHIC POSITION: In 22 m thick, light tan, grad-
ing upward to light gray, massive, fine-grained sandstone within
43 m Miocene section.

ZONE: Entoptychus-Gregorymys Concurrent-range Zone.

AGE: Later Arikareean, early Miocene.

MATERIAL: Right mandible, P4-M2, I; left mandible
P4-M3, fragmentary incisor. Posterior parts of both rami miss-
ing. AMNH 56333.

DISCUSSION: Judging from the illustrations of this specimen,
it may be distinct from Promylagaulus ovatus, P. montanensis,
P. riggsi, and P. lemhiensis. Lingual enamel is present on the
occlusal surface at a stage of wear in which the occlusal outline is
oval, unlike the condition in P. ovatus. In late stages of wear in
P. ovatus, the originally rectangular outline remains until the lin-
gual enamel disappears. The Wyoming specimen more closely
resembles P. montanensis and P. lemhiensis in this character.

However, the size is smaller than that of P. montanensis, P. lem-
hiensis, or P. riggsi and the dentinal tracts seem to be less exten-
sive than in P. montanensis, judging from the greater continuity
of enamel surrounding the heavily worn molars. It appears to be
more hypsodont than P. riggsi.

That this occurrence seems to be within the Entoptychus-Greg-
orymys Concurrent-range Zone is based upon the presence of En-
toptychus as well as the association with what may be Pleurolicus
(McKenna and Love 1972:17-21). Although these geomyoids
are not contemporaneous in the John Day Formation, the range of
Entoptychus commences immediately above the range of Pleuro-
licus (Fisher and Rensberger 1972:24).

PHYLETIC RELATIONSHIPS
OF THE SPECIES

The greater size of the species of Promylagaulus from the
Rocky Mountains and the fact that the most primitive form, P.
riggsi, is the smallest suggests that the group may have origi-
nated in the Great Plains. P. ovatus, the youngest known form in
the Plains, was only slightly larger than P. riggsi but conspicu-
ously advanced in hypsodonty. The species occurring farthest
west, P. lemhiensis, is the largest but is only slightly more hyp-

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 312:1-18.

Rensberger: Early Miocene Promylagaulus

sodont than P. riggsi. These relationships suggest (Fig. 9) that an
early divergence from the primitive stock may have dispersed
westward and radiated again in the Rocky Mountain region. The
amount of time involved in this dispersal, diversification and evo-
lution was apparently not very great, judging from the occur-
rences of both P. montanensis and P. ovatus in the Entopty-
chus-Gregorymys Concurrent-range Zone, probably low in that
zone, and P. lemhiensis in deposits not much older.

RELATIONSHIPS OF PROMYLAGAULUS

McGrew (1941:26), followed by subsequent workers, placed
Promylagaulus in the Mylagaulidae. He believed, although with
a degree of uncertainty regarding several characters, that Promy-
lagaulus gave rise to Mylagaulodon and therefore to later myla-
gaulids. As McGrew noted, the upper premolar of P. riggsi
displays the same general features as FMNH P26266 and is in-
termediate in degree of hypsodonty between the latter and Menis-
comys hippodus. Wilson (1960:54) noted an apparent difference
in the number of lakes in the molar talonid of Promylagaulus and
Mesogaulus, but was uncertain as to whether wear in the speci-
men of Promylagaulus accounted for the discrepancy. Other
characters in the worn dentition of P. riggsi raise at least some
doubt about a relationship to the mylagaulids. The better repre-
sented taxa, P. ovatus and P . lemhiensis, unknown at the time of
those earlier studies, exhibit features which differ more clearly
from those of the mylagaulids.

The mesostyle in P4, an early character in aplodontids and
which in P. riggsi can be interpreted as almost worn away, is
much smaller in Promylagaulus than in Mylagaulodon (FMNH
P26266). The posterolabial fossette of P4 is small and is worn
away well before the other two major fossettes disappear in P .
ovatus, whereas in the mylagaulids this structure is as persistent
as the other fossettes, and in Mylagaulodon is large. The labial
inflection of P4 is undivided in P. ovatus and P. lemhiensis but is
divided in both Meniscomys and the mylagaulids (for the latter
see Black and Wood 1956, Fig. 1). The posterolabial fossettid of
the lower molars, the one queried by Wilson, is absent or vesti-
gial in the molars of Promylagaulus, relatively larger in Menis-
comys, and still larger in Mesogaulus (Wilson 1960, Fig. 59) and
Mylagaulus (McGrew 1941, Fig. 12). However, the posterolabial
inflection above the fossettid is formed differently in Mesogaulus
than in either Meniscomys or Promylagaulus, for in occlusal view
(Black and Wood 1956, Figs. 8B,E) the channel is elongate, ex-
tends strongly posterolinguad, and the anterolabial margin of the
hypoconid doesn’t hook toward the mesoconid but joins the base
of the protoconid to close the fossettid. In both Meniscomys and
Promylagaulus the posterolabial fossettid is small and oval, and
is formed by union of the anterolabial process of the hypoconid
with the mesoconid.

One of the distinctive characters of Promylagaulus is the pos-
terolabial flattening of the hypoconid in P4-M3. The hypoconid
is usually rather protuberant posterolabially in Mesogaulus (Wil-
son 1960, Figs. 56-59; Black and Wood 1956, Figs. 6, 8), but in
the unworn pattern of the molars may extend anterolabiad some-
what as in Promylagaulus .

On the other hand, Promylagaulus shares with the later myla-
gaulids reduction of the labial prominence of the stylar cusps in
the upper cheek teeth and the tendency for the occlusal outlines
to become oval. It also shares relatively large P;J and, unlike the
aplodontines, persistent fossettids. Although the reduction of sty-
lar structures may have occurred independently, these events may

have emerged from an early common adaptive trend and affinity.

The morphology of the mylagaulids and Promylagaulus seems
therefore to suggest that the mylagaulids may possibly have
evolved from a lineage leading eventually to Promylagaulus, as
opposed to others going toward other aplodontoids, but if so, the
ancestral form for Promylagaulus and the mylagaulids was so
primitive that the posterolabial inflection of the lower molars was
still unclosed — a stage more primitive than that of any aplodon-
tid presently known from the Meniscomys Concurrent-range
Zone.

Promylagaulus may represent a progressive group that was
part of a rather profuse radiation of aplodontoid rodents in the
early Miocene just prior to and during the Meniscomys Concur-
rent-range Zone, of which only a few groups, not Promylagaul-
us, survived into the middle Miocene.

ACKNOWLEDGMENTS

I wish to thank the following people for providing access to
specimens used in this study: Drs. Lawrence G. Barnes, Theo-
dore Downs, and David P. Whistler, LACM; Drs. Philip E.
Bjork and Morton Green, SDSM; Dr. William D. Turnbull,
FMNH; Dr. Robert W. Fields and Mr. Ralph Nichols, UM; and
Dr. John A. White, ISU. I am grateful to Drs. Craig C. Black
and Mary R. Dawson of the Carnegie Museum, Dr. Donald L.
Rasmussen of the Amoco Production Company, and Dr. David
P. Whistler of the LACM, who read the manuscript and offered
suggestions which greatly improved it. Mr. and Mrs. Ben Witte
and Mr. David Nunnallee were of much assistance in the field
work that produced the specimens from the Deep River Forma-
tion. This study was supported in part by N.S.F. Grant GB
35959.

LITERATURE CITED

Black, C.C. and A.E. Wood 1956. Variation and tooth re-
placement in a Miocene mylagaulid rodent. Jour. Paleontol-
ogy 30:672-684.

Fisher, R.V. and J.M. Rensberger. 1972. Physical stratigraphy
of the John Day Formation, central Oregon. Univ. Calif.
Publ. Geol. Sci., 101:45 p.

Harksen, J.C. 1970. Geology of the Sharps Corner quadrangle.
In: Macdonald, J.R., 1970. Review of the Miocene Wound-
ed Knee faunas of southwestern South Dakota. Los Angeles
County Mus. Nat. Hist. Bull. 8:82 p., 2 maps.
Macdonald, J.R. 1963. The Miocene faunas from the Wounded
Knee area of western South Dakota. Amer. Mus. Nat. Hist.
Bull. 125:141-238.

. 1970. Review of the Miocene Wounded Knee faunas

of southwestern South Dakota. Los Angeles County Mus.
Nat. Hist. Bull. 8:82 p., 2 maps.

Macdonald, L. 1972. Monroe Creek (early Miocene) microfos-
sils from the Wounded Knee area. South Dakota. South Da-
kota Geol. Surv. Rept. Investig. 105:43 p.

McGrew, P.O. 1941 The Aplodontoidea. Field Mus. Nat. Hist.
Geol. Ser. 9:1-30.

McKenna, M.C. and J.D. Love. 1972. High-level strata contain-
ing early Miocene mammals on the Big Horn Mountains,
Wyoming. Amer. Mus. Novitates 2490:31 p.

Nichols, R. 1976. Early Miocene mammals from the Lemhi Val-
ley of Idaho. Tebiwa 18:9-47.

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 312:1-18.

Rensberger: Early Miocene Promylagaulus

Rensberger, J.M. 1971. Entoptychine pocket gophers (Mamma-
lia, Geomyoidea) of the early Miocene John Day Formation,
Oregon. Univ. Calif. Publ. Geol. Sci. 90:209 p.

. 1975. Haplomys and its bearing on the origin of the

aplodontoid rodents. Jour. Mammalogy 56:1- 14.

. (manuscript). Aplodontoid rodents of the John Day

Formation .

Stirton, R.A. 1935. A review of the Tertiary beavers. Univ.

Calif. Publ. Bull. Dept. Geol. Sci. 23:391- 457.

Wilson, R.W. 1960. Early Miocene rodents and insectivores
from northeastern Colorado. Univ. Kansas Paleontological
Contr. Vertebrata 7:92 p.

Accepted for publication June 1, 1978.

Contrib. Sci. Natur. Hist. Mus. Los Angeles County. 1979. 312:1-18.

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