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CONTRIBUTIONS
IN SCIENCE
LOS
ANGELES
COUNTY
MUSEUM
December 31, 1966
TABLE OF CONTENTS
and
AUTHOR INDEX
1965 - 1966
Nos. 86-121
Los Angeles County Museum of Natural History • Exposition Park
Los Angeles, California 90007
2
Contributions in Science
TABLE OF CONTENTS
No. 86. Tooth terminology and variation in sharks with special reference
to the sand shark, Car char ias taurus Rafinesque, by Shelton P.
Applegate. 18 pp., 5 figs. April 9, 1965.
No. 87. Frog- like vertebrae from the lower Permian of southeastern Utah,
by Peter Paul Vaughn. 18 pp., 1 fig. June 28, 1965.
No. 88. Geolab is wolff i, a new fossil insect ivore from the late Oligocene
of South Dakota, by J. R. Macdonald. 6 pp., 1 fig. June 28, 1965.
No. 89. A new South American toe biter (Hemiptera, Belostomatidae ), by
Arnold S. Menke. 4 pp. , 2 figs. June 28, 1965.
No. 90. Normichthys yahganorum , a new searsiid fish from Antarctic waters,
by Robert J. Lavenberg. 7 pp. , 2 figs. June 28, 1965.
No. 91. Observations on captive and wild Atlantic bottlenosed dolphins,
T ursiops truncatus, in the northeastern Gulf of Mexico, by Melba
C. Caldwell, David K. Caldwell, and J. B. Siebenaler. 10 pp., 1
fig. June 28, 1965.
No. 92. The Barstovian Camp Creek fauna from Elko County, Nevada, by J. R.
Macdonald. 18 pp., 7 figs. April 4, 1966.
No. 93. A key to the species of Ophiuroidea (Brittle Stars) of the Santa
Monica Bay and adjacent areas, by Richard A. Boolootian and David
Leighton. 20pp., 31 figs. April 4, 1966.
No. 94. Pliocene birds from Chihuahua, Mexico, by Hildegarde Howard. 12
pp., 1 fig. April 4, 1966.
No. 95. Observations on the behavior of wild and captive false killer
whales, with notes on associated behavior of other genera of
captive delphinids, by David H. Brown, David K. Caldwell, and
Melba C. Caldwell. 32 pp., 13 figs. April 4, 1966.
No. 96. A new Peromyscus from the late Pleistocene of Anacapa Island,
California, with notes on variation in Peromyscus nesodytes Wilson,
by John A. White. 8 pp. , 4 figs. April 4, 1966.
No. 97. A new species of Heterant hidium from California (Hymenoptera :
Megachi 1 idae ), by Roy R. Snelling. 8 pp., 1 fig. May 5, 1966.
No. 98. Studies on North American bees of the genus Hylaeus 1. Distribution
of the western species of the subgenus Prosopis with descriptions
of new forms (Hymenoptera: Col let idae), by Roy R. Snelling. 18
pp., 3 figs. May 5, 1966.
No. 99. Hie California species of Philorus: taxonomy, early stages and
descriptions of two new species (Diptera: Blepharocer idae ) , by
Charles L. Hogue. 22 pp., 10 figs. May 5, 1966.
No. 100. A new genus of Fissurell idae and a new name for a misunderstood
species of west American Diodora, by James H. McLean. 8 pp. , 1
fig. May 5, 1966.
No. 101. A possible ancestor of the Lucus Auk (Family Mancallidae) from
the Tertiary of Orange County, California, by Hildegarde Howard.
8 pp., 1 fig. May 5, 1966.
No. 102. Anew Syrrhophus from Mexico (Amphibia: Leptodacty 1 idae ) , by
James R. Dixon and Robert G. Webb. 5 pp. , 1 fig. May 5, 1966.
No. 103. A new species of Boetica from the Pliocene of California, by
George P. Kanakoff. 4 pp., 3 figs. May 5, 1966.
Bound by DOBBS BROS. LIBRARY BINDING CO., INC., St. Augustus
Table of Contents
3
j>
j _____
«. No. 104.
D
No. 105
3* No. 106.
fl)
3
No. 107.
No. 108.
No. 109.
No. 110.
No. 111.
No. 112.
No. 113.
No. 114.
No. 115.
No. 116.
No. 117.
No. 118.
No. 119.
No. 120.
No. 121.
Observations on the distribution, coloration, behavior and audible
sound production of the spotted dolphin, Stenella plagiodon (Cope),
by David K. Caldwell and Melba C. Caldwell. 28 pp., 13 figs.
July 22, 1966.
Comparison of the early Permian vertebrate faunas of the Four
Corners region and north-central Texas, by Peter Paul Vaughn.
13 pp., 1 fig. July 22, 1966.
New distribution data for M artarega, Buenoa and Abedus, including
the first record of the genus Uartarega in the United States
(Hemiptera: Notonect idae, Belostomat idae ) , by Arnold S. Menke
and Fred S. Truxal . 6 pp. , 1 fig. July 22, 1966.
Two fossil birds from the lower Miocene of South Dakota, by
Hildegarde Howard. 8 pp., 1 fig. July 22, 1966.
Sounds and behavior of captive Amazon freshwater dolphins, Inia
geoffrensis, by Melba C. Caldwell, David K. Caldwell, and William
E. Evans. 28 pp., 10 figs. July 25, 1966.
A new Haliotid from Guadalupe Island, Mexico (Mollusca: Gastro-
poda), by Robert R. Talmadge. 4 pp., 2 figs. October 27, 1966.
Galeus piperatus , a new shark of the family Scyl iorhinidae from
the Gulf of California, by Stewart Springer and Mary H. Wagner.
9 pp., 2 figs. October 27, 1966.
A new subspecies of the Aztec mastiff bat, Molossus aztecus
Saussure, from southern Mexico, by Alfred L, Gardner. 5 pp. ,
Nov. 9, 1966.
The taxonomy and nomenclature of some North American bees of the
genus Centris with descriptions of new species (Hymenoptera :
Anthophoridae), by Roy R. Snelling. 33pp., 1 fig. Oct. 27, 1966.
A new pelobatine frog from the lower Miocene of South Dakota with
a discussion of the evolution of the Scaphiopus-Spea complex,
by Arnold G. Kluge. 26 pp., 8 figs. Dec. 28, 1966.
Additional avian records from the Miocene of Sharktooth Hill,
California, by Hildegarde Howard. 11 pp., 1 fig. Dec. 28, 1966.
Late Tertiary radiation of viperfishes (Chauliodontidae ) based on
a comparison of Recent and Miocene species, by Jules M. Crane, Jr.
29 pp., 13 figs. Dec. 28, 1966.
Recognition of the cancel lari id genus Neadmete Habe, 1961, in the
west American fauna, with description of a new species from the
Lomita Marl of Los Angeles County, California, by George P.
Kanakoff and James H. McLean. 6 pp., 2 figs. Dec. 28, 1966.
A new species of Architectonica from the Santa Susana Mountains,
Ventura County, California, by J. Alden Sutherland. 4 pp., 2 figs.
Dec. 28, 1966.
A new toe biter from Mexico (Belostomatidae, Hemiptera), by A. S.
Menke. 6 pp. , 5 figs. Dec, 28, 1966.
Additional fish remains, mostly otoliths, from a Pleistocene
deposit at Playa Del Rey. California, by John E. Fitch. 16 pp. ,
12 figs Dec. 31, 1966.
A new species of Dioptopsis from California (Diptera: Blepharo-
ceridae), by Charles L. Hogue. 5 pp. , 9 figs. Dec. 31, 1966.
Summer food of four species of lizards from the vicinity of White
Sands, New Mexico, by James R. Dixon and Philip A. Medica. 6 pp.,
4 figs, Dec. 31, 1966,
4
Contributions in Science
AUTHOR INDEX
Applegate, Shelton P.
Boolootian, Richard A.
Brown, David H.
Caldwel 1 , David K.
Cal dwel 1 , Melba C .
Crane , Jules M. , Jr .
Dixon, James R.
Evans, William E.
Fitch, John E.
Gardner, Alfred L.
Hogue, Charles L.
Howard, Hildegarde
Kanakoff, George P.
Kluge, Arnold G.
Lavenberg, Robert J.
Leighton, David
Macdonald, J. R.
McLean, James H.
Medica, Philip A.
Menke, Arnold S.
Siebenaler, J, B.
Snelling, Roy R.
Springer, Stewart
Sutherland, Alden
Talmadge, Robert R.
Truxal, Fred S.
Vaughn, Peter Paul
Wagner, Mary H.
Webb, Robert G.
White, John A.
No.
86
No.
93
No.
95
Nos .
91,95,104,108
Nos .
91,95,104,108
No.
115
Nos .
102,121
No.
108
No.
119
No.
111
Nos .
99,120
Nos .
94,101,107,114
Nos .
103,116
No.
113
No.
90
No.
93
Nos .
88,92
Nos .
100,116
No.
121
Nos .
89,106,118
No.
91
Nos .
97,98,112
No.
110
No.
117
No.
109
No.
106
Nos .
87, 105
No.
110
No.
102
No.
96
e les CONTRIBUTIONS
Z fill & IN SCIENCE
Number 86 April 9, 1965,
TOOTH TERMINOLOGY AND VARIATION IN SHARKS WITH
SPECIAL REFERENCE TO THE SAND SHARK, C ARCH ARIAS
TAURUS RAFINESQUE.
By Shelton P. Applegate
Los Angeles County Museum
Exposition Park • Los Angeles, Calif. 90007
CONTRIBUTIONS IN SCIENCE is a series of miscellaneous tech-
nical papers in the fields of Biology, Geology and Anthropology, published
at irregular intervals by the Los Angeles County Museum. Issues are num-
bered separately, and numbers run consecutively regardless of subject mat-
ter. Number 1 was issued January 23, 1957. The series is available to scien-
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MANUSCRIPT FORM.— (1) The 1960 AIBS Style Manual for Biological
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PROOF.— Authors will be sent galley proof which should be corrected and re-
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David K. Caldwell
Editor
TOOTH TERMINOLOGY AND VARIATION IN SHARKS WITH
SPECIAL REFERENCE TO THE SAND SHARK, CARCHARIAS
TAURUS RAFINESQUE.
By Shelton P. Applegate1
Abstract: Heterodonty in teeth is common in sharks. The
use of three new terms is advocated: alternates, medials and
posteriors. Dental formulae can be used in classifying recent and
fossil sharks. In Carcharias taurus Rafinesque, tooth length was
found to be proportional to total length of the shark. Unasso-
ciated fossil teeth may be identified through the erection of arti-
ficial tooth sets if the teeth can reasonably be referred to a single
species.
Introduction
In current studies of recent sharks the implications of the tooth termi-
nology and dental formulae which were proposed and used by Maurice Leriche
(1905, 1910, 1926) have not yet received the attention and use which they
warrant. This is due to a need to demonstrate that these tooth types and dental
formulae are truly significant characters that can, in fact, be useful in classi-
fying species and higher taxa. Once the approach pioneered by Leriche has
been validated, as attempted in this paper, it should be possible to project the
results obtained from studying fossil and recent shark dentition into higher and
higher taxonomic categories. This will lead, hopefully, to a better understanding
of generic and familial characters and to a better comprehension of selachian
evolution.
The aims of this paper are to give a general discussion of the use of tooth
types and dental formulae in sharks and to attempt to analyze the range of
variation in the teeth of the sand shark Carcharias taurus Rafinesque. If tooth
morphology and number varied widely between individuals of the same species
then the use of tooth characters to delineate species of fossil carchariids and
related sharks would be unwise and the whole rationale behind the use of tooth
types and dental formulae would be weakened. However, the variation within
the sample of Carcharias taurus is not of the magnitude that would cast doubt
on the validity of using tooth types or formulae in this shark. I therefore
contend that it is possible to distinguish species of fossil carchariids by their
teeth alone once the proper tooth type and probable position in the jaws of the
fossil teeth has been determined.
iAssociate Curator of Vertebrate Paleontology, Los Angeles County Museum.
4
Contributions in Science
No. 86
Acknowledgments
Thanks are due Dr. Lionel Walford, director of the Sandy Hook Marine
Laboratory at Sandy Hook, New Jersey, for permission to use the facilities of
the Laboratory. Mr. John Casey, head of the Laboratory’s Shark Project, was
particularly helpful in making contacts with the local fishermen and in the
capture of the four of the largest sand sharks. Reade Wood served as my field
assistant and aided greatly in the dissection of specimens. Dr. Bobb Schaeffer,
Department of Vertebrate Paleontology, American Museum of Natural
History, has aided in the preparation and editing of this paper, as has my wife,
Anne Chase Applegate. The photographs by Raymond Rigsbee were made
with the help of a grant from the Duke Faculty Research Fund. Dr. Joseph
Waters has assisted in the analysis of the statistics. The graph used in this paper
was prepared by Dorothy Kresch. The whole study, including publication, was
made possible through the aid of a grant from the National Science Foundation,
G-24538, An Investigation into the Interrelationships of Selected Modern
Shark Families.
Discussion of Heterodonty
The basis for tooth terminology and dental formulae in sharks rests on the
widespread occurrence of heterodonty among fossil and recent sharks. Hetero-
donty, although normally not applied to sharks’ teeth, is the logical term used
to express the radical change in size and shape of the teeth found in a shark’s
jaw. The normal reduction in size alone from the front to the rear of the mouth
does not indicate heterodonty.
A heterodont condition as is here defined is well documented for the
older Paleozoic and early Mesozoic sharks, particularly the hybodonts (Wood-
ward, 1891). On the other hand, the older cladodonts and their relatives need
a more detailed investigation before any similar generalization can be made.
One major factor in studying the occurrence of heterodonty in Paleozoic
sharks is that many of the species are described from a single tooth, a fact that
can be demonstrated by an examination of Woodward, 1891. Such a state of
affairs would tend to support an early appearance of homodonty whether it
ever existed in these sharks or not.
In recent and fossil sharks true homodonty, i.e., where the teeth in a jaw
are all the same shape and show no abrupt change in size, is a rare phenomena.
It may exist in recent Rhincodon and Cetorhinus. There is some evidence for
this condition in the Orectolobidae and to a lesser extent in the Scyliorhinidae.
Chlamydoselachus, the primitive fringed-gilled shark, has what could be
considered a homodont condition except that it has a single row of medial
teeth which are unique. If a true homodont tooth condition ever preceded a
heterodont condition it has yet to be demonstrated in the fossil record.
Heterodonty in sharks involves a number of distinct variations. A primary
type of heterodonty occurs when the upper teeth are quite different from the
lower teeth. In the family Pseudotriakidae and in some Scyliorhinidae not only
1965
Shark Tooth Terminology and Variation
5
Figure 1. Jaw of Carcharias taurus, field number 7, LACM number F 105, total
length 205 cm, from Sandy Hook Bay, New Jersey. A = anteriors, I = intermediate,
L=laterals, P=posteriors, S=symphyseal. Approximately half natural size.
6
Contributions in Science
No. 86
is the tooth shape distinct in different positions but the teeth of the lower jaw
are arranged in a completely different manner ( see Bigelow and Schroeder,
1948 : fig. 40) . In the Hexanchidae the lower teeth are long, flat and blade-like,
while the upper teeth are needle-like. A similar and possibly related condition
exists in the majority of species now placed in the families Squalidae and
Dalatiidae. The reverse condition exists when the upper teeth are blade-like
and the lower teeth are more narrowly pointed. This condition is well displayed
in the majority of the Carcharhinidae and in all of the Sphyrnidae. The latter
type of heterodonty has been developed independently in Carcharodon and
in some of the fossil species of Isurus — /. hastalis Agassiz for example. An even
more fundamental type of heterodonty occurs when individual teeth in an
upper or lower jaw vary widely in size and shape from their neighbors. This
common condition exists in the Heterodontidae, Hexanchidae, Carchariidae
(including the family Scapanorhynchidae) , Isuridae, Alopidae, Carcharhini-
dae and Sphyrnidae. Slight tooth differentiation occurs in Chlamydoselachidae
as has been discussed. Tooth differentiation is weakly defined in the Orectolo-
bidae, Scyliorhinidae, Squalidae, Dalatiidae, Echinorhinidae, Squatinidae, and
Pristophoridae.
Such regional differences in the shark jaws as has just been noted can be
best treated through the use of terms of a positional nature. A nomenclature of
this type already exists, for in 1905 Leriche coined such terms for the teeth
that occur in Carcharias ferox and later ( 1 905, 1 908, 1910, 1 926) , he extended
the use of these names to species belonging to other families, i.e., Isuridae,
Alopiidae, Hexanchidae, Squatinidae, Scyliorhinidae, and Carcharhinidae.
Leriche did not exhaust the applications of these terms nor their possible modi-
fications. There has been a wide use of Leriche’s names since 1905 by British
and French paleontologists; therefore any attempt to completely abandon
Leriche’s terms, no matter how technically desirable, would only confuse the
already lengthy literature. There is no reason why these names may not be
modified and new terms added if there is a real need for them. In time the
terminology of tooth types should become stable and will with common usage
give us a valuable tool in working with both recent and fossil sharks.
Leriche (1905) used the tooth names symphysaires anterieures, inter-
mediates, and laterales to describe the different teeth in Carcharias ferox
(Risso). An approximate English translation of these terms would be symphy-
seals, anteriors, intermediates and laterals. It is suggested that the term
posteriors be substituted for the posterior laterals as used by White in 1931.
In considering the teeth in the Scyliorhinidae, Triakidae, Pseudotriakidae,
Carcharhinidae, Sphyrnidae, Hexanchidae, Squalidae, Dalatiidae and Hetero-
dontidae it has become apparent that there is a need for another term to
designate the median teeth which occur in the symphyseal area and are distinct
from the symphyseal teeth as used in the Carchariidae. Obvious terms for these
teeth are median or medial teeth definable as small teeth of the symphyseal
1965
Shark Tooth Terminology and Variation
7
Figure 2. Symphyseal, anteriors, intermediates, and laterals of C archarias taurus
number 11, LACM F 106, adult female caught off of Lewes, Delaware. The teeth
are from the right side and are shown in an internal view. Approximately .8 natural
size.
8
Contributions in Science
No. 86
Figure 3. LACM F 106. A=first upper anterior, B=first lower anterior, C=
second upper anterior, D = second lower anterior, E = third upper anterior, F =
third lower anterior. Approximately natural size.
1965
Shark Tooth Terminology and Variation
9
area with at least one tooth in a medial position and the other teeth, if present,
identical to this median tooth. If the teeth adjacent to the median tooth are
similar in size and shape they should be considered median teeth also. Median
teeth may be symmetrical, or asymmetrical as in the recent Prionace glauca,
and then be oriented to the left or right.
In many of the Carcharhinidae there are small teeth which occur in the
symphyseal area that are neither medial, nor symmetrically arranged as the
symphyseals. These small teeth occur in oblique tooth rows of 2, 3, 4, or 5
teeth. Since there is an alternation of these teeth in the last position, the term
alternate tooth is suggested to cover these teeth.
Symphyseal teeth are the small asymmetrical teeth which lie on either
side of the symphysis (Fig. 2) and never in the center of the jaw. In C ar-
charias taurus such teeth are limited to the lower jaw. The teeth which might
be called upper symphyseals are here interpreted as being first upper anterior
teeth as they resemble these teeth in both shape and size.
Anterior teeth are situated on both sides of the upper and lower jaws in
C archarias; these teeth differ widely from the symphyseals (Figs. 2 and 3). In
C. taurus, there are three upper rows of anteriors. The smallest anterior is
borne in the first or more medial upper row or file (the latter term was used
by Leriche) . The lower jaw also possesses three rows of anteriors on either side
of the symphysis; these teeth lie to the outside of the symphyseals. The largest
tooth in the jaws of C. taurus is the second lower anterior tooth. The total height
of the anteriors is approximately twice their greatest root width. The lateral
edges of the tooth crown of the anteriors are nearly parallel for a short distance
before narrowing to an attenuated point. When these teeth are viewed from
the side there is a pronounced S-shaped curve of the crown. The total anterior
tooth even when it is in the first position in a file or row inclines inward into
the mouth. The two roots of the anteriors form an acute angle which is greater
than that of the symphyseals, but less than in the lateral teeth. In the anteriors
the largest root points toward the symphysis; in the symphyseals on the other
hand the larger root points away from the symphysis.
In the small immature specimens at hand the first upper anterior lacks
denticles, confirming the observation of Bigelow and Schroeder (1948: 99).
In the upper jaw the teeth termed intermediates occur just lateral to the
anteriors; there is only a single file of intermediates on each side of the upper
jaw. This tooth (Fig. 2) has a small triangular crown. The roots are asym-
metrical. The longest branch of the two roots points toward the symphysis.
There are apparently no denticles on this tooth in very young specimens, a
feature shared with the first upper anterior as discussed above.
The teeth designated as laterals occur to the rear and lateral to the inter-
mediates in the upper jaw and to the rear and lateral to the anteriors in the
lower jaw. Laterals are shown in Figure 2. Seen in side view the crown of
the laterals is straight; seen in anterior view the crowns are lower than those
10
Contributions in Science
No. 86
Figure 4. Posteriors of LACM F 106. Upper teeth, A=number 1, B=number 6,
C = number 11. Lower teeth, D = number 1, E = number 5, F = number 10.
One should note the great amount of chipping and wear in these teeth. Approxi-
mately 10 times natural size.
1965
Shark Tooth Terminology and Variation
11
of the anteriors. The roots of the laterals characteristically form obtuse angles
with each other. The total tooth height is almost equal to the greatest width.
The lower lateral teeth, when viewed anteriorly have a straight axis. The upper
laterals have crowns with a curved axis; this curve is directed toward the
corners of the jaw. Occasionally the lower posterior laterals may show a
marked posterior curving of the crown axis. There is a tendency in C. taurus
for the upper laterals to bear two denticles on each side of the tooth in contrast
to the usual one.
The small teeth in back of the laterals are called posteriors (Fig. 4) . They
are the most variable of the teeth in Carcharias. The crown may have a straight
or curved axis. The denticles may even rival the crown in size. The greatest
width of the roots is frequently more than the height of the tooth. There is no
marked extension of the root beyond the base of the crown as occurs in the
laterals.
In order to compare the tooth types mentioned in the sand shark (such as
alternates and medials) with those seen in other shark families, it may be said
that in 16 families of living sharks, excluding the Rhincodontidae and Cetor-
hinidae, medials are lacking in only four, the Carchariidae, Isuridae, Alo-
piidae and Echinorhinidae. Alternate teeth are known only from the Scylior-
hinidae, Triakidae, Carcharhinidae, Sphyrnidae and Echinorhinidae. Anteriors
are lacking in the Echinorhinidae. Intermediates are known only in the Car-
chariidae, Isuridae and Alopiidae. Laterals are lacking in Heterodontus for in
this species the anteriors are followed by flat posterior teeth. Posterior teeth
occur in all of the families.
Functional Relationships
The teeth in C. taurus serve to puncture, slice and crush the fish on which
this shark feeds. In feeding on its prey the shark must hold, immobilize (often
sever in half) and move its food to the stomach via the pharyngeal cavity
through the short esophagus.
The anteriors serve to puncture and kill or stun the prey; their strong
inward inclination makes it easy for food to be held and moved into the mouth.
A turn of the head or body would place the prey under the laterals where it
would be sliced and swallowed. The posteriors must serve to hold and crush the
food. The latter action is indicated by the great amount of wear that the
posteriors receive.
I believe that the symphyseals and intermediates function mainly to break
up the anterior teeth into patches; this would reduce the number of teeth
puncturing the prey, a factor making for both rapid and deep penetration of
these fangs as well as quick removal of the prey after it has been caught and
killed. Lack of serrations may be a factor in freeing prey from the teeth.
It is interesting to note that an examination of the articulation of the two
jaws in C. taurus leads to the conclusion that lateral movement of one jaw in
relation to another is all but impossible. Food must be moved by the move-
12
Contributions in Science
No. 86
ment of the whole jaw, or turning the head or the whole body in relation to
the food.
Most of the fishes taken from the stomachs of twelve sand sharks were
menhaden ( Brevoortia tyrannus, between 8 and 10 inches long). Each had
been chopped into two parts, the tails of each fish showing punctures that,
from their size and spacing, were made by the anteriors. The severing of the
fish must have been accomplished by the laterals as the cut in each case was
straight and clean.
Replacement and Number of Teeth
In Carcharias there is continuous replacement of teeth throughout the life
of the shark (Breder, 1942; Cadenat, 1962). As the teeth in the last position
fall out they are replaced by those behind; the last part of a tooth to calcify is
the tip of the root. Complete roots indicate that the tooth had reached the last
position; therefore, the paleontologist can be sure that he is examining mature
teeth by inspecting the tips of the roots. The tooth bud and the teeth in the
process of formation are hidden by a gum-like membrane through which the
teeth rupture. Once they have passed through this membrane, the teeth may be
considered to be functional. At any given time there are usually two functional
rows of anteriors, intermediates and laterals followed by from 4 to 6 rows of
small posteriors. The large number of posterior teeth no doubt add appreciably
to the total crushing area of these teeth.
In the present Carcharias sample the number of teeth in the upper jaw
varies from 38 to 55 and in the lower jaw from 34 to 44. This is a greater
variation in tooth number than has been reported in the past (Garman, 1913;
Bigelow and Schroeder, 1948). The use of total tooth number as a taxonomic
character in the Carchariidae therefore has little validity.
Dental Formula and Terminology
A more helpful tool is the variation in the number of teeth in the different
tooth types; this can be best revealed by the use of a dental formula similar
to that used by Leriche (1905, 1910, 1926) , Desbrosses (1930) and Dartvelle
and Casier (1943). Such a formula uses the first letter of each tooth type
followed by the number of teeth of this type in the first row (Fig. 1 ). If a tooth
is missing in the first position the one behind it is counted.
A horizontal line separates the teeth in the upper jaw from those of the
lower. As an example let us take the teeth of Carcharias taurus as figured by
Bigelow and Schroeder (1948: 95). This left portion of the jaw would be
written as follows:
A3 II F7 P16
SI A3 15 PI 3
The total numerical variation of the tooth types in the series of twelve
specimens of Carcharias taurus used in the present study may be expressed by
the following formula.
1965
Shark Tooth Terminology and Variation
13
P6-19 L6-7 II A3 A3 II L6-8 P8-15
P4-14 L5^6 A3 sT~" SI A3 L545 P8-13
From this formula it is evident that neither the symphyseals, intermediates and
or the anteriors varied in number. The variation of the laterals and posteriors
is shown in Table 1.
TABLE 1
The variation in the number of lateral and posterior teeth
in Carcharias taurus.
Laterals Posteriors
Upper Lower Upper Lower
Specimen
Right
Left
Right
Left
Right
Left
Right
Left
No.
No.
No.
No.
No.
No.
No.
No.
No.
1.
6
6
5
5
8
10
8
9
2.
6
6
5
6
10
10
9
9
3.
7
6
5
5
9
8
9
9
4.
6
6
5
5
10
10
9
8
5.
6
6
5
5
10
9
8
8
6.
6
6
5
5
19
15
14
11
7.
6
7
6
6
11
15
4
11
8.
6
6
5
5
12
12
12
12
9.
6
7
5
6
9
10
10
8
10.
7
7
5
6
8
10
10
9
11.
7
8
5
5
13
11
13
13
12.
7
7
5
6
9
10
12
12
The mode of the lower laterals is five per side and in the upper laterals it is
six per side. These modes occur in 79% and 66% of the sample respectively.
The upper posteriors vary from 8 to 19, which is a range of 1 1 teeth as opposed
to a range of 3 teeth for the laterals. The upper posterior teeth average 10.7 per
side; the mode per side is 10. The lower posteriors have an average of 9.8 per
side with a variation from 4 to 14 with a range of 10.
In some of the specimens of C. taurus the addition of posterior teeth
during the life of the shark may be demonstrated; several single teeth were
followed in the replacement series by a double row of immature teeth. The
deletion of teeth in a jaw is more difficult to demonstrate although a case of
deletion may be reflected in a specimen in the comparative anatomy collections
at Duke University in which no intermediates are present even though a space
for these teeth exists. Since this specimen lacks data one cannot be sure the
intermediates have not been removed; however close examination suggests that
this is unlikely.
In reconstructing the dentition of fossil sharks several terms were needed.
14
Contributions in Science
No. 86
First “tooth set”; that is, a single complete row of all types of mature teeth from
both sides of the upper and lower jaws, arranged in their natural order as they
occurred in life. A “natural tooth set” is one which shows the natural order of
tooth arrangement. Such a “natural tooth set” occurs obviously in living sharks;
in fossils it occurs only under exceptional conditions of preservation where the
teeth are still in place in the jaws. It is, of course, more likely that a partial
“natural tooth set” will be found with only a few teeth in place. An “associated
tooth set” occurs when a number of tooth types are found which can be re-
ferred to one specimen. An “artificial tooth set” may be erected when a num-
ber of tooth types from one locality may be considered to belong to one species.
In doing this, comparisons are made with known related natural sets as well as
associated sets. As with the “associated tooth set” tooth positions can only be
inferred. Occasionally material from more than one locality may be used in
“artificial sets” when the chance of confusion with a closely related species
is negligible.
Once we know what tooth types exist for a set of a fossil species then we
can assess more accurately the specific limits of these types and the likelihood
of calling different tooth types from the same set different species becomes
more remote. If carefully used and applied, the use of “sets” should allow more
exactitude in determination of species.
Body Length and Tooth Height
It is evident from an examination of Table 2, that there is a general
increase in total tooth height which coincides with a similar increase in the
total length of the shark. In this case total length is the measurement from the
tip of the snout to the tip of the tail, and total tooth height is that measurement
from the tip of the tooth to the tip of the largest root. These measurements of
tooth height and fish length were plotted against each other. A definite linear
relationship was found in all cases. An example of such a plot is shown in
Figure 5. The total height of the second lower anterior tooth was plotted against
the total length of 10 sharks having these teeth. It may be added that the
second anterior is the largest tooth in the jaws although the third anterior is
slightly larger in one jaw. The constancy of this linear relationship at once
suggests the possibility of being able to predict the total length of a shark once
the total height of a particular tooth of one of the tooth types is known. Once
his procedure has been established in recent sharks then it might be possible to
compute the total length of fossil sharks such as Isurus hastalis (Agassiz),
Hemipristis serra Agassiz and Carcharodon megalodon Agassiz.
One might conclude that the tooth bud or even earlier germinal layer
increases in size as the shark grows larger, so that at any one time the functional
tooth size is a reflection of the size of the fish.
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TOTAL LENGTH OF SHARK
16
Contributions in Science
No. 86
TABLE 2
Total length of ten sand sharks compared with total tooth height
LOWER JAW
Total tooth height in cm.
Total length
in cm.
right left
No.
PI
LI
A3
A2
A1
SI
SI
A1
A2
A3
LI
PI
1.
112
.27
.71
1.23
1.49
1.46
.65
.79
1.46
1.53
.98
.71
.30
2.
113
.26
.86
1.27
1.55
1.61
.79
.78
1.52
1.65
1.25
.82
.23
4.
118
.35
.99
1.27
1.77
1.51
.79
.75
1.52
1.66
1.25
.88
.35
6.
148
.41
1.21
1.50
2.27
2.18
1.13
.98
2.22
2.14
1.77
1.35
.41
7.
205
.35
1.38
1.73
2.38
2.21
1.13
1.12
2.16
2.34
1.85
1.48
.62
8.
207
.45
1.52
2.36
2.90
2.78
1.28
1.25
2.73
2.85
1.52
.34
9.
227
.71
1.81
2.64
3.03
3.00
1.70
1.74
2.98
3.06
2.66
1.88
—
10.
252
.70
1.62
2.15
3.20
2.88
1.50
1.55
2.53
3.00
1.46
.85
11.
272
.60
2.27
2.69
3.54
3.50
1.78
1.64
3.35
3.27
2.95
2.01
.69
12.
273
.62
1.89
2.61
3.44
3.33
1.89
1.82
(2.69)
3.02 ■
2.00
.50
UPPER JAW
ri
ght
left
No.
PI
LI
A3
A2
A1
SI
SI
A1
A2
A3
LI
PI
1.
112
.30
.81
.33
1.07
1.35
1.02
1.03
1.35
1.19
.31
.66
.31
2.
113
.37
.85
.41
1.28
1.30
1.11
1.20
1.29
.40
.89
.38
4.
118
.37
1.09
.50
1.23
1.44
1.19
1.45
.51
.91
.38
6.
148
.42
1.12
.69
1.60
1.71
1.59
1.69
1.69
1.72
.73
1.21
.43
7.
205
.49
1.27
.75
1.83
2.04
1.70
1.70
2.00
1.77
.81
1.31
.52
8.
207
.69
1.42
.98
1.82
2.46
2.10
2.10
2.47
.89
1.29
.52
9.
227
1.80
1.07
2.32
2.99
2.23
2.93
.92
1.69
.65
10.
252
.79
1.70
.83
2.55
2.72
2.31
2.28
2.55
2.50
.84
1.80
.66
11.
272
.65
2.02
1.15
2.64
2.93
2.54
2.99
2.43
1.11
1.95
.88
12.
273
1.06
2.22
1.17
2.75
2.60
2.65
3.11
2.70
1.06
2.06
1.02
The total tooth height (greatest distance from tip of root to tip of tooth) is
measured in centimeters. It should be noted that sharks number 3 and 5 with total
lengths of 113 and 140 centimeters were not used in this chart for they were broken.
The capital letters at the top of each column stand for the respective tooth types as
described in the text.
1965
Shark Tooth Terminology and Variation
17
Conclusions
Leriche’s terminology and dental formulae give the student of fossil and
recent sharks a meaningful method of describing and studying sharks’ teeth.
Heterodont dentition as defined in this paper is a common phenomena in
recent and fossil sharks.
To Leriche’s tooth terms should be added medials, alternates and
posteriors as distinctive tooth types.
The restrictive nature of tooth types above a familial level suggests a
natural grouping of sharks which may be phyletic.
The functional use of teeth in Carcharias taurus can be correlated with
tooth type.
Completely formed root tips indicate mature teeth.
In Carcharias taurus, total tooth number for the upper and lower jaw is
not a reliable specific character; however the number of the symphyseals,
anteriors and intermediates is constant. Numerical variation is confined to the
laterals and posteriors.
In C. taurus tooth length is directly proportional to total length.
The use of artificial tooth sets is an effective way to treat unassociated
fossil teeth.
The results of this study of tooth variation in C. taurus suggest that there
is much information to be gained by extending such studies to other recent
species of sharks. Essential to such studies are collections of jaws accurately
identified with reliable locality, sex, and size data. For each species as many
jaws as is practical should be examined.
18
Contributions in Science
No. 86
Literature Cited
Bigelow, H. B., and Schroeder, W. C.
1948. Sharks, In Fishes of the Western North Atlantic, Part I, Mem. Sears
Found, for Marine Research, pp. 59-576.
Breder, C. M.
1942. The shedding of teeth by Carcharias littoralis (Mitchell). Copeia,
1942(1): 42-44.
Cadenat, J.
1962. Notes d’ Ichtyologie ouest-africaine. XXXVIII. — Documents pour ser-
vir a la recherche des mecanismes de deplacement et de remplacement
des dents chez les Requins, Bulletin de L’Instut. Francais d’Afrique
Noire. Tome XXIV, Serie A. 2:551-579, 26 pis.
Dartvelle, E., and Casier, E.
1943. Les Poissons fossiles du Bas-Congo et des regions voisine, Tervueren (A
Annales, Musee Congo Belgique), Parts I and II, pp. 1-255, 76 figs., pis.
I-XXII.
Desbrosses, P.
1930. Presence du Squale feroce: Odontaspis jerox Agassiz dans le Golfe de
Gascogne, Bulletin Societe Zoolique, France, T. 55, pp. 232-235, 5 figs.
Garman, S.
1913. The Plagiostomia (sharks, skates and rays). Mem. Mus. Comp. Zook,
Harvard, 36: 1-528.
Leriche, M.
1905. Les Poissons Eocene de la Belgique, Memoires du Musee Royal His-
toire Naturelle Belgique Bruxelles, T. Ill, pp. 49-228, pis. IV-XII, text
figs. 9-64, 1 series, Mem. No. 11, 1905.
1906. Contributions a l’etude des Poissons fossils du Nord de le France,
Memoires de la Societe Geologique du Nord. T. V, 1906, pp. 1-430, pis.
I-XVII, 78 text figs.
1908. Note sur des Poissons paleocenes et eocenes des environs de Reims
(Marne), Annales de la Societe Gelogique du Nord, T. XXXVII, 1908,
pp. 230-232, fig. 1.
1910. Les Poissons Oligocenes de la Belgique, Memoires Musee Royal His-
toire Naturelle Belgique Bruxelles, T. V, pp. 231-363, figs. 65-159, pis.
XIII-XXVII.
1926. Les Poissons neogene de la Belgique, Memoires Musee Royal Histoire
Naturelle Belgique Bruxelles, Memoire 32, 1926, pp. 368-472, pis.
XXVIII XLI, text figs. 161-228.
1938. Contribution a l’etude des Poissons fossiles des pays riverains de la
Mediterranee americaine (Venezuela, Trinite, Antilles, Mexique) Mem-
oires de la Societe Paleontologique Suisse, vol. LXI, pp. 1-41, pis. I-IV,
8 figs.
White, E. I.
1931. The Vertebrate Faunas of the English Eocene. London. British Museum
(Natural History), XIV -f- 1-123 p., 34 figs., 16 pis.
Woodward, A. S.
1891. Catalogue of the fossil fishes in the British Museum, London, 1:1-474,
figs. 1-12, pis. I-XVII.
■ \ .
I ( f
LOS
ANGELES
COUNTY
MUSEUM
Number 87
CONTRIBUTION*
IN SCIENCE
June 28, 196
FROG-LIKE VERTEBRAE FROM THE LOWER
PERMIAN OF SOUTHEASTERN UTAH
By Peter Paul Vaughn
Los Angeles County Museum • Exposition Park
Los Angeles, Calif. 90007
FROG-LIKE VERTEBRAE FROM THE LOWER PERMIAN
OF SOUTHEASTERN UTAH1
By Peter Paul Vaughn2
Abstract: A new genus of Permian amphibians, Lasalia,
is based on the new species L. cutlerensis. The holotype of the
species is a remarkably frog-like vertebra found in the undif-
ferentiated Cutler sediments of Lisbon Valley, south of the town
of La Sal in southeastern Utah; another vertebra from the same
locality is referred to the species. The associated fauna indicates
a Lower Permian ( Wolfcampian ) horizon. The vertebrae are
amphicoelus, but they are not notochordal— this is noteworthy
for an early Permian amphibian. In the holotypic vertebra there
are long, hollow transverse processes that arise from the pedicels
of the neural arch; there is an indication that the ribs also had
capitular attachments. Lasalia is compared with other Paleozoic
vertebrates including the lepospondyls, with the Triassic Proto-
batrachus, with the Jurassic N otobatmchus, and with the living
amphibians. Close resemblances are found only among the
salientians. The ways in which Lasalia differs from the known
salientians— costal capitular attachment, relatively larger cen-
trum, other, minor points— are such as could easily have been
mitigated in the interval of time under consideration. It is not
claimed, but only suggested, that Lasalia is related to the
Salientia. If it is so related, a re-evaluation of the antiquity of
the vertebral structure of those living frogs generally considered
to be the most primitive may be required. Knowledge of
Lasalia does not offer much assistance in the search for the
ancestors of the Salientia. It is pointed out that the amphibians
were a highly diverse group even as early as the Permian.
INTRODUCTION
In the course of a long-range study of the vertebrates of the Permian
Cutler Group of the Four Corners region, several large pieces of rock that
contain many scattered bones were taken from a conglomerate in the undif-
ferentiated Cutler sediments of Lisbon Valley, about twelve miles south-
southeast of the town of La Sal, San Juan County, Utah. The Cutler beds in
this area are exposed along the southwestern flank of a salt anticline (see
Elston, Shoemaker and Landis, 1962). Random splitting and acid-etching of
the blocks of matrix in the laboratory disclosed many more bones; among
them are two small vertebrae quite unlike any others described from the
Permian. Further preparation and study of these vertebrae have revealed
their remarkably frog-like nature. Although it is not possible at this time
definitely to identify any other bones from the same site as pertaining to
the same species, it would seem to be more than worthwhile to describe and
discuss these vertebrae now, especially in view of the fact that the only pre-
vious notice of any pre-Triassic creature in any way really like frogs is the
description, by Griffiths (1963), of a trackway from the Lower Permian
Ecca beds of South Africa,
This study was supported by National Science Foundation grant NSF GB-1014.
2Research Associate, Los Angeles County Museum; and Department of Zoology,
University of California, Los Angeles.
1965
Lower Permian Frog-like Vertebrae
3
A NEW GENUS AND SPECIES OF AMPHIBIANS
Because of the evident importance of this heretofore unknown kind of
Permian vertebra, it seems desirable to establish a new species, for which a
new genus must be set up. The two vertebrae are so obviously unique among
Permian specimens that it seems unlikely that there would be any difficulty
in the correct referral of future finds to the species. It seems that a new
family should be established for the reception of this animal, but a formal
diagnosis of this family is best postponed until such time as its fossil record
becomes better known.
Class AMPHIBIA
Subclass and order uncertain
LASALIDAE, new family
This family is based on the new genus Lasalia, described below.
Lasaliq, new genus
Type species —Lasalia cutlerensis , new species.
Diagnosis— (The diagnosis pertains to vertebrae presumably in the an-
terior part of the column; a description of possible serial changes within
the column may be found in succeeding sections.) The length of the centrum
of the vertebra is not much greater than the width of the centrum. The
centrum is amphicoelus but not notochordal; its center is composed of can-
cellous bone. The surface of the centrum is pitted by small, irregularly
spaced, longitudinally oriented foramina. A gently rounded ridge on the
anterior half of the lateral surface of the centrum fades into the general
surface of the centrum posteriorly; the anterior part of this ridge is carried
away laterally from the general surface of the centrum by a fairly promi-
nent buttress; a round, shallow pit occurs on the lateral surface of this ridge
near its anterior end. There is no neurocentral suture, at least not in the
adult. The neural canal is elliptical in cross-section, wider than high. The
neural arch and spine are of moderate height; the spine is prolonged pos-
teriorly as a median process lying between the postzygapophyses; this proc-
ess and the postzygapophyses terminate posteriorly on about the same trans-
verse plane. The neural spine also projects forward slightly beyond its base.
Long, hollow transverse processes, circular in cross-section and of somewhat
greater diameter distally than proximally, arise from the bases of the pedi-
cels of the neural arch and extend laterally and somewhat posteriorly. The
base of the transverse process is separated from the buttress on the side of
the centrum by a short groove that faces anteriorly and somewhat laterally.
4
Contributions in Science
No. 87
Lasalia cutlerensis , new species
Figure 1
Holotype — UCLA VP 1670, a single vertebra.3 This vertebra is nearly
complete, but the prezygapophyses and a portion of the anterior part of the
centrum are lacking due to the initial exposure of this specimen by means
of a random cut through the matrix with a diamond saw. The vertebra
is very fragile and, after the photograph (Fig. 1) was taken, further prepa-
ration broke the vertebra in such a way that the distal half of the left
transverse process and parts of the left postzygapophysis and posterior por-
tion of the neural spine lie in one piece of matrix while the rest of the verte-
bra lies in another piece. In addition, the anteroventral lip of the centrum is
separated from the other parts— due to the initial cut with the saw.
Horizon and locality— The holotype, the referred specimen, and asso-
ciated specimens were collected on July 25, 1964, by a field party under
my supervision. The locality is in Lisbon Valley, in SE14 SE1/* SEx/4 sec. 20,
T. 30 S., R. 25 E., San Juan County, Utah. The site of collection is in the
Cutler sediments, about 200 feet above their contact with the underlying
Hermosa Formation as these units are bounded in the U. S. Geological Sur-
vey map of the Mount Peale 4 NW Quadrangle (see Weir, Puffett and
Dodson, 1961). The specimens were collected from a dull red conglomerate
about 25 cm. thick composed of very fine-grained sandstone pebbles, mostly
in the size range from 0.5 to 2 cm., embedded in a matrix of medium-
sized quartz grains covered with iron oxide, the whole held together by a
calcareous cement; there are also scattered flakes of mica. The bones and
fragments of bones occur among the quartz grains, between the pebbles.
The total aspect of the conglomerate suggests a stream-channel deposit. The
excellent preservation of such objects as a thin maxillary bone of a small
pelycosaur, without any signs of wear, indicates the primary nature of the
deposit. The associated fauna, which will be described below, indicates a
Lower Permian (Wolfcampian) horizon, probably equivalent to the Hal-
gaito Shale of the differentiated part of the Cutler Group in the vicinity of
Monument Valley (see Baars, 1962; Vaughn, 1962).
Referred specimen— UCLA VP 1673, a vertebra broken obliquely
through the centrum and neural arch. This vertebra comes from the same
locality as the holotype, and although it differs from the holotype in minor
ways to be noted below, its general similarity and especially its lack of a
continuous notochordal canal justify reference to Lasalia cutlerensis.
3The abbreviation “UCLA VP” stands for University of California, Los Angeles,
Vertebrate Paleontology.
1965
Lower Permian Frog-like Vertebrae
5
Figure 1. The vertebra UCLA VP 1670, holotype of Lasalia cutlerensis, new genus,
new species. The vertebra is seen in anterior view at a magnification of approxi-
mately seven times. Due to the initial exposure of the vertebra by means of a cut
with a diamond saw, the anterior end of the centrum is not seen, but the cancellous
bone of the center of the centrum is clearly visible. The right transverse process
is bent backwards, and details of its structure cannot be as easily made out as can
those of the left process.
Diagnosis— At this time the specific is the same as the generic diag-
nosis. Further characterization of the species may be gained from the meas-
urements and more detailed description given in the following section.
Description— Some further description of the holotype, beyond the
general features from which the generic diagnosis was drawn, is in order
here. Due to damage in exposure of the specimen, the length of the centrum
cannot be stated with great accuracy, but it must have been about 4 to 5
mm. The greatest width across the centrum, in a line that takes in the but-
tresses of the anterior ends of the lateral ridges, is about 4.3 mm. Posterior
to the ridges, the width is only about 3 mm. The ventral surface of the
centrum is flattened in its central area; this flattened part forms an obtuse
angle with the lateral surface of the centrum on either side. The antero-
ventral lip of the centrum that was separated from the rest of the vertebra
by the saw cut is concave above; this demonstrates the concave nature of
the anterior end of the centrum. The concavity of the posterior end of the
centrum was revealed by grinding with a small abrasive tool. In the can-
cellous bone of the center of the centrum, there is no continuous canal that
would indicate that the notochord was continuous. The width of the neural
canal is 1.8 mm. The median posteriorwards projection of the neural spine
acts as a prolonged roof for the neural canal; this projection is separated
from the postzygapophyses by rounded notches. The postzygapophysis of
at least the left side (details on the right side are not visible) seems to
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have had a concave hind border. The left transverse process (again, details
on the right side are obscure) is 3.6 mm. long. The diameter of the distal
end of the process is about 1.5 mm.; near its proximal end, the process is
somewhat flattened and has a height of only about 1 mm. The circular
opening at the distal end of the process faces mostly laterally but also
slightly ventrally. On at least the left transverse process there is a ridge
along the anterior border. I suspect that the small, round pit near the
anterior end of the lateral surface of the ridge that runs along the side
of the centrum served as the facet for the capitulum of the rib. If there was
a bony costal capitulum in L. cutler ensis, the rib must have been unusual
in appearance, with its tuberculum widely separated from the capitulum.
Possibly the capitulum was represented only by cartilage or by a liga-
mentous attachment to the facet on the centrum. In any event, if the facet
does represent a capitular attachment, this would indicate that there was
no intercentrum. The groove that separates the base of the transverse proc-
ess from the lateral ridge on the centrum probably carried the spinal nerve.
The referred vertebra, UCLA VP 1673, is broken obliquely through
the centrum and neural arch, but details of the center of the centrum, the
neural canal, the transverse processes, and the dorsal surfaces of the post-
zygapophyses are clear. As in the holotype, there is no continuous noto-
chordal canal; the center of the centrum is composed of cancellous bone. It
is also clear that the front of the centrum is concave, but the difficulty in
preparation of this fragile specimen makes it impossible to describe the
nature of the posterior end of the centrum. The length of the centrum
cannot be stated, but its width seems to be the same as the width behind
the lateral ridges in the centrum of the holotype, about 3 mm. Details are
not clear, but there seems to be no sign of ridges along the sides of the
centrum— this would be a difference from the holotype. The transverse
processes come off the vertebra at a sharper angle than in the holotype;
they are directed posteriorly at an angle of about 45 degrees. The transverse
processes are shorter than in the holotype, only about 2.3 mm. long, and
they are only 1 mm. in diameter at their distal ends. As in the holotype,
the transverse processes arise from the bases of the pedicels of the neural
arch, are hollow, are somewhat flattened dorsoventrally at their proximal
ends and circular in cross-section in their distal parts, and each bears a
ridge along its anterior border. A notable difference from the holotype is
that in the referred specimen the posterior borders of the postzygapophyses
meet in a centrally placed, rounded notch; there is no posteriorwards pro-
longation of the neural spine between the postzygapophyses. The posterior
borders of the postzygapophyses are perhaps slightly concave, but not as
markedly as was observed in one of the postzygapophyses of the holotype.
The lesser length and diameter of the transverse processes make it seem
that the referred specimen is from a position in the vertebral column pos-
terior to the serial position of the holotype. If there really were no lateral
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Lower Permian Frog-like Vertebrae
7
ridges on the centrum of the referred specimen, this would indicate that
the more posterior ribs lacked capitula. Unfortunately, there is no way of
knowing whether or not the two vertebrae are from the same individual
animal; they were not close to one another in the matrix.
There are other small bones in the collection from the same rocks—
finely sculptured bits of dermal bone, tiny, simply shaped limb elements,
and so on— that may possibly pertain to Lasalia cutlerensis, but this is
unsure. It may be hoped that future field work will uncover an articulated
specimen. This seems to be unlikely in the conglomerate itself, but it may
be possible to find such a specimen in sandstones associated with the con-
glomerate, and I certainly intend to continue the search.
ASSOCIATED FAUNA
From the same rocks from which the holo type and the referred specimen
of Lasalia cutlerensis were taken were also collected: teeth of pleuracanth
sharks, parts of palaeoniscoid fishes, crossopterygian scales, a vertebra of
the lepospondylous amphibian Diplocaulus sp., a nasal bone referable to
the rhachitomous labyrinthodont amphibian Eryops sp., a neural spine of
an edaphosaurian pelycosaur, and bones of another kind of pelycosaurian
reptile, probably an eothyridid. Besides these elements there are, of course,
many unidentified bones and parts of bones.
The pleuracanth teeth are like those of Xenacanthus texensis Cope from
the Wichita Group of northcentral Texas, quite unlike those of X. platyp-
ternus (Cope) from the higher Clear Fork Group of Texas (see Hotton,
1952). Teeth of Xenacanthus aff. X. texensis have also been reported from
the Halgaito Shale of the Cutler Group in the region of Monument Valley
(Vaughn, 1962). The palaeoniscoid scales are of several different kinds;
some resemble the scales of Progyrolepis tricessimilaris Dunkle from the
Wichita Group of Texas; similar scales have been reported from the Hal-
gaito Shale (Vaughn, 1962). Many small, conical teeth with distinct
enamel caps have also been recovered from the conglomerate; presumably
these also pertain to palaeoniscoids. The crossopterygian scales are like
those of Ectosteorhachis nitidus Cope from the Wichita Group of Texas;
E. nitidus is not found in the overlying Clear Fork Group. Ectosteorhachis
aff. E. nitidus has been reported from the Halgaito Shale (Vaughn, 1962).
Neither pleuracanth nor crossopterygian remains have been found in the
higher Organ Rock Shale of the Cutler Group, and of the few palaeonis-
coid scales known from the Organ Rock, none resemble those in Progyro-
lepis (Vaughn, 1964).
The vertebra referable to the lepospondylous amphibian Diplocaulus
sp. (UCLA VP 1674) consists of the complete centrum 3.5 mm. long and a
part of the neural arch. Diplocaulus is known from rocks as old as late
Pennsylvanian and is found in both the Wichita and Clear Fork Groups of
Texas, but this is the first report of Diplocaulus from the region of the
Four Corners.
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The nasal bone referable to Eryops sp., UCLA VP 1668, fits exactly
the description of the nasal bone of E. megacephalus Cope given by Sawin
(1941). The nasal bone from Lisbon Valley is only about three-quarters as
large as that in the specimen of E. megacephalus figured by Sawin; Sawin’s
specimen is from the upper part of the Wichita Group of Texas. Small
vertebral parts referred to Eryops sp. have been reported from the Halgaito
Shale (Vaughn, 1962).
UCLA VP 1669 is a small pelycosaurian maxillary bone, of about 50
mm. length, with a dentition very much like that of the eothyridid Bald-
winonus trux Romer and Price. B. trux is known from El Cobre Canyon
in nothern New Mexico, from beds that are probably of Wolfcampian
horizon (see Vaughn, 1963a), but in view of the fact that the only known
specimen of B. trux has a maxillary bone of about three times the length of
UCLA VP 1669, it would seem unwise to use this comparison for estimation
of horizon. The collection from Lisbon Valley also includes a fragment of
a scapulocoracoid that has the general aspect of this element as it is known
in ophiacodont pelycosaurs— to which group the eothyridids apparently
belong— and other pelycosaurian elements in the collection may pertain to
the Lisbon Valley Peothyridid too. Because there seems to be a good chance
that more parts of this small pelycosaur will be found in future field work,
a description of the maxillary bone is postponed.
The edaphosaurian neural spine, UCLA VP 1667, is quite similar in
structure and size to those of the small Edaphosaurus novomexicanus Willis-
ton and Case known from the Cutler beds of northern New Mexico. These
New Mexican beds are generally considered to be equivalent to the lower
parts of the Wichita beds of Texas (Langston, 1953; Romer, 1960; Vaughn,
1963a).
In sum, the evidence points to equivalence of the type locality of
Lasalia cutlerensis to some horizon in the lower part of the Wichita Group
of Texas, and also to probable equivalence to the Halgaito Shale of the
Cutler Group in the vicinity of Monument Valley. The lower part of the
Wichita Group is of Wolfcampian (Lower Permian) horizon (see Dunbar,
et al., 1960), and the same is probably true of the Halgaito Shale (Vaughn,
1962, 1964). Unless contradictory evidence presents itself, it would seem
that Lasalia cutlerensis lived in early Permian times.
Mention may be made of another fossiliferous level in the Cutler sedi-
ments of the same general area of Lisbon Valley. From a cross-bedded sand-
stone in SW*/4 NEV4 NW14 sec. 34, T. 30 S., R. 25 E., my field party recov-
ered, among other items, a maxillary and other bones, UCLA VP 1671, of a
fairly large sphenacodontid pelycosaur and a complete tibia, UCLA VP
1672, apparently also of a sphenacodontid. This locality is about 1200 feet
above the Hermosa Formation. Preliminary study of these materials shows
similarity to specimens from the Organ Rock Shale of the Cutler Group in
Monument Valley. In Monument Valley, the divisions of the Cutler Group
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Lower Permian Frog-like Vertebrae
9
are, in ascending order, Halgaito Shale, Cedar Mesa Sandstone, Organ Rock
Shale, and De Chelly Sandstone (see Baars, 1962). If one calculates the aver-
age thicknesses of these formations from the data given by Baker (1936),
then it may be seen that the level of the type locality of Lasalia cutlerensis
falls easily within the boundaries of the Halgaito Shale and the level of the
large sphenacodontid maxillary bone falls easily within the boundaries of
the Organ Rock Shale. This is, admittedly, a naive approach to stratigraphy,
but, taken along with the faunal evidence to date, it does suggest that it
may soon be possible to correlate beds of the undifferentiated Cutler sedi-
ments of Lisbon Valley with those of the differentiated Cutler Group of
Monument Valley.
COMPARISON WITH OTHER FORMS
The vertebrae of Lasalia cutlerensis must be compared, first of all,
with vertebrae of other Paleozoic forms. Although completely ossified,
imperforate centra are not unknown among Paleozoic fishes— they occur
in an Upper Devonian lungfish (Jarvik, 1955)— I know of no Paleozoic
fish whose vertebrae even remotely resemble those of L. cutlerensis. The
absence of a notochordal canal and the unitary construction of the centrum
would seem to rule out any close connection with the labyrinthodont am-
phibians of the Paleozoic. The lack of a notochordal canal also makes it
seem highly improbable that we are dealing with any sort of reptile; as
Romer (1956:223) has pointed out, “a notochordal type of centrum was
. . . present in all cotylosaurs and was unquestionably characteristic of ances-
tral reptiles . . . .” The above comparisons are drawn on basic structure; nor
is there any resemblance of the vertebrae of L. cutlerensis to any of these
groups in details of structure.
Next to be considered are those late Paleozoic amphibians usually
banded together as the Lepospondyli; these are the Aistopoda, Nectridia,
Lysorophia, and Microsauria. It may be noted that various authors have
argued for reptilian relationships for the Microsauria; the literature is too
complicated to go into here, and the matter is not really germane to the
present discussion. Lasalia cutlerensis is similar to the lepospondyls in the
unitary construction of the centrum. Williams (1959) has given reason to
believe that the lepospondylous centrum, like the centrum in living gym-
nophionans, urodeles and anurans, is intersegmental in position and there-
fore equivalent to the pleurocentrum— the dominant central element in
amniotes. If I am correct in my interpretation of the pit on the lateral
ridge of the centrum in L. cutlerensis as a capitular facet, then it seems quite
likely that the centrum in L. cutlerensis was also intersegmental, and this
would heighten the similarity to the lepospondyls. With this, the similarity
ends. As far as I know, the centrum is notochordal in all lepospondyls,
and an examination of details of structure reveals further differences. In the
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aistopods and nectridians, the transverse process is given off near the center
of the centrum, rather than from the pedicel of the neural arch as in
Lasalia. Further, in the aistopods the transverse processes are relatively short;
and although nectridian processes may be long, there is usually a doubling
of the process so that distally there are two articular facets, one above the
other— much as in urodeles. Nectridians are also distinguished by the great
anteroposterior elongation of their neural spines. In the lysorophians the
neural arch occurs, even in adults, in two lateral halves, and although
the transverse processes arise from the pedicels of the neural arch, they are
much shorter than those in Lasalia. Elongate transverse processes of the kind
seen in L. cutlerensis are not known among the microsaurs; indeed, micro-
saurian vertebrae often look much like those of captorhinomorph cotylosaurs.
The neurocentral suture in the vertebrae of microsaurs and lysorophians
has also been pointed out, from time to time, as a feature that distinguishes
these animals from other lepospondyls and the living amphibians, but this
seems to be unreliable; Gregory, Peabody and Price (1956) found the value
of this feature to be overridden by other structural relationships, and Wil-
liams (1959) has noted that neurocentral sutures normally occur even in
young Liopelma hochstetteri, a living frog.
Parsons and Williams (1962, 1963) have presented an impressive list of
characters held in common by the orders of living amphibians— the Gym-
nophiona, Urodela, and Anura. Chief among these in terms of potential
usefulness to the paleontologist is the common possession of pedicellate
teeth, teeth in which two distinct portions, a pedicel and a crown, are both
truly parts of the tooth and are both composed primarily of dentine. These
authors use these resemblances to show that all the modern amphibians are
closely related, in contradistinction to various theories of polyphyletic ori-
gins discussed in their papers, and they revive Gadow’s name Lissamphibia
to include the three modern orders. After consideration of earlier theories
on the origins of the modern amphibians, and after a search for the
“protolissamphibian” among the various groups of Paleozoic vertebrates,
including the lepospondyls, Parsons and Williams conclude (1963:48) that
“it is impossible, on the basis of our present knowledge, to put forward
even a tentative theory on which group of Paleozoic amphibians are to be
considered ancestral to the Recent orders,”
There is no fossil record of gymnophionans. The earliest undoubted
record of urodeles is in the Upper Jurassic (Hecht and Estes, 1960). Re-
cently, I described some tiny vertebrae found in a coprolite from the Lower
Permian of New Mexico; the caudal vertebra that I figured has a large
haemal arch and, as I pointed out, is very similar to caudal vertebrae in
living urodeles (Vaughn, X 963b)— the dorsal vertebrae in the same copro-
lite, by the way, are not at all like the vertebrae of Lasalia cutlerensis.
Kuhn (1964) has designated this caudal vertebra as the holotype of Vaughn-
iella urodeloides, a name that expresses his opinion that this animal may
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Lower Permian Frog-like Vertebrae
11
actually represent a Permian forerunner of the urodeles. I am inclined to
disagree with Kuhn’s interpretation of the central element as the inter-
centrum, but possibly he is correct in his phylogenetic assignment. If so, it
would seem that the lissamphibians were separated from other amphibians
much earlier than has been generally thought, and this would be consonant
with the studies by Parsons and Williams. If lissamphibians really were a
distinct group as early as the Permian, the lack of a better early record may
be due simply to the probably lesser frequency with which very small non-
marine vertebrates were preserved, and if preserved, the lesser frequency
with which they are discovered. The earliest usually accepted record of
salientians is in the Lower Triassic, in the form of the unique specimen of
Protobatrachus massinoti Piveteau from Madagascar. Protobatrachus shows
a mixture of primitive amphibian and advanced anuran features; as many
as sixteen presacral vertebrae were retained and the caudal vertebrae were
not fused into a urostyle, but the characteristically compound frontoparietal
bone of anurans was present. Hecht (1962) questions the relationship of
Protobatrachus to the anurans, largely on the basis of an alternative inter-
pretation of the homologies of the enlarged tarsal bones, and he suggests
the possibility, but does not insist, that the frog-like features of Proto-
batrachus are of independent, convergent origin. Griffiths (1963:275),
however, feels that “the phylogenetic relationship of Protobatrachus to
the Anura is established.” Griffiths has also examined the question of the
ontogenetic stage of the specimen of P. massinoti and (1963:277) con-
cludes, “The present thesis ... is that the Salientia evolved in water and
that when they finally emerged on land they were already pre-adapted
for jumping. If this view is correct the functional paradoxes . . . are re-
solved and Protobatrachus may be interpreted, with equal validity, either
as an adult or a late metamorphic stage.” Protobatrachus is usually classi-
fied under Proanura. Hecht (1963) says that the earliest known true frog
—that is, a member of the Anura— comes from the Lower Jurassic of Ar-
gentina and that the major frog adaptations were completed by the Middle
Jurassic. With regard to the possible pre-Triassic record of frogs, it must
be noted that Gregory’s (1950) reinterpretation of the skull roof in the
Pennsylvanian Amphibamus = Miobatrachus casts serious doubt on Watson’s
(1940) thesis that the frogs were derived from among such temnospondylous
labyrinthodont amphibians, although Griffiths (1963) feels there is still
much merit in this thesis. I have already mentioned the trackway from
the Lower Permian Ecca beds of South Africa described by Griffiths (1963),
who interprets these prints as made by fore limbs alone while the hind
limbs were supported by water, and who is inclined to ascribe the trackway
to a proanuran, If Griffiths is correct in this, it is further evidence that
the lissamphibians were separated from other amphibians at least as long
ago as the early Permian.
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There is no resemblance between the vertebrae of Lasalia cutlerensis
and the vertebrae of gymnophionans; among other things, there are no
elongated transverse processes in the gymnophionans. As for the urodeles,
their vertebrae are nectridian-like in the placement of the transverse proc-
esses midway along the centrum and in that the transverse process usually
is doubled and bears two articular facets. It is only among the salientians
that vertebrae closely similar to those of L. cutlerensis are found.
The sole specimen of Protobatrachus massinoti consists of a mould in
a nodule that has been separated into slab and counterslab. The restoration
given by Piveteau (1937:fig. 2) of the first, three vertebrae shows that the
second and third may have been like the holotype of L. cutlerensis, but
details are lacking, and there are some dissimilarities. For one difference,
the transverse process is longer in Lasalia, and in this way Lasalia is more
like true frogs. A more basic difference lies in the fact that the ribs in
Protobatrachus, as in true frogs, are single-headed whereas, as I have noted,
a facet on the centrum of the holotype of L. cutlerensis indicates that the
ribs in Lasalia were equipped not only with tubercula but also with at least
vestiges of capitula. If Lasalia and Protobatrachus are related, the indica-
tions are that the salientians lost the capitulum altogether sometime in the
interval between early Permian and early Triassic. Such a loss would be in
keeping with the general tendency to reduction of ribs in the Salientia;
among living frogs, according to Noble (1931:233), “No ribs appear as
distinct ossifications . . . higher than the Pipidae, although bits of cartilage
are frequently found on the ends of the diapophyses.” Goodrich (1930:
80-81) says “The reduced single-headed rib of Anura is attached to a
‘transverse process/ apparently a lateral outgrowth of the neural arch, but
passing below the vertebral artery. It is doubtful, however, whether the
vertebral artery in the various groups [of living amphibiansl is strictly ho-
mologous and constant in position . . . , and the evidence does not allow us to
decide whether the process represents a parapophysis which has shifted
dorsally, or a true diapophysis.” If it is true that Lasalia is related to the
Anura, it would seem that the anuran transverse process is truly a diapo-
physis. In Anura, the centrum is variously ossified. Griffiths (1963) uses
the terms ectochordal, stegochordal, and holochordal. In the ontogeny of
ectochordal centra, the entire perichordal sheath is chondrified and then
ossified; the result is an ossified cylinder with a notochordal canal. In
stegochordal centra, ossification is limited to the dorsal part of the peri-
chordal sheath. In holochordal centra, the notochord is completely replaced
by osteoid tissue; the result is a solid centrum. The Liopelmidae, gen-
erally acknowledged as the most primitive family of living frogs, have an
ectochordal centrum, and Griffiths says (1963:260) that “ Protobatrachus
. . . shows conclusively that primitive salientian vertebrae consisted of a
chain of bony, spool-shaped centra, pierced by a persistent notochord
The centrum in Lasalia may be described as holochordal, and the possession
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Lower Permian Frog-like Vertebrae
13
by this Permian animal of this supposedly advanced kind of centrum would
seem to argue against relationship of Lasalia to the Salientia. However, the
matter is not entirely clear. Hecht (1962:41) says only that in the specimen
of P. massinoti, “The vertebral impressions are well separated by large inter-
vertebral pieces of matrix. Some of these pieces of matrix, which have re-
placed the non-ossified intervertebral elements, bear conical anterior or
posterior projections, perhaps an indication that the notochord was con-
tinuous or nearly so.” These pieces of matrix may be seen in the enlarged
photographs presented by Piveteau (1937:pl. 1, fig. 4; pi. 2, fig. 1); they
certainly do not conclusively demonstrate a continuous notochordal canal.
It is obvious that the centra in Protobatrachus were amphicoelus, as in
Lasalia, and it is possible that they were not notochordal, again as in Lasalia.
In his study of the almost completely known upper Middle Jurassic or
lower Upper Jurassic frog N otobatrachus degiustoi from Argentina, Reig
(1957) describes the vertebrae as amphicoelus but not notochordal, and
Hecht (1963) notes that a re-examination of the materials indicates that the
centra are procoelus, that is, that the intervertebral elements became joined
to the posterior ends of the centra. This tends to cast further doubt on
the generally accepted theory that primitive salientians are necessarily char-
acterized by a continuous notochord. Reig’s illustration of a vertebra of
N. degiustoi in anterior view (1957:fig. 7) shows it to be fairly similar to
the holotypic vertebra of Lasalia cutler ensis, but there are differences. In
N. degiustoi, the centrum is narrower, the neural canal is larger, the neural
spine is higher, the transverse processes are as long but are not expanded
at their distal ends and do not bear articular cups, and of course there are
no capitular facets. It must be noted that the figured vertebra of N. degiustoi
was taken from the “lumbar” region; farther anteriorly, on the second
through fifth vertebrae, the places of the distal portions of the transverse
processes are occupied by well developed ribs. Indeed, Reig believes that
the transverse processes of the posterior vertebrae were formed partly
through the fusion of primitive ribs to the vertebrae. This implies another
dissimilarity to L. cutler ensis, the holotypic vertebra of which must now be
compared with the more anterior vertebrae in N otobatrachus. At this level
of the column, one difference is removed in that the transverse processes
in N otobatrachus bear articular cups at their distal ends, but these proc-
esses are much shorter than those in Lasalia. It is, of course, quite possible
that N otobatrachus, despite its early age, already represents one of many
lines of anuran radiation; Hecht (1963:20) says, “N otobatrachus represents
a generalized form with certain primitive features but which does not
clearly align itself with any of the generally considered primitive living
frogs, such as Liopelma, Ascaphus, or the discoglossids.”
Remarkably, the closest resemblances to the vertebrae of Lasalia cutler-
ensis are found among living frogs. Because of the vast interval in time
between the Permian and the Recent, it would be unreasonable to look
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No. 87
for signs of affinity with any specific family of living frogs, and the
comparison will be made in general terms although a few specific forms
will be singled out for details of similarity. The differences must be listed
first. None of the living frogs, of course, have capitular facets on their
centra; indeed, ribs are virtually absent in all except the primitive groups—
this matter has received comment above. In even holochordal forms, the
ossification of the centrum may be less complete than in Lasalia ; in Rana,
for example, much of the osteoid tissue of the center of the centrum is
finely woven, spongy, rather than composed of firm trabeculae of cancel-
lous bone such as are seen in Lasalia cutlerensis. The varieties of articular
surfaces at the ends of the centra will not be considered; the differences
result from the differential fate of the intervertebral bodies (see Griffiths,
1963), and great variation must have occurred in these surfaces throughout
the evolution of the frogs. In general, the centrum in living frogs is rela-
tively smaller, and the neural canal is relatively larger, than in Lasalia.
With this, the differences end, and the features in Lasalia are matched in
one or another of the living frogs. The surface of the centrum is relatively
smooth in most frogs but in, say, Scaphiopus there may be seen irregular
pitting not much different from what is seen in Lasalia. In Scaphiopus
and other living frogs may also be seen a posteriorwards prolongation of
the neural spine similar to the condition in the holotypic vertebra of
L. cutlerensis. In the vertebra that is referred to L. cutlerensis , this poster-
iorwards prolongation is lacking, but it will be remembered that this re-
ferred vertebra has transverse processes of lesser length and diameter than
the holotype and it also seems to lack facets for costal capitula; this seems
to indicate derivation from a more posterior position in the vertebral col-
umn. This suggests an analogy to the condition in, say, Rana, in which the
posteriorwards projection is strongly developed in the anterior part of the
column but becomes much less pronounced in the posterior vertebrae. The
neural spines in Rana are lower but in, for example, Ceratophrys they
are as high as in Lasalia. In some living frogs the transverse processes are
shorter than in Lasalia, and in others they are longer. Although the trans-
verse processes are longer in such a frog as Rana catesbiana, they are very
similar to the processes in L. cutlerensis, especially if one considers the fourth
vertebra in R. catesbiana. The similarity extends even to the presence of a
ridge along the anterior border of the process. The processes in R. catesbiana
are hollow like those in L. cutlerensis except that in R. catesbiana the
distal ends are plugged with calcified tissue; another minor difference is
that the distal ends of the processes are somewhat flattened dorsoventrally
in R. catesbiana.
The differences noted make it clear that the vertebrae in Lasalia
cutlerensis are far from exactly similar to those of any living frog, but the
resemblances are strikingly close, and the dissimilarities are such as could
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Lower Permian Frog-like Vertebrae
15
reasonably be expected to have been mitigated in the interval of time
under consideration. It is far from my intention to claim that Lasalia was
a salientian or even a forerunner of the salientians, but I do stress that
the vertebrae of L. cutler ensis are remarkably like those seen in salientians,
and I do suggest the possibility of a phylogenetic connection. Should fu-
ture finds support this suggestion, we may need to revise our opinion as
to the degree of antiquity of various skeletal characteristics of those living
frogs generally considered to be the most primitive, for example, the
Liopelmidae. That these frogs are generally acknowledged as very gen-
eralized is undoubtedly justified, but it may become necessary to inquire
into the possibility of origin through secondary simplification of features
such as ectochordal centra. As Ritland has summed it up in his study of
the liopelmid Ascaphus (1955:279-280), “Although frequently stated to be
the most primitive anuran, in reality Ascaphus is not primitive. Few frogs
or toads are more highly specialized for life in a unique but uniform
habitat . . . Ascaphus together with Leiopelma unquestionably possesses
many generalized features, especially in those systems which function pri-
marily in locomotion and support . . . Nevertheless, they are unmistakably
anurans, with the majority of the typical anuran specializations.”
ORIGINS
Even if we were to assume that the Permian Lasalia represents the
forerunners of the salientians, we should be very little nearer to an under-
standing of the ultimate origins of this group; we are left with the theories
already current. Watson (1940) suggested that temnospondylous labyrin-
thodont amphibians like the Pennsylvanian Amphibamus are the most likely
ancestors of the salientians and, notwithstanding Gregory’s (1950) rein-
terpretation of the elements of the posterior skull table in Amphibamus,
some authors (Eaton, 1959; Griffiths, 1963) still feel that the sources of the
Salientia are to be found among such temnospondyls. They may be right,
but Williams (1959) has noted that the pleurocentral nature of the frog
centrum suggests that if the salientians really are related to the labyrintho-
dont amphibians, the affinities are more likely with the anthracosaurs than
with the temnospondyls. No anthracosaur yet known seems a likely candi-
date for such relationship, but Panchen (1959) has demonstrated that some
of the amphibians assigned to the Temnospondyli, namely, the plagiosaurs,
actually have a pleurocentral, rather than a basically intercentral, centrum.
The plagiosaurs are mainly a Triassic group as far as known, but Panchen
has described an Upper Permian member. This is not to suggest that the
plagiosaurs had anything to do with the salientians, but it does show that
the pleurocentral centrum occurred in more Permian amphibians than we
have thought, and this may eventually come to have some bearing on the
16
Contributions in Science
No. 87
origins of the Salientia. Meanwhile, I must subscribe to the opinion of Par-
sons and Williams (1963) that we cannot, with any confidence, point to any
known group of Paleozoic amphibians as ancestral to the Recent orders.
If Lasalia and Vaughniella (mentioned in the preceding section) really
are related to the lissamphibians of modern times, the term “Amphibia”
includes a staggering array of different structural types even as early as the
Permian. It would include not only the lissamphibians but also: the lepo-
spondyls, a highly varied and probably, taxonomically, artificial assemblage;
the labyrinthodont amphibians, comprising temnospondyls, plagiosaurs,
anthracosaurs and perhaps also the plesiopodans (see Eaton, 1960); the sey-
mouriamorphs, classified sometimes as labyrinthodonts, sometimes as co-
tylosaurian reptiles; seymouriamorph-diadectid intermediates such as
Tseajaia (see Vaughn, 1964); and, according to Romer’s re-evaluation
(1964), Diadectes, an animal formerly classified with the cotylosaurian
reptiles. Relationship of the labyrinthodonts, seymouriamorphs and diadec-
tids seems to be adequately documented, but this is not sufficient reason
to deny the existence of an amnion in all these forms; an amnion may well
have been present in the more reptile-like members, particularly the diadec-
tids. Further, this large group has not been clearly shown to be related to
the lepospondyls, and the lissamphibians would seem to comprise still an-
other entity. The term “Amphibia” is useful in a generally descriptive sense,
but in the sense of designation of discrete phylogenetic units, it may prove
to have about the same value as the term “Pisces.”
ACKNOWLEDGMENTS
I extend my thanks to my able field assistants, Bruce Bartholomew,
David Berman, Timothy Brown, and Thomas Kauffman.
LITERATURE CITED
Baars, D. L.
1962. Permian System of Colorado Plateau. Bull. Amer. Assoc. Petrol.
Geol., 46:149-218.
Baker, A. A.
1936. Geology of the Monument Valley-Navajo Mountain region, San Juan
County, Utah. Bull. U. S. Geol. Survey, 865:1-106.
Dunbar, C. O et al.
1960. Correlation of the Permian formations of North America. Bull. Geol.
Soc. Amer., 71:1763-1806.
Eaton, T. H., Jr.
1959. The ancestry of modern Amphibia: a review of the evidence. Univ.
Kansas Publ. Mus. Nat. Hist., 12:155-180.
1960. A new order of fishlike Amphibia from the Pennsylvanian of Kansas.
Univ. Kansas Publ. Mus. Nat. Hist., 12:217-240.
Elston, D. P., E. M. Shoemaker, and E. R. Landis
1962. Uncompahgre front and salt anticline region of Paradox Basin, Colo-
rado and Utah. Bull. Amer. Assoc. Petrol. Geol., 46:1857-1878.
1965
Lower Permian Frog-like Vertebrae
17
Goodrich, E. S.
1930. Studies on the Structure and Development of Vertebrates. London:
MacMillan and Co., 837 pp.
Gregory, J. T.
1950. Tetrapods of the Pennsylvanian nodules from Mazon Greek, Illinois.
Amer. J. Sci., 248:833-873.
Gregory, J. T., F. E. Peabody, and L. I. Price
1956. Revision of the Gymnarthridae, American Permian microsaurs. Bull.
Peabody Mus. Nat. Hist., 10:1-77.
Griffiths, I.
1963. The phylogeny of the Salientia. Biol. Rev., 38:241-292.
Hecht, M. K.
1962. A reevaluation of the early history of the frogs. Part I. Syst. Zool.,
11:39-44.
1963. A reevaluation of the early history of the frogs. Part II. Syst. Zool.,
12:20-35.
Hecht, M. K., and R. Estes
1960. Fossil amphibians from Quarry Nine. Postilla, Peabody Mus. Nat.
Hist., 46:1-19.
Hotton, N., Ill
1952. Jaws and teeth of American xenacanth sharks. J. Paleont., 26:489-500.
Jarvik, E.
1955. The oldest tetrapods and their forerunners. Sci. Monthly, 80:141-154.
Kuhn, O.
1964. Cyrtura Jaekel aus dem Solnhofener Schiefer ist ein Nachzugler der
Temnospondyh (Amphibia, Labyrinthodontia ) . Neues Jahrbuch fur
Geologie und Palaontologie, Stuttgart, Monatshefte, 1964, 11:659-664.
Langston, W., Jr.
1953. Permian amphibians from New Mexico. Univ. Calif. Publ. Geol. Sci.,
29:349-416.
Noble, G. K.
1931. The Biology of the Amphibia. London and New York: McGraw-
Hill Co., 577 pp.
Panchen, A. L.
1959. A new armoured amphibian from the Upper Permian of East Africa.
Philos. Trans. Roy. Soc. London, B, 242:207-281.
Parsons, T. S., and E. E. Williams
1962. The teeth of Amphibia and their relation to amphibian phylogeny.
J. Morph., 110:375-389.
1963. The relationships of the modern Amphibia: a re-examination. Quart.
Rev. Biol., 38:26-53.
Piveteau, J.
1937. Un amphibien du Trias inferieur; essai sur Torigine et revolution
des amphibiens anoures. Annales de Paleontologie, Paris, 26:133-177.
Reig, O. A.
1957. Los anuros del Matildense. Acta Geologica Lilloana, Buenos Aires,
1:231-297.
Ritland, R. M.
1955. Studies on the post-cranial morphology of Ascaphus truei. II. Myol-
ogy. J. Morph., 97:215-282.
18
Contributions in Science
No. 87
Romer, A. S.
1956. Osteology of the Reptiles. Chicago: Univ. Chicago Press, 772 pp.
1960. The vertebrate fauna of the New Mexico Permian. New Mexico
Geol. Soc. Guidebook of Rio Chama Co., 11th Field Conf., pp. 48-54.
1964. Diadectes an amphibian? Copeia, 1964 (4) :718-719.
Sawin, H. J.
1941. The cranial anatomy of Eryops megacephalus. Bull. Mus. Comp.
Zool., 88:407-463.
Vaughn, P. P.
1962. Vertebrates from the Halgaito tongue of the Cutler Formation, Per-
mian of San Juan County, Utah. J. Paleont., 36:529-539.
1963a. The age and locality of the late Paleozoic vertebrates from El Cobre
Canyon, Rio Arriba County, New Mexico. J. Paleont., 37:283-286.
1963b. New information on the structure of Permian lepospondylous verte-
brae—from an unusual source. Bull. So. Calif. Acad. Sci., 62:150-158.
1964. Vertebrates from the Organ Rock Shale of the Cutler Group, Permian
of Monument Valley and vicinity, Utah and Arizona. J. Paleont.,
38:567-583.
Watson, D. M. S.
1940. The origin of frogs. Trans. Roy. Soc. Edinburgh, 40:195-231.
Weir, G. W., W. P. Puffett, and C. L. Dodson
1961. Preliminary geologic map and section of the Mount Peale 4 NW
Quadrangle, San Juan County, Utah. U. S. Geol. Survey, Min. Invest.,
Map MF-151.
Williams, E. E.
1959. Gadow’s arcualia and the development of tetrapod vertebrae. Quart.
Rev. Biol., 34:1-32.
LOS
ANGELES
COUNTY
MUSEUM
Nmber 88
CONTRIBUTIONS
IN SCIENCE
June 28, 1965
GEOLABIS WOLFF I, A NEW FOSSIL INSECTIVORE FROM
THE LATE OLIGOCENE OF SOUTH DAKOTA
By J. R. Macdonald
I
■ f .
Los Angeles County Museum
• Exposition Park
• Los Angeles, Calif. 90007
CONTRIBUTIONS IN SCIENCE is a series of miscellaneous tech-
nical papers in the fields of Biology, Geology and Anthropology, published
at irregular intervals by the Los Angeles County Museum. Issues are num-
bered separately, and numbers run consecutively regardless of subject mat-
ter. Number 1 was issued January 23, 1957. The series is available to scien-
tists and scientific institutions on an exchange basis. Copies may also be
purchased at a nominal price.
INSTRUCTIONS FOR AUTHORS
Manuscripts for the LOS ANGELES COUNTY MUSEUM CONTRIBU-
TIONS IN SCIENCE may be in any field of Life or Earth Sciences. Acceptance of
papers will be determined by the amount and character of new information and the
form in which it is presented. Priority will be given to manuscripts by staff members,
or to papers dealing with specimens in the Museum’s collections. Manuscripts must
conform to CONTRIBUTIONS style and will be examined for suitability by an
Editorial Committee. They may also be subject to critical review by competent
specialists.
MANUSCRIPT FORM.— (1) The 1960 AIBS Style Manual for Biological
Journals is highly recommended as a guide. (2) Typewrite material, using double
spacing throughout and leaving ample margins, on only one side ot 8V2 x 11 inch
standard weight paper. (3) Place tables on separate pages. (4) Footnotes should be
avoided if possible. (5) Legends for figures and unavoidable footnotes should be
typed on separate sheets. Several of one kind may be placed on a sheet. (6) Method
of literature citation must conform to CONTRIBUTIONS style— see number 50 and
later issues. Spell out in full the title of non-English serials and places of publication.
(7) A factual summary is recommended for longer papers. (8) A brief abstract
should be included for all papers. This will be published at the head of each paper.
ILLUSTRATIONS. — All illustrations, including maps and photographs, should
be referred to as “figures!’ All illustrations should be of sufficient clarity and in the
proper proportions for reduction to CONTRIBUTIONS page size. Permanent ink
should be used in making line drawings and in lettering (do not type on drawings);
photographs should be glossy prints of good contrast. Original illustrations will not
be returned unless specifically requested when the manuscript is first submitted. Au-
thors may also request their engravings at this time.
PROOF. — Authors will be sent galley proof which should be corrected and re-
turned promptly. Changes after the paper is in galley will be billed to the author. Un-
less specially requested, page proof will not be sent to the author. 100 copies of each
paper will be given free to a single author or divided equally among multiple authors.
Orders for additional copies should be sent to the Editor at the time corrected galley
proof is returned; appropriate forms for this will be included when galley is sent.
David K. Caldwell
Editor
GEOLAB IS WOLFF I, A NEW FOSSIL INSECTIVORE FROM
THE LATE OLIGOCENE OF SOUTH DAKOTA
By J. R. Macdonald1
Abstract: A new species of Geolabis is described from the
upper part of the Poleslide member of the Brule formation in the
White River Badlands of South Dakota. This record extends the
range of the genus to the late Oligocene.
As part of a project to develop the early Miocene fauna of the Wounded
Knee Area, Shannon County, South Dakota, a portion of the 1964 field season
was devoted to prospecting the upper 200 feet of the Poleslide member of the
Brule formation which conformably underlies the basal Miocene Sharps for-
mation in this area. This collecting was the beginning of an attempt to fill in
the faunal gap between the late Whitneyan faunas and the early Arikareean
faunas of this region. This collecting indicates that the faunal gap is not real
but results from either a lack of thorough collecting in these beds or the non-
reporting of collected material which lies unpublished in museum drawers.
The specimen described below extends the range of Geolabis into the later
part of the Whitneyan. It represents a species which became a “giant” in its
line, and it serves to re-emphasize that, despite more than a hundred years of
collecting in the White River Badlands, there is still much to be learned of the
White River fauna.
The field work in South Dakota during 1964 was supported by NSF Grant
GN-3. The photographs were made by Mr. Armando Solis, Museum Photog-
rapher for the Los Angeles County Museum (LACM).
The new form may be described as follows:
INSECTIVORA Bowdich, 1821
Erinaceidae Bonaparte, 1838
Geolabidinae McKenna, 1960
Geolabis Cope, 1884
Geolabis Cope, 1884 o, p. 807
1Curator of Vertebrate Paleontology, Los Angeles County Museum.
SMITHS:
msTiTumis
hit
sJIUti
4
Contributions in Science
No. 88
Figure 1. Geolabis wolffi new species, LACM 9582. Fragment of right maxillary with
P4-M2; labial, crown, and lingual views. (To be viewed with a stereoscope. Ca. X 5.)
1965
New Fossil Insectivore from South Dakota
5
Geolabis wolffi,2 new species
Figure 1
Type— LACM No. 9582, fragment of right maxillary with P4 - M2.
Type Locality — LACM 1990, Wolff Ranch Badlands, Shannon County,
South Dakota.
Horizon— Poleslide member of Brule formation in grey zone from 175
feet to 135 feet below base of Sharps formation. Late Oligocene.
Diagnosis — P4 four-rooted with well developed hypocone; M1 with pro-
toconule and metaconule; M2 with deeply incurved labial wall, anterior cingu-
lum extends linguad beyond protocone, and there is a small cingular cusp on
the anterior cingulum opposite the hypocone.
Description— P4 with very tall paracone dominating tooth; metacone rep-
resented by ridge sweeping posterad and labad to join metastyle; protocone
much smaller than paracone, joined near apex by shelf-like anterior cingulum
which extends labad to the parastyle; hypocone less developed than on molars
but supported by root; posterior cingulum not shelf-like and not prominent;
conical parastyle extends anterad of transverse base of crown, with small lin-
gual satellite style separated by small notch on anterior cingulum; labial cingu-
lum relatively weakly developed, rising to apex on side of parastyle to form
incipient mesostyle; metastyle small, broken away. M1 with large sub-equal
paracone and metacone, paracone conical, metacone with crest to metastyle as
in P4, protocone worn flat, with cingula-like shelves extending labially to base
of parastyle and metastyle; small ridges extend from protocone-parastyle ridge
and protocone-metastyle ridge to base of paracone and metacone represent
protoconule and metaconule; anterior cingulum shelf-like at base of crown,
extends from point below lingual edge of paracone to antero-lingual “corner”
of protocone, with small cuspule at lingual end; posterior cingulum broader
than anterior, extending along base of crown from below labial edge of meta-
cone to postero-lingual “corner” of protocone, hypocone worn but well de-
fined; labial cingulum widely expanded, ridge extending from base of paracone
to antero-labial corner expands to form twinned parastyles which are separated
on labial wall by notch, metastyle worn, elongated postero-labially. M2 similar
to M1 except labial wall compressed antero-posteriorly so parastyle and meta-
style expand labially leaving deep re-entrant at center of labial wall; anterior
cingulum more shelf-like.
Discussion— McKenna (1960) has thoroughly reviewed the Geolabidinae
and indicated the supposed synonymy within the subfamily. The specimen de-
scribed above differs from the two forms known from upper dentitions: Geo-
labis marginalis (Cope) and Metacodon mellingeri Patterson and McGrew
(1937)— which McKenna (1960:135) believes to be a junior synonym of G.
rhynchaeus.
2For Mr. Otto Wolff of Rapid City and Rockyford, South Dakota.
6
Contributions in Science
No. 88
Geolabis wolffi differs basically from both species in the presence of a
strong hypocone on P4 with a supporting root and the development of well-
defined cusp on the lingual end of the anterior cingulum of M2. The large size
may or may not be significant as a distinguishing character.
Other minor differences may be noted among the three species of Geo-
labis, but they are overshadowed by those listed above.
Table of Measurements
Transverse
Anteroposterior
diameter from
diameter from
parastyle-meta-
parastyle to
style labial
border to proto-
cone—hypocone
border
metastyle
P1
2.9
3.2
M1
ca. 3.0
3.8
M2
2.6
4.1
Literature Cited
Cope, E. D.
1884o. The Vertebrata of the Tertiary formations of the West. Book I. Report
U. S. Geolog. Survey of the Territories, F. V. Hayden, U. S. Geologist in
Charge, Washington, D. C., pp. i-xxxv, 1-1009, pis. 1-lxxva.
McKenna, M. C.
1960. The Geolabidinae. A new subfamily of early Cenozoic Erinaceoid In-
sectivores. Univ. Calif. Pubis, in Geol. Sci., 37 (2) : 131-164, 6 figs.
Patterson, V., and P. O. McGrew
1937. A soricid and two erinaceids from the White River Oligocene. Field
Mus. Nat. Hist., Geol. Ser., 6:245-272, 15 figs.
LOS
i1
ANGELES
COUNTY
MUSEUM
CONTRIBUTIONS
IVdiC IN SCIENCE
Number 89 June 28, 1965
A NEW SOUTH AMERICAN TOE BITER
(HEMIPTERA, BELOSTOMATIDAE)
By A. S. Menke
Los Angeles County Museum • Exposition Park • Los Angeles, Calif. 90007
A NEW SOUTH AMERICAN TOE BITER
(HEMIPTERA, BELOSTOMATIDAE)
By A. S. Menke1
Abstract: A new species of toe biter, Belostoma rhom-
boides, is described from French Guiana, South America. Among
material recently received from South America, an interesting
specimen of Belostoma was found which has proved to be a new
species. The terminology used in the description has been ex-
plained by Lauck (1962) and Lauck and Menke (1961).
Belostoma rhomboides, new species
Figures 1 and 2
Holotype female — Length 21.5 mm., width 12.75 mm., width of head 5
mm., length of head 3 mm., width of pronotum 7 mm., length of pronotum 3
mm.; proportions of head as follows: length of anteoculus:interoculus (4:3.5),
eye length:eye widthimaximum interocular width (3.5:3. 5:9.5), length of
beak segment I: II (3.5: 4.5).
Structural characters .—Base of clypeus reaching ocular line; beak stout,
segment I about one half as broad as long; antenna four segmented, II and III
bearing long finger-like projections; eye globose, outer margin rounded; pro-
sternal keel not developed, prosternum flattened and V-shaped, apex of V di-
rected posteriorly; front tarsus one-segmented (true segments II-III fused);
clavus with a few faint veins; corium with a prominent network of veins; em-
bolium very broad, strongly curved at middle (Fig. 1); membrane narrow,
greatest width equal to greatest width of clavus (Fig. 1) ; flight wings fully de-
veloped; outer margin of ventral laterotergites not continuous, but serrate,
margin of each tergite expanding posteriorly (Fig. 2); visible laterotergites II-
V with a narrow band of long silky hair, remainder of abdominal venter cov-
ered with short, dense setae.
Distinctive markings — Posterolateral angle of pronotum with a large, pale,
yellow brown spot; base of clavus and corium, and embolium at embolial frac-
ture, pale yellow brown; remainder of dorsum typically dark brown; venter and
legs dark brown but anterolateral angle of ventral laterotergites IV-VI with a
pale yellow brown spot (Fig. 2).
Distribution— Known only from holotype female. French Guiana (no
further data). Type deposited in the Los Angeles County Museum, Los
Angeles.
iResearch Associate, Los Angeles County Museum; Department of Entomology,
University of California, Davis.
1965
New South American Toe Biter
3
Figures 1 and 2, dorsal and ventral aspect, respectively, of holotype of Belostoma
rhomboides, new species.
SMfTHStfr
mi O
4
Contributions in Science
No. 89
Discussion —Belostoma rhomboides can be separated readily from all
other known Belostoma by the one-segmented front tarsus, form of prosternum,
narrow forewing membrane and the serrate abdominal margin. The only other
belostomatids that have a one-segmented tarsus are the species of Horvathinia
and Diplonychus urinator (Dufour). Belostoma rhomboides does not fit any
of the species groups outlined by Lauck (1962) and probably should be placed
in a group by itself. The affinities of this species are not clear. The small eyes
and narrow membrane are suggestive of the genus Abedus but the air straps are
typically Belostoma. The discovery of a male should shed some light on the re-
lationships of rhomboides since the genitalia are diagnostic in most of Lauck’s
species groups.
Literature Cited
Lauck, D. R.
1962. A monograph of the genus Belostoma, Part I, Introduction and B. den-
tatum and subspinosum groups. Bull. Chicago Acad. Sci., 11(3) : 34-81.
Lauck, D. R., and A. S. Menke
1961. The higher classification of the Belostomatidae. Ann. Entomol. Soc.
Amer., 54:644-657.
LOS
ANGELES
COUNTY
MUSEUM
Dumber 90
CONTRIBUTIONS
5c\lU IN SCIENCE
June 28, 1965
NORMICHTHYS YAHGANORUM , A NEW SEARSIID FISH
FROM ANTARCTIC WATERS
By Robert J. Lavenberg
Los Angeles County Museum
Exposition Park
Los Angeles, Calif. 90007
CONTRIBUTIONS IN SCIENCE is a series of miscellaneous tech-
nical papers in the fields of Biology, Geology and Anthropology, published
at irregular intervals by the Los Angeles County Museum. Issues are num-
bered separately, and numbers run consecutively regardless of subject mat-
ter. Number 1 was issued January 23, 1957. The series is available to scien-
tists and scientific institutions on an exchange basis. Copies may also be
purchased at a nominal price.
INSTRUCTIONS FOR AUTHORS
Manuscripts for the LOS ANGELES COUNTY MUSEUM CONTRIBU-
TIONS IN SCIENCE may be in any field of Life or Earth Sciences. Acceptance of
papers will be determined by the amount and character of new information and the
form in which it is presented. Priority will be given to manuscripts by staff members,
or to papers dealing with specimens in the Museum’s collections. Manuscripts must
conform to CONTRIBUTIONS style and will be examined for suitability by an
Editorial Committee. They may also be subject to critical review by competent
specialists.
MANUSCRIPT FORM.— (1) The 1960 AIBS Style Manual for Biological
Journals is highly recommended as a guide. (2) Typewrite material, using double
spacing throughout and leaving ample margins, on only one side of 8V2 x 11 inch
standard weight paper. (3) Place tables on separate pages. (4) Footnotes should be
avoided if possible. (5) Legends for figures and unavoidable footnotes should be
typed on separate sheets. Several of one kind may be placed on a sheet. (6) Method
of literature citation must conform to CONTRIBUTIONS style — see number 50 and
later issues. Spell out in full the title of non-English serials and places of publication.
(7) A factual summary is recommended for longer papers. (8) A brief abstract
should be included for all papers. This will be published at the head of each paper.
ILLUSTRATIONS.— All illustrations, including maps and photographs, should
be referred to as “figures!’ All illustrations should be of sufficient clarity and in the
proper proportions for reduction to CONTRIBUTIONS page size. Permanent ink
should be used in making line drawings and in lettering (do not type on drawings);
photographs should be glossy prints of good contrast. Original illustrations will not
be returned unless specifically requested when the manuscript is first submitted. Au-
thors may also request their engravings at this time.
PROOF. — Authors will be sent galley proof which should be corrected and re-
turned promptly. Changes after the paper is in galley will be billed to the author. Un-
less specially requested, page proof will not be sent to the author. 100 copies of each
paper will be given free to a single author or divided equally among multiple authors.
Orders for additional copies should be sent to the Editor at the time corrected galley
proof is returned; appropriate forms for this will be included when galley is sent.
David K. Caldwell
Editor
NORMICHTHYS YAHGANORUM, A NEW SEARSIID FISH
FROM ANTARCTIC WATERS
By Robert J. Lavenberg1
Abstract: A new species of searsiid fish, Normichthys
yahganorum, is described from two specimens obtained in the
southeastern Pacific Ocean. The new species is the third known
member of the genus. N. yahganorum differs from both previ-
ously known species, N. operosa and N. campbelli, in having
fused gill filaments. Other features utilized to distinguish the
three species include longitudinal scale rows, ventral rays and
gill rakers.
In the exploratory investigations of the Antarctic biota by members of the
department of biological sciences of the University of Southern California, the
United States Antarctic Research Vessel USNS Eltanin has undertaken several
cruises along the Chilean coast in the southeast Pacific Ocean. The ship usually
departs from Valparaiso, and proceeds south to 40° where biological opera-
tions in the Antarctic begin. Among the fishes collected off southern Chile dur-
ing Cruises 5 and 15 are two moderate-sized searsiids. The combination of
dermal pits above the lateral line canal and the absence of photophores readily
diagnoses these individuals as members of the genus Normichthys Parr ( 1960) .
In identifying these two slickheads, an unusual arrangement of the gill
filaments was noted. This characteristic and several other meristic features
were noted that distinguish the Antarctic forms from all other known species
of the genus. The material differs so markedly from the other Normichthys
that I consider them representatives of a distinct species.
The material has been deposited in the fish collections of the Los Angeles
County Museum (LACM). The new species may be known as:
Normichthys yahganorum, new species
Figures 1 and 2
Holotype— LACM 10264; immature male; 95.3 mm. in standard length (SL);
off southern Chile, approximately 60 miles W and just S of Isla Gamblin (45°
OF S, 76° 33' W at beginning of haul); Eltanin station 215; 10-foot midwater
trawl (IKMWT); maximum depth of trawl 1100 m., over a bottom of 3180
m.; 14 September 1962.
Paratype— LACM 10265; immature female; 76 mm. in SL; off southern
Chile (38° 00' S, 74° 48' W at beginning of haul); Eltanin station 1286; 10-
1Assistant Curator of Ichthyology, Los Angeles County Museum.
4
Contributions in Science
No. 90
Figure 1. Normichthys yahganorum, new species, Paratype, LACM 10265. Imma-
ture female, 76 mm. SL, illustrating the slender shape of young individuals.
Figure 2. Normichthys yahganorum, new species, Holotype, LACM 10264. Imma-
ture male, 95 mm. SL.
1965
New Antarctic Searsiid Fish
5
foot IKMWT; maximum depth of trawl 2350 m., over a bottom of 4660 m.;
trawl fished between 2045 and 0330 hours; 2 October 1964.
Diagnosis — A Normichthys differing from N. operosa and N. camp belli
in having the gill filaments fused and forming a flap-like extension of tissue
from the gill arch instead of no fusion of gill filaments; short gill filaments
present on periphery of tissue flap in yahganorum; in having smaller scales,
1 1 1-117 in the longitudinal series rather than 80-90 ( operosa ) or 65-71 (camp-
belli)', in having seven ventral rays rather than six ( operosa ) or eight (camp belli);
and in having an intermediate number of gill rakers, 6-8+1 + 16-17 rather than
7+20-21 ( operosa ) or 5-6+1 + 12-14 (campbelli).
Description — Body strongly compressed, tapering slightly to caudal pe-
duncle in larger individuals and more strongly in smaller individuals; greatest
depth just anterior to ventral fins, depth tapering more strongly from origin of
dorsal to caudal peduncle in larger than in smaller forms, caudal peduncle
depth variable with size of individual; depth slightly increased by a moderately
sharp, short fleshy dorsal comb as in the Platytroctinae, and a similar but less
distinct ventral fleshy portion extending through length of anal fin base; dorsal
and ventral combs probably more prominent in smaller individuals; greatest
width at head directly behind eye. Dorsal, anal, and procurrent rays moderate-
ly elevated, ventral fins not elevated. Shoulder organ inconspicuous, small
basal portion lying directly above the insertion of the pectoral fins, short tube
extending posteriorly over five scale rows. Two dermal pits just above lateral
line canal, their position above and midway between angle of preopercle and
shoulder organ. Cleithra protruding from body on ventral side of body between
gill membranes, a flap of scaled tissue surrounding protruding cleithra. Anal
papilla strongly tapered. Head moderately pointed; dorsal comb structure
originating at nape directly above preopercle; flattened in nape region but con-
cave in interorbital space; roof of skull with a wide extent in interorbital space
but narrowing sharply just anterior to orbits; frontals laterally flattened, ex-
tending slightly over margin of eye, upper lateral surface rugose; dorsal profile
descending in a gentle slope from posterior interorbital region to tip of snout;
ventral profile following a straight line between slightly protruded cleithra and
posterior margin of lower jaw, lower jaw rising in a gentle slope from posterior
margin to snout tip; snout pointed, ending at junction with forward directed
premaxillary tusks. Snout length greater than interorbital width at mid-orbits,
both less than eye diameter. Nares flapless. Jaws of moderate length, pointed;
two supramaxillaries; upper jaw shorter than lower jaw; posterior edge of
maxillary extending just behind pupil; teeth on premaxillary well developed and
uniserial, a pair of tusks directed anteriorly; maxillary dentition weaker than
premaxillary, teeth small and uniserial; dentitional pattern of dentary like that
of maxillary, a short mid-dentary tooth row present; one pair of elongate teeth
on head of vomer; palatine toothless; tongue without teeth but covered with
numerous spinous papillae. Teeth of lower jaw insert inside upper jaw series
when mouth is closed.
Scales cycloid, thin and oval in shape; small and adherent, completely
covering the body; head scaleless; heavily marked by annuli, a few ridge-like
6
Contributions in Science
No. 90
furrows on scales suggesting radii; 111 to 117 scales in a longitudinal series
along lateral line, 19 scale rows above lateral line and 16 scale rows below;
lateral line semi-distinct, 31 to 34 pores present along its longitudinal extent;
a small series of lateral line pores present over urostyle; a single pore present
in epidermis below each body scale.
Gill rakers moderately long, constituting about five per cent of SL. Gill
filaments fused along entire extent of gill arch giving rise to a broad flap of gill
filament tissue, gill filament tissue flap about two to three per cent of SL at its
greatest width on lower limb of arch; small pseudobranch present; a large
white gland present under gill cover in region of preopercle.
Dorsal and anal fins subequal, anal origin slightly posterior to dorsal
origin; origin of dorsal nearer to caudal fin than snout, first three or four rays
anterior to anal origin but closer to anal origin than to ventral origin; ventral
fins originate nearer to caudal fin than snout; pectoral fin base inserted about
one-fifth of way up side of body, its position horizontal with body plane, pec-
toral rays short and slender, length of rays about equal to length of base; ven-
tral rays short, although slightly damaged they apparently equal length of
pectoral fin rays.
Inner surface of peritoneum slightly pigmented with various shades of
brown, from light tan to dark brown, in a reticulate pattern. A thin-walled
stomach present. Four large pyloric caeca, the first and third branched.
Counts and Measurements —The following counts are for both specimens.
Dorsal rays 19, anal rays 17, pectoral rays 16, ventral rays 7, branchiostegal
rays 7, gill rakers 6-8+1 + 16-17, and vertebrae 44. Measurements for the
specimens are given in Table 1 .
Remarks —Nor michthys yahganorum represents the first occurrence of
this genus in the Antarctic region of the Pacific Ocean. Although the new species
is quite distinct from N. operosa Parr (1951) and N. campbelli Lavenberg
(1965), it shares certain characteristics with these species including the ab-
sence of photophores, the presence of dermal pits, subequal dorsal and anal
fins, and a thickened ventral abdominal wall. The distinctness of N. yahganor-
um is shown in several features including the reduced number of dermal pits;
only two pits are present in N. yahganorum while in the other species the num-
ber ranges from three to seven. There is no pore in the body scales of the new
species as reported in N. operosa (Parr, 1960). A pore exists in the epidermis
beneath each scale. The lateral line is distinct in N. yahganorum but reduced
and indistinct in N. operosa and N. campbelli. A striking feature of N. yah-
ganorum is the development of a dorsal and ventral keel similar to that of the
Platytroctinae. This keel or comb is weakly developed but present.
N. yahganorum and N. operosa have dermal pits lying equidistant be-
tween the top of the gill slit and the shoulder organ. The dermal pits of N.
campbelli are just anterior to the top of the gill slit.
In all three species of Normichthys the upper branchiostegal rays are
broad and flattened while the lower rays are slender ray-like structures.
This species is named for the Yahgan Indians, archipelagic shellfish
gatherers of Tierra del Fuego, who practiced shellfish conservation and avoid-
1965
New Antarctic Searsiid Fish
7
ed exhausting their food supply. Normichthys yahganorum occurs in deep
water along the Chilean coastline where this South American Indian culture
once flourished.
Distribution— The three species currently recognized in the genus Norm-
ichthys have widely separated geographic ranges. N. operosa occurs in the
eastern Atlantic Ocean. N. campbelli inhabits the midwaters of the eastern
north Pacific Ocean, off southern California and Baja California. N. yahga-
norum apparently lives in deep water in the Antarctic region of the south-
eastern Pacific Ocean.
Acknowledgments.—! am grateful to William A. Bussing, David K. Cald-
well and Jay M. Savage for their critical review of the manuscript. The mate-
rial was collected aboard the USNS Eltanin by Hugh H. DeWitt, Thomas Hop-
kins and Richard F. McGinnis. The photographs were made by the Museum
staff photographer, Armando Solis. The work was financed in part by National
Science Foundation Research Grant G- 19497, under sponsorship of the
United States Antarctic Research Program.
TABLE 1
Measurements of Normichthys yahganorum, new species,
expressed in mm. ; figure in parentheses is per cent of SL.
LACM 10264 LACM 10265
Character
Holotype
Paratype
Standard Length
95.3
76.0
Head Length
28.7
(30.1)
25.1
(33.0)
Snout Length
7.7
( 8.1)
6.0
( 7.9)
Eye Diameter
8.5
( 8.9)
8.2
(10.8)
Interorbital Width at Mid-orbits
8.4
( 8.8)
6.1
( 8.0)
Maxillary Length
13.9
(14.6)
11.5
(15.1)
Mandible Length
15.9
(16.7)
13.4
(17.6)
Predorsal Length
60.6
(63.6)
47.2
(62.1)
Dorsal Fin Base Length
19.0
(19.9)
14.7
(19.3)
Preanal Length
64.1
(67.3)
48.6
(63.9)
Anal Fin Base Length
18.2
(19.1)
13.1
(17.2)
Prepectoral Length
32.3
(33.9)
25.4
(33.4)
Preventral Length
53.0
(55.6)
40.4
(53.2)
Pre-shoulder Organ Length
33.2
(34.8)
26.8
(35.3)
Body Depth Just Anterior To Ventral Fins
24.1
(25.3)
15.4
(20.3)
Least Depth Caudal Peduncle
10.2
(10.7)
5.3
( 6.9)
Caudal Peduncle Length
18.6
(19.5)
15.6
(20.5)
Literature Cited
Lavenberg, R. J.
1965. A new species of searsiid fish, Normichthys campbelli, from the eastern
North Pacific Ocean. Bull. So. Calif. Acad. Sci., 64(1): 22-26.
Parr, A. E.
1951. Preliminary revision of the Alepocephalidae, with the introduction of a
new family, Searsidae. Amer. Mus. Novitates, (1531) : 1-21.
1960. The fishes of the family Searsidae. Dana Rept., 51: 1-104.
LOS
ANGELES
COUNTY
MUSEUM
Number 91
CONTRIBUTIONS
HXhi IN SCIENCE
June 28, 1965
OBSERVATIONS ON CAPTIVE AND WILD ATLANTIC
BOTTLENOSED DOLPHINS, TURSIOPS T RUN CATVS, IN
THE NORTHEASTERN GULF OF MEXICO
by Melba C. Caldwell, David K. Caldwell and J. B. Siebenaler
Los Angeles County Museum
Exposition Park
Los Angeles, Calif. 90007
CONTRIBUTIONS IN SCIENCE is a series of miscellaneous tech-
nical papers in the fields of Biology, Geology and Anthropology, published
at irregular intervals by the Los Angeles County Museum. Issues are num-
bered separately, and numbers run consecutively regardless of subject mat-
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David K. Caldwell
Editor
OBSERVATIONS ON CAPTIVE AND WILD ATLANTIC
BOTTLENOSED DOLPHINS, TURSIOPS TRUNCATUS, IN
THE NORTHEASTERN GULF OF MEXICO1
By Melba C. Caldwell,2 David K. Caldwell3 and J. B. Siebenaler4
Abstract: Data are presented concerning scouting behavior
by wild specimens; learning by observation by captive animals;
directional swimming of captives; evidence for stress-caused ul-
cers in captive specimens; ingestion of foreign objects by both
captive and wild cetaceans; seasonal micro-distribution of wild
animals; and a shark bite and infestations of the whale barnacle
Xenobalanus globicipitus on wild specimens. Included also are
additional records for the pigmy sperm whale, Kogia breviceps,
and Cuvier’s beaked whale, Ziphius cavirostris, in the north-
eastern Gulf of Mexico.
The present knowledge concerning the ecology and behavior of cetaceans
is still distressingly incomplete, even for a species as much studied as the Atlan-
tic bottlenosed dolphin, Tursiops truncatus (Montagu). Because we have ac-
cumulated a number of fragmentary observations on this species from ob-
servations in the wild and during the procedures of capture, training and sub-
sequent captive existence at Florida’s Gulfarium, Fort Walton Beach, which
seem to contribute to a better understanding of its biology, we take this op-
portunity to record them here. The observations are naturalistic and oppor-
tunistic, but they point the way in some cases for possible experimental pro-
cedures, and it may never be possible to investigate some of these behaviors
further in the laboratory under any conditions.
Observations on scouting behavior by wild tursiops truncatus: One in-
stance of scouting of a barrier in the open ocean has previously been reported
for Tursiops gilli Dali (Eberhardt and Evans, 1962:326; Evans and Dreher,
1962:220; Dreher and Evans, 1964:383). Their findings report the detach-
ment of a scout from a school. The scout made several echolocation runs on
an artificial barrier, each time returning to the group after the run. As the
barrier was passable at one end, the school finally avoided the barrier and
continued on its way.
1Partial support for certain phases of this study was received through grants from
the National Institute of Mental Health (MH-07509-01 ) and the National Science
Foundation (GB-1189).
2Research Associate, Los Angeles County Museum, also University of Southern Cali-
fornia, Antarctic Research Program.
3Curator of Ichthyology, Los Angeles County Museum: also Research Associate,
Florida State Museum, and Collaborator in Ichthyology, Institute of Jamaica.
4Curator and General Manager, Florida’s Gulfarium, Fort Walton Beach, Florida.
smiths)* hi) ft Bb:
>*LSTlTirP'; '
4
Contributions in Science
No. 91
Additional data on scouting behavior is available from the capture pro-
ceedings of Tursiops truncatus near Fort Walton Beach and Destin, Florida.
At one time the practice at the Gulfarium was to catch a school of dolphins in
one of the many dead-end bays in the area. These bays were ones with a nar-
row mouth and shallow water. A net was strung completely across the mouth
of the bay after a school of dolphins had entered. The second largest male
(sexes were determined after the animals were captured; individuals were
identified by marks and fin shapes), which came to be called by the fishermen
the “Lieutenant’’ detached himself from the group and scouted the net. He
then returned to the group. Following this, the largest male, closely followed
by a large female, charged the net. Sometimes this procedure was successful
and the group followed them through the opening, either a break in the net
caused by the charge, or a spot where the top of the net was pulled down by
the charge. Usually, however, the entire school could be captured. Only rare-
ly did an animal leap over the net. Also, in a former North Carolina fishery,
T. truncatus were taken commercially by net. It was said that they rarely
jumped the net when encircled, but if one did, the others followed (Clark,
1887:309). Net scouting behavior by T. truncatus during capture attempts on
the east coast of Florida was also suggested by remarks made in a popular ac-
count (Chapin, 1962:26). Present capture procedure at the Gulfarium in-
cludes encircling schools of dolphins either in open water or from the open
beach. Although the scouting behavior is not as easy to observe under these
circumstances, it often does occur.
Evidence for learning by observation by captive tursiops truncatus:
Learning by observation rarely has been shown in animal groups other than
man (Beach, 1947). Chimpanzees do, however, learn faster if permitted to
observe problem solving from experienced chimpanzees (Crawford and
Spence, 1939; Darby and Riopelle, 1959). Also, monkeys have learned a new
habit of washing sweet potatoes before eating by observing a young member
of their group perform this unusual behavior (Miyadi, 1959:859).
At the Gulfarium, naive animals are placed in tanks with trained animals.
These untrained animals are back-up performers for the trained dolphins, and
the training period for the back-up animals is considerably reduced by this
procedure.
One striking example of this learning by observation occurred at the Gulf-
arium. A dolphin was used to begin the show by raising a flag. This was done
by training the animal to leap, grasp and pull a ball suspended over the water.
This trained animal was sent away and a fresh animal was trained to perform
the act. However, the trainer erred slightly and the new dolphin was condi-
tioned in such a way that it learned to leap and raise the flag by striking the
ball with its snout, instead of grasping it in its teeth as it was supposed to do.
This animal later died, and another female, “Belinda’’ a tank mate and back-
up animal for the animal that was incorrectly trained, took over the act im-
mediately and without training. Furthermore, she performed the trick by the
1965
Behavioral Observations on Cetaceans
5
same incorrect procedure of striking the ball with her snout instead of grasp-
ing it. Belinda was then trained to perform the act correctly by seizing the ball
and this became a part of the show.
An additional prop, a flash camera, was subsequently installed on the
flag-raising apparatus. This new prop frightened Belinda, and she refused to
perform. However, a young male that had been in the tank for eight months
took over for the two performances that Belinda missed, flashing the camera
by seizing and pulling the ball. The young male’s previous training had con-
sisted solely of being taught to play basketball, an entirely unrelated trick.
After the two shows, Belinda resumed her old place in the act after her initial
fear of the new prop was abated.
Directional swimming by captive cetaceans: For a period of about 10
years, from 1954 until the winter of 1964, all of the newly-introduced T.
truncatus at the Gulfarium swam counter-clockwise, regardless of the direction
of the current flow in the tanks. Charles Emmett of the Aquarium of Niagara
Falls, New York, and Donald McSheehey of the Aquatarium, St. Petersburg,
Florida, also reported this same experience ( pers . conver., with Siebenaler).
However, out of seven animals captured in the winter of 1964, and observed
at the Gulfarium or by Marjorie Siebenaler elsewhere at Fort Walton Beach,
six swam clockwise and only one counter-clockwise. McSheehey (pers. con-
ver., 1965, with Siebenaler) also stated that, contrary to his past experience,
two dolphins captured on 1 April 1965 swam clockwise. We have no explana-
tion for this behavioral change as the locations of capture have not been
changed.
To further complicate the picture, all of the freshly-introduced T. trunca-
tus (from Florida) that we observed during studies at Marineland of the
Pacific, near Los Angeles, swam in a clockwise direction.
Through 1964, only counter-clockwise swimming was performed at the
Gulfarium by several captive spotted dolphins, Stenella plagiodon (Cope) , held
at different times, and by a captive adult male pigmy sperm whale, Kogia
breviceps (Blainville) , which stranded near Fort Walton Beach several years
ago. The latter animal apparently constitutes the sixth record for this species
in the Gulf of Mexico (see Caldwell, Inglis and Siebenaler, 1960).
Evidence for ulcers caused by stress in captive animals: Unusual psycho-
logical stress can have dire effects on captive dolphins. In the early period dur-
ing 1957 when the Gulfarium was open to the public but still not fully struc-
turally complete, the seawater system of the main dolphin tank was being re-
worked and additions being made. For several weeks the animals were sub-
jected to a constant barrage of pounding on the attached water pipes. The noise
was deafening to human divers working underwater in the tank at times when
this was happening. Consequently it must have been even more disturbing to
the sensitive auditory apparatus of the dolphins. Shortly thereafter, about six
of the dolphins died. Autopsy by a local veterinarian and a local medical doc-
tor, and corroborated by the Gulfarium biologists, revealed the probable cause
6
Contributions in Science
No. 91
of death of all of the dolphins to be duodenal ulcers. There was no question, at
least, that the ulcers were present and severe. Brown and Norris (1956:320)
cited a case of death in a captive Pacific common dolphin, Delphinus bairdi
Dali, due to a perforated gastric ulcer. They attributed the perforation to a
refusal to feed because of the death of another animal to which the dolphin
was closely attached. Brown and Norris also reported two cases of healed
gastric ulcers in two D. bairdi that were autopsied in the laboratory.
In the fall of 1964, an adult male T. truncatus died at the Gulfarium after
spending several years in captivity there; first arriving as a very young animal.
This animal became somewhat of a pet, but had undergone intermittent train-
ing for the show and at times had performed certain trained acts in the main
show tank. When not in the show tank, he was kept in a small training tank,
sometimes in isolation from other dolphins, sometimes not. When in the main
show tank, particularly during the last year of his life, he had received rather
rough treatment by the larger females. Upon autopsy, he was found to have
perforated gastric ulcers which, it was postulated, contributed to his death if
it was not the actual cause.
Ingestion of foreign objects by captive and wild cetaceans: Captive dol-
phins are prone to ingest all manner of inedible objects (Brown, et al., 1960).
Stomach contents of autopsied animals at the Gulfarium include gravel, metal
nuts, coins (including one as small as a dime), a lady’s broach, and plastic
stripping from the inside of the tanks. The stripping measured up to 18 inches
long and two to three inches wide.
This inclination to ingest such items as the plastic stripping led to a cata-
strophic mass mortality among the Gulfarium’s trained show animals during
the latter part of November, 1964. Through a series of unfortunate circum-
stances, two of the main show animals, an adult male and female, and an
adult female back-up animal all ingested large quantities of this stripping and
died. Balls of the plastic up to four inches in diameter were found impacted
in the first stomach of these animals. In one instance the stomach, normally
five to six inches in diameter, was distended to nearly 12 inches with the im-
pacted balls of plastic. Another animal, an adult female, was saved when the
trainers forced her to regurgitate the contents of her stomach after a liberal
dose of mineral oil. This animal regurgitated nearly ten quarts of the balls of
plastic and, in addition, an 8-inch lady’s comb.
Such seemingly perverse ingestion is not confined to captive dolphins.
Petit, Lomont and Theodorides (1956) found fragments of both marine and
terrestrial plants, including wood, foliage from a plantain, black poplar foli-
age, palm foliage and root stock (all in the total amount of 780 grams) im-
pacted in the anterior stomach of a wild specimen reported as Tursiops tursio
Fabr. Mr. Robert L. Brownell, Jr., stated (pers. conversation , 1965) that he
once found a piece of paper wadded into a two-inch ball, along with seaweed,
squid beaks and roundworms, in the stomach of a 5.5-foot male Pacific striped
dolphin, Lagenorhynchus obliquidens Gill. The animal had stranded alive at
1965
Behavioral Observations on Cetaceans
7
the Santa Monica pier, California, on 29 August 1963. A wide variety of in-
edible objects also has been recovered from the stomachs of wild sperm whales,
Physeter catodon Linnaeus; the list includes rocks, sand, a glass fishing buoy,
a coconut, wood, an apple, a shoe, a plastic bag and bailing wire (Caldwell,
Caldwell and Rice, 1965).
Seasonal micro-distribution of tursiops truncatus: While these dolphins
are present in the Fort Walton Beach— Destin area in Florida throughout the
year, there is a definite tendency for seasonal variation in microhabitat in that
region.
During the winter, from December to mid-March, these dolphins move
far up into the shallow bayous which constitute the headwaters of Choctawha-
tchee Bay. The waters of these bayous are fresh at these times, and the dol-
phins apparently are there feeding on small clupeid fishes that spawn there at
that time and on sea trout {Cy noscion) . During this period in the bayous, the
dolphins develop eruptions on their skin which are like those which develop
when these animals are kept captive in fresh water for several weeks. This
suggests, therefore, that the animals do not move in and out of the bayous into
salt water, but instead remain in them for extended periods.
During the rest of the year, Tursiops are found in the deep passes between
the bay and the open Gulf, and in and just behind the surf zone along the open
beach and up to a mile or so offshore.
There is almost no exception to these seasonal distributions, and dolphins
are absent from the bayous in summer and absent from the passes and beaches
in winter.
Infestations of the cosmopolitan obligate cetacean barnacle, xenobalanus
globicipitus Steenstrup: Although the list of hosts for this unusual barnacle is
long, we have found but one reference (Barnard, 1924:96) listing it from a
species of Tursiops . The host was reported as T. catalaniae (Gray) from
Natal.
A school of six adults and two young T. truncatus was captured on 30
June 1964 just off East Pass, near Destin. The two unweaned young, a 51 -inch
male and an 80-inch female, each with milk in their stomachs, had a massive
infestation of these barnacles attached along the entire posterior part of their
caudal flukes and a few of the barnacles attached near the distal tips of the
dorsal and both pectoral flippers. The barnacles on the caudal flukes were most
abundant on the dorsal surface, covering an area up to three inches from the
posterior edge of the flukes, as well as covering the posterior edge itself. None
of the adults from the school were infested.
Siebenaler has seen these barnacles on other T. truncatus, including
adults, caught in the vicinity of Destin over a period of nearly ten years. Mr.
F. G. Wood, presently of the Naval Ordinance Test Center at Point Mugu,
California, told us (in late 1964) that he had seen these barnacles on Atlantic
bottlenosed dolphins collected on the upper east coast of Florida. At Point
Mugu, through the courtesy of Dr. Sam H. Ridgway, we saw additional speci-
8
Contributions in Science
No. 91
Figure 1. Nearly-healed scar of shark bite on a living Atlantic bottlenosed dolphin,
Tursiops truncatus. See text for details.
mens of this barnacle taken from an adult T. Truncatus collected on 10 Octo-
ber 1963 at Gulfport, Mississippi. We were told that the infestation on this
animal was light.
A young, 12-foot, 8-inch female Cuvier’s beaked whale, Ziphius caviro-
stris Cuvier, that stranded alive near Fort Walton Beach on 10 December 1964
also had an infestation of these barnacles on the posterior upper surface of the
caudal flukes, just to the left of center. Apparently this is the first record of
this barnacle from this species of cetacean, and the record of the cetacean
itself is of interest as it is only the fourth for the Gulf of Mexico (see Gunter,
1954).
A TURSIOPS TRUNCATUS with a healed shark bite scar: A 6-foot, 3-inch
male dolphin captured during early July in water less than five feet deep just
off East Pass at Destin had a fresh but nearly-healed scar from the bite of a
large shark (Fig. 1 ) . In greatest dimension, the bite measured 10% inches wide
and 9% inches in gape, and was located on the left antero-dorsal side of the
body just anterior to a vertical from the left flipper and behind the blowhole.
While Gray (1964: 20) stated that Tursiops with evidence of shark bites are
not uncommon, one wonders how a shark could successfully attack such a
fast-moving and agile animal. However, most sharks are capable of strong
bursts of speed and the position of the wound in the case of the Destin dolphin
1965
Behavioral Observations on Cetaceans
9
suggests that the attacking shark may have been able to attack partly from
above and to the side of the dolphin and perhaps was thus able to briefly pin
the mammal against the shallow bottom or on one of the many shallow sandy
bars in the region where the dolphins are most often found.
With the aid of Dr. Shelton P. Applegate, Los Angeles County Museum,
an attempt was made to identify the attacking shark. By a process of elimina-
tion based on the size of the bite scar, the arrangement of the teeth and their
number and probable shape from the punctures, as well as the known habits
and distribution of Gulf of Mexico sharks (see Bigelow and Schroeder, 1948),
Dr. Appplegate concluded that the two most likely prospects are the great white
shark, Carcharodon carcharias (Linnaeus), and the bull shark, Carcharhinus
leucas (Muller and Henle). The bull shark is the most likely prospect of the
two because of its abundance and ecological requirements in the northern
Gulf of Mexico.
Literature Cited
Barnard, K. H.
1924. Contributions to the crustacean fauna of South Africa. Annals So.
African Mus., 20 (1): 1-103.
Beach, Frank A.
1947. Do they follow the leader? Natural History, 56(8) :356-359, 379-383.
Bigelow, Henry B., and William C. Schroeder
1948. Sharks. In John Tee-Van, et al., editors, Fishes of the western North
Atlantic. New Haven: Mem. Sears Foundation, Bingham Oceanogr.
Lab., 1(1): 59-546.
Brown, David H., and Kenneth S. Norris
1956. Observations of captive and wild cetaceans. J. Mammal., 37(3) : 3 1 1-
326.
Brown, David H., Rankin W. McIntyre, C. A. Delli Quadri and Robert J. Schroeder
1960. Health problems of captive dolphins and seals. J. Amer. Veterinary
Med. Assn., 137(9) :534-538.
Caldwell, David K., Melba C. Caldwell and Dale W. Rice
1965. Behavior of the sperm whale, Physeter catodon L. In K. S. Norris,
editor, Whales, dolphins, and porpoises. Proc. 1st Internatl. Symp. on
Cetacean Biol., Univ. Calif. Press. To appear in late 1965.
Caldwell, David K., Anthony Inglis and J. B. Siebenaler
1960. Sperm and pigmy sperm whales stranded in the Gulf of Mexico. J.
Mammal., 41(1) : 136-138.
10
Contributions in Science
No. 91
Chapin, Henry
1962. The remarkable dolphin & what makes him so. New York: Young
Scott Books, 96 p.
Clark, A. Howard
1887. The porpoise fishery. In G. Brown Goode, The fisheries and fishery
industries of the United States. Washington: U. S. Comm. Fish and
Fish., sec. V, vol. II, pt. XVI, pp. 308-310.
Crawford, H. P., and K. W. Spence
1939. Observational learning of discrimination problems of chimpanzees.
J. Comp. Psychol., 27:133-147.
Darby, C. L., and A. J. Riopelle
1959. Observational learning in the rhesus monkey. J. Comp. Physiol. Psy-
chol., 52:94-98.
Dreher, J. J., and William E. Evans
1964. Cetacean communication. In William N. Tavolga, editor, Marine Bio-
Acoustics. New York: Pergamon Press, pp. 373-393.
Eberhardt, Robert L., and William E. Evans
1962. Sound activity of the California gray whale, Eschrichtius glaucus. J.
Audio Engineering Soc., 10(4) : 324-328.
Evans, William E., and J. J. Dreher
1962. Observations on scouting behavior and associated sound production by
the Pacific bottlenosed porpoise ( Tursiops gilli Dali). Bull. So. Calif.
Acad. Sci., 61(4) :217-226.
Gray, William B.
1964. Porpoise tales. New York: A. S. Barnes and Co., 1 1 1 p.
Gunter, Gordon
1954. Mammals of the Gulf of Mexico. In Paul S. Galtsoff, coordinator, Gulf
of Mexico. Its origin, waters and marine life. U. S. Fish and Wildlife
Serv., Fish. Bull., 55(89) :543-551.
Miyadi, Denzaburo
1959. On some new habits and their propagation in Japanese monkey groups.
Proc. 15th Internatl. Congr. Zool., London (1958), pp. 857-860.
Petit, B., H. Lomont and J. Theodorides
1956. Contenu stomacal aberrant ayant provoque une obstruction intestinale
chez dauphin ( Tursiops tursio Fabr.). Vie et Milieu, 7(3) :422-424.
wrm
LOS
ANGELES
CONTRIBUTIONS
COUNTY
MUSEUM
IN SCIENCE
Dumber 92
April 4
THE BARSTOVIAN CAMP CREEK FAUNA
FROM ELKO COUNTY, NEVADA
By J. R. Macdonald
Los Angeles County Museum of Natural History • Exposition Park
Los Angeles, California 90007
CONTRIBUTIONS IN SCIENCE is a series of miscellaneous technical papers
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Editor
THE BARSTOVIAN CAMP CREEK FAUNA
FROM ELKO COUNTY, NEVADA
By J. R. Macdonald1
Abstract: The late Miocene Camp Creek fauna from the
Raine Ranch formation in Elko County, Nevada, is described. The
fauna includes Canids, Felids, Gomphotheriids, Equids, Came-
lids, and Cervids which are not determinate at the generic level.
In addition there is Parahippus sp., Protohippus sp., ?Brachycrus
sp., Hesperocamelus stylodon, Aepycamelus sp., and ?Merycodus
sp. The degree of hypsodonty in the cranium of a juvenile Proto-
hippus sp. suggests a post-Sheep Creek and pre-Barstow age for
the fauna.
Introduction
The fossil-bearing rocks at the junction of Susie Creek and Camp Creek,
about twenty-one miles northeast of Carlin, Nevada, have been mentioned
briefly in the literature several times since Sharp (1939) discussed the “Hum-
boldt formation” of northeastern Nevada. In this paper he refers to the locality
and its fossils (pp. 151-152) and states, “All of the material collected has been
placed in the hands of Dr. Chester Stock, of the department of geology, Cali-
fornia Institute of Technology, Pasadena, California!’ The Vertebrate Paleon-
tological Collections of the California Institute of Technology were purchased
by the Los Angeles County Museum of Natural History after Dr. Stock’s un-
timely death. Unfortunately, the Camp Creek material has not been found in
this collection and must be presumed lost.
Van Houten (1956), in his review of the Cenozoic sedimentary rocks of
Nevada, indicates the Camp Creek locality on two maps (figs. 1 and 2) but
only mentions the area in passing.
Lovejoy (1959) used the term “Camp Creek fauna” for the first time and
reported on a small suite of specimens which he had collected and given to The
American Museum of Natural History (AMNH numbers 45823 through
45827).
Regnier (1960) reviewed the geology of the Cenozoic rocks of the Car-
lin, Nevada, area and abandoned the use of the term “Humboldt formation!’
“. . . on account of its imprecision” (p. 1191). The sediments from which the
Camp Creek fauna has been collected are referred to the “upper member” (un-
named) of the Raine Ranch formation (pp. 1195 and 1197).
Collecting by the Los Angeles County Museum of Natural History in this
area was initiated in 1958 under the direction of Dr. Theodore Downs and
through the instigation of Mr. Timothy M. Doheny, who originally reported
to the Museum an occurrence of fossil vertebrates near his ranch in Nevada. It
Senior Curator of Vertebrate Paleontology, Los Angeles County Museum of Nat-
ural History.
1
2
Contributions in Science
No. 92
was through the support of Mr. Doheny and his friend and Museum benefac-
tor, Mr. Vernon Barrett, that two Museum field trips were made to the Camp
Creek area and environs in 1958 and 1959.
Acknowledgments
Acknowledgment is made to the members of the field parties who col-
lected the material described herein, to Mr. Timothy Doheny and Mr. Vernon
Barrett, who supported the work in the field, to Miss Mary Butler who pre-
pared the figures, to Miss Laurie Bryant for curatorial assistance, and to Mrs.
Eileen Macdonald for editing the manuscript.
Abbreviations which accompany specimen numbers have the following
meanings: LACM— Los Angeles County Museum of Natural History; AMNH
—American Museum of Natural History; FAM— Frick Laboratory; UCMP—
University of California Museum of Paleontology, Berkeley.
All specimen measurements are in millimeters.
FAUNAL LIST
Canidae, genus indeterminate
?Felidae, genus indeterminate
Gomphotheriidae, genus indeterminate
Parahippus sp.
Protohip pus sp.
Equidae, genus indeterminate
?Brachycrus sp.
Hesperocamelus styiodon Macdonald
Aepycamelus sp.
Camelidae, genus indeterminate
?Merycodus sp.
Cervidae, genus indeterminate.
AGE OF THE CAMP CREEK FAUNA
The genera of this fauna range from the early Miocene through the middle
Pliocene. Parahippus is known throughout the Miocene; Protohippus (as used
by Quinn, 1955), from middle Miocene to middle Pliocene; Brachycrus from
middle to late Miocene; Hesperocamelus styiodon heretofore only from the
early Pliocene but presumed to have earlier occurrence (Macdonald, 1949:
190); Aepycamelus extends from the middle Miocene through the early
Pliocene; and the long ranging Merycodus is found from the middle Miocene
through the middle Pliocene.
A close approximation of the age of the fauna is found only in the cranium
1966
Late Miocene Mammals from Nevada
3
of a juvenile horse which is herein referred to Protohippus sp. The crown
height of the M1 suggests that it is post-Sheep Creek and pre-Barstow in age—
an indication, therefore, of a mid-Barstovian (late Miocene) age for the fauna.
CARNIVORA Bowdich, 1821
Canidae Gray, 1821
Canidae, genus indeterminate
Referred specimens : LACM 5444, a jaw fragment; LACM 9327, distal
end of a metapodial; LACM 9328, distal end of a radius. LACM Locality
1535. 2
The fragment of the right mandible (5444) indicates an animal about the
size of the modern coyote. The roots of several premolars are preserved. The
general aspect suggests a canid with crowded but not overlapping or rotated
premolars.
The distal end of the metapodial (9327) suggests an animal slightly smaller
than a Great Dane.
The distal end of the radius (9328) is about the size of that of a coyote and
could be from the same individual as the jaw fragment.
Felidae Gray, 1821
?Felidae, genus indeterminate
Referred specimens'. LACM 5442, proximal phalanx. LACM Locality
1535.
This proximal phalanx is probably from a felid about the size of the mod-
ern mountain lion.
PROBOSCIDEA Illiger, 1811
Gomphotheriidae Cabrera, 1929
Gomphotheriidae, genus indeterminate
Referred specimen : LACM 5435, an enamel fragment. LACM Locality
1535.
This fragment of the enamel from a cusp is the only record of a probo-
scidean from this fauna.
PERISSODACTYLA Owen, 1848
Equidae Gray, 1821
Parahippus Leidy, 1858
Parahippus Leidy, 1858:26.
-Details as to precise localities are on file in the permanent records of the Section of
Vertebrate Paleontology, Los Angeles County Museum of Natural History.
4
Contributions in Science
No. 92
Parahippus, species indeterminate
Referred specimen : LACM 7355, an unworn P2. LACM Locality 1535.
This isolated tooth has very little enamel, but there is enough to show that
there was a more extensive enamel covering during life. The enamel is very
slightly crenulated; the anterior cingulum extends down and across the para-
lophid to a point near the base of the tooth below the protoconid; the posterior
cingulum extends from the hypoconulid to the base of the tooth at its postero-
labial corner, but does not extend across the base of the hypoconid; and the
metaconid and the metastylid are not separated.
This specimen is similar in size and shape to an isolated P2 (LACM (CIT)
1045) from the Sucker Creek fauna of Oregon which Scharf ( 1935:105-106)
referred to Parahippus avus (Marsh). However, the Sucker Creek specimen
has a well developed cingulum extending from the base of the protoconid
across the base of the hypoconid and up to the hypoconulid. Until more is
known about the variability of cinguli, it would not be prudent to suggest that
these specimens represent conspecific forms.
The tooth also seems close to Parahippus crenidens Scott (1893); how-
ever again the amount of variation in cement and cinguli must be better under-
stood before an assignment can be made.
Measurements of P2, Parahippus sp.
Greatest antero-posterior diameter 20.0 mm.
Greatest transverse diameter 12.5 mm.
Protohippus Leidy, 1858
Protohippus Leidy, 1858:26.
Protohippus, species indeterminate
Referred specimens : LACM 4254, cranium with I1-3, dP1-4, and M1,
scapula, humerus, radius, two tibiae, metacarpals II, III, and IV, and metatar-
sals II and III; LACM Locality 1286. LACM 5445, a lower cheek tooth and a
fragment of an upper cheek tooth, LACM Locality 1535. AMNH 45827, par-
tial cranium with L-M3, and questionably associated skeletal fragments.
The positive assignment of the juveniles to a species is always a question-
able procedure unless a large population has been collected from a single lo-
cality representing individuals of all ages. The cranium and partial skeleton of
the colt (LACM 4254) presents this difficulty. The facial characters of the in-
dividual change with growth, and the single permanent tooth is little better
than an isolated tooth for making an identification.
This colt has well cupped incisors which are just beginning to wear, the
deciduous premolars are heavily worn, the dP4 has only 7.6 mm. of crown re-
maining at the mesostyle, and the M1 is unworn. The cranium is nearly com-
plete and uncrushed, although the lacrymal fossae are probably deeper than in
1966
Late Miocene Mammals from Nevada
5
life due to collapse of the thin bone of the facial region. The left M1 has been
removed for measurement and sectioning.
The M1 was sectioned 10.1 mm. below the top of the mesostyle; the saw
blade had a thickness of 2.3 mm. The top of the cut has an open pre- and post-
fossette and an unconnected protocone. The crown surface of the lower sec-
tion, which represents the tooth at the one-third wear stage, has a well con-
nected protocone, closed fossettes, and a closed hypoconal groove.
The rapid connection of the protocone to the protoloph at an early stage of
wear and the simplicity of the enamel borders of the fossettes suggests this
form should be referred to Protohippus rather than Merychippus. Protohippus
Figure 1. Protohippus sp., LACM 4254, cranium, dorsal view. XV6.
Figure 2. Protohippus sp., LACM 4254, cranium, lateral view. XV3.
6
Contributions in Science
No. 92
is used as defined by Leidy (1858), redefined by Osborn (1918), and as resur-
rected by Quinn (1955).
A comparison with the protohippines at hand shows this form to have an
M1 somewhat larger than the type of Protohip pus tehachapiensis (Buwalda and
Lewis, 1955) from the Hemingfordian Phillips Ranch fauna, P. tehachapiensis
has a more open hypoconal groove, a broader connection of the protocone and
the protoloph at what appears to be about the half-wear stage, and a strong
arcate distortion of the fossettes.
The cheek teeth of Protohippus intermontanus (Merriam, 1915) from the
Barstow fauna are, in general, much more massive, larger, and higher crowned
than the M1 of the Camp Creek colt. The styles are more prominent and the
protocone has its long axis antero-posteriorly oriented instead of being slightly
on the oblique; the anterior cingulum is greatly reduced or lacking; and the
Figure 3. Protohippus sp., LACM 4254, cranium. A. Palatal view showing dP-dP4
and M1. XVi. B. Crown of sectioned M1. XV3.
enamel borders are somewhat more complex, but the plicaballin is at the same
stage of reduction. Merriam (1915:50-51) referred this species to Mery chip-
pus rather than Protohippus because of the small amount of cement on some
referred milk teeth. In 1919 he assigned the species to the subgenus Proto-
hippus.
Lacking mature material, the only realistic age assignment for this speci-
men is post-Sheep Creek and pre-Barstow, as the height of the crown of the M1
falls between the crown heights of species from these two faunas.
The partial cranium collected by Lovejoy in 1956 (AMNH 45827) and
the questionably associated skeletal fragments represent an old individual with
heavily worn dentition. Oblique crushing has greatly reduced the lacrymal
fossa on the left side and over-emphasized it on the right side. The incisors re-
tain their cups; the canines are worn flat at the tips; P2 is heavily worn; P2-4
1966
Late Miocene Mammals from Nevada
7
heavily worn, most of the cement gone from fossettes, fossette borders without
plications, protocone and hypocone remain separated; P3-4 show prehypoconal
groove, parastyle and mesostyle strongly developed; M1 with protocone and
hypocone connected, post protoconal valley forming lake; M2 similar to P4;
M3 showing oval protocone, hypoconal groove forming a lake.
Juvenile Protohippus, LACM 4254
Measurements and Analytical Characters (Downs, 1961).
Condylobasal length
273.5
Basal Length
258.0
Basilar length
254.0
Palitar length
124.3
Palital length
128.3
Length of tooth row
154.5
Length dP1-4
91.4
Post glenoid length
ca. 30.0
Interorbital constriction
51.4
Zygomatic breadth
110.9
M1:-
height of crown
34.9
shape of protocone
elongate oval
anteroposterior diameter of protocone
5.2
transverse diameter of protocone
ca. 3.7
development of cingulum
prominent
anteroposterior diameter of tooth
19.0
transverse diameter of tooth
17.9
tooth size
340.1
number of plications
1 (pli protoconule)
length of pli protoconule
ca. 1.1
tooth curvature
39.0
connection of protoconule and metaloph
closed
degree of tapering of crown
tr. dia. at base 20.8
tr. dia. crest 18.1
pli caballin
very weak
hypoconal groove
closed
Metacarpal III:
length
162.5
transverse diameter proximal end
21.1
Metatarsal III:
length
180.2
transverse diameter proximal end
20.2
transverse diameter collateral process
22.0
transverse diameter distal end
18.9
8
Contributions in Science
No. 92
The lightly worn isolated lower cheek tooth (LACM 5445) is heavily cov-
ered with cement, has a slight taper, and the metaconid and metastylid are sepa-
rated for at least half of the length of the tooth.
Equidae, genus indeterminate
Referred specimens : LACM 5371, a left astragalus, LACM Locality 1537.
LACM 5446, the distal end of a right tibia, a left astragalus, a left calcaneum,
two lateral proximal phalanges, a proximal phalanx, median phalanx, and a
right cuboid, LACM Locality 1535.
The isolated astragalus has an anteroposterior length of 44.1 mm. The
material from Locality 1535 was associated and may well belong to a single
individual. The tibial fragment has a transverse diameter at the distal end of
32.65 mm.; the length of the astragalus is 36.9 mm.; the calcaneum is broken
but fits the astragalus; the two lateral proximal phalanges are of a size to corre-
spond to the other elements; the proximal central phalanx is 38.0 mm. long;
the median central phalanx is worn but articulates with the proximal phalanx;
the right cuboid is 23.5 mm. long. This material is equivalent in size to the as-
sociated elements found with the LACM cranium of Protohippus sp. at Local-
ity 1286.
ARTIODACTYLA Owen, 1848
Merycoidodontidae Thorpe, 1923
Brachycrus Matthew, 1901
Brachycrus Matthew, 1901:397.
?Brachycrus, species indeterminate
Referred specimen : LACM 5370, a slightly worn P3. LACM Locality
1537.
This isolated tooth resembles the P3’s of Brachycrus buwaldi (Merriam)
and B. siouense (Sinclair) shown in figures 6 and 8 of Schultz and Falkenbach
(1940). The B. buwaldi specimen (FAM 34467) is from Green Hills, Bar-
stow, California, and the B. siouense specimen (FAM 36113) is from the
“Lower Snake Creek” deposits in Antelope Draw, Sioux County, Nebraska.
The variation shown in the premolars of Oreodonts precludes a definite
identification of this tooth. It is suggestive of this genus, and the age of the two
species which it resembles falls within the possible limits of the Camp Creek
fauna.
Camelidae Gray, 1821
Hesperocamelus Macdonald, 1949
Hesperocamelus Macdonald, 1949: 186.
1966
Late Miocene Mammals from Nevada
9
Hesperocamelus stylodon Macdonald, 1949
Hesperocamelus stylodon Macdonald, 1949:186.
Referred specimen : LACM 4318, skull and partial skeleton. LACM Lo-
cality 1535.
Figure 4. Hesperocamelus stylodon Macdonald, LACM 4318, cranium, lateral view.
X%.
This specimen is 17% greater in length but otherwise does not differ great-
ly from the type specimen (UCMP 35382) from the Chalk Spring fauna in
northern Elko County. The type specimen is without lower jaws, so this speci-
men adds some information to the dental characteristics of the species. The L
is missing, the I2 3 are spatulate, the lower canine is small, slightly recurved,
Figure 5. Hesperocamelus stylodon Macdonald, LACM 4318, cranium, palatal view
with L-M3. X3/s.
10
Contributions in Science
No. 92
and laterally compressed. The Pt is laterally compressed and very slightly re-
curved. P2 has the anterior minor cusp slightly hooked lingually, the major cusp
is low, the crest thickens posterad of the major cusp and the posterior border
is slightly inflected. P3 is similar to P2 but larger, the anterior and posterior in-
flections are more pronounced, there is a small lake on the crown within the
posterior inflection. The M12 are non-diagnostic. The M3 has a strongly cres-
centic talonid. This may or may not be characteristic of the species. Additional
material will be required to show whether the crescentic talonid of the M3 is
distinctive or a variable element as it is in most other camels.
The Age of Hesperocamelus styldon
When describing the Chalk Springs fauna from northeastern Elko Coun-
ty, which includes the type of H. stylodon, I assigned the entire fauna to the
Clarendonian age. Stratigraphically below the horizon of the type were frag-
ments of horse cheek teeth which I believed to be referable to Pliohippus.
Above the type horizon is a good Clarendonian fauna including Eucastor le-
contei (Merriam), Neohipparion near occidentale, and a Pliohippus showing
Figure 6. Hesperocamelus stylodon Macdonald, LACM 4318, left ramus with L,-M3,
lateral view. X 2/7.
In discussing the relationship of H. stylodon to the more advanced species,
H. alexandrae (Davidson) from the Barstow fauna (Macdonald, 1949:190),
I was concerned over the apparent occurrence of a primitive species ( H . stylo-
don) in a higher horizon. I assumed that H. stylodon represented a primitive
line that lingered on in time. The Camp Creek occurrence indicates a long tem-
poral span for this species and validates my earlier assumption. (Macdonald,
1949: fig. 11.)
1966
Late Miocene Mammals from Nevada
11
Table 1.
Measurements Hesperocamelus stylodon Macdonald
Length of cranium — pmx — occiput
Diastema, I1-2
Diastema I2-3
Diastema I3 — canine
Diastema canine — P 1
Diastema P1-2
UCMP 35382
(type)
321.5
LACM 4318
373.0
Diastema C-Pi
Diastema P. .
right
left
right
left
l3
113.7
116.4
132.5
130.7
alveolus I1
5.4
5.1
6.4
7.0
I2
4.5
4.5
5.2
5.2
7.6
7.2
9.6
7.8
I3
5.2
5.3
6.1
6.4
10.3
14.2
19.6
canine
5.0
5.1
ca. 6.5
16.7
ca. 19.5
Pi
6.5
6.8
ca. 8.7
7.5
10.5
13.4
11.7
P2
12.5
12.7
12.4
11.7
P3
16.3
16.2
15.6
15.9
Pi
15.6
15.9
16.4
16.2
Mi
20.0
20.6
24.6
24.8
M2
26.7
27.5
35.1
35.6
M3
ca. 34.0
31.4
37.5
37.5
/c
ca. 8.7
ca. 14.3
Pi
9.3
21.7
P2
11.9
Pa
14.7
P4
17.1
16.6
M1
21.1
M2
31.9
31.8
M3
44.6
46.5
Alveolar length canine — Mu
Alveolar length Pi-M3
Length P2-M3
Length Mr3
Length of tibia
Length of humerus
Length of metacarpal (crushed)
Length of metatarsal (crushed)
Length of calcaneum
Length of astragalus
Length of proximal phalanges
Length medial phalanx
360.0
369.0
373.5
ca.201.7
ca. 178.9
ca. 146.1
98.8 ca. 102.2
408.0
290.0
317.0
307.0
104.5
70.3
81.8
80.9
44.5
49.6
12
Contributions in Science
No. 92
Aepycamelus Macdonald, 1956
Aepycamelus Macdonald, 1956:198.
Aepycamelus sp.
Referred specimen : LACM 4095, mandibles and partial skeleton. LACM
Locality 1286.
The lower dentition resembles that of the paratype (LACM (CIT) 2819)
of Aepycamelus stocki (Henshaw) 1942 from the Tonopah fauna. The para-
type lacks the L-Pj which are present in the Camp Creek specimen. The in-
cisors are large and spatulate, the canine fairly massive, slightly recurved, and
with strongly developed lingually curving anterior flange. Pi is laterally com-
pressed with slight anterior and posterior blades. P2 is low crowned, unworn,
with a slight linguad curve to the anterior “cusp!’ P3 is slightly worn, the an-
terior end curves sharply lingually, at this stage of wear there is a double pos-
terad-lingual re-entrant which isolates a small lingual cusp and forms a spur on
the posterior border. The P4 is well worn, the anterior is hooked lingually at
about 30°, there is an elongated median posterior lake opening at the most
posterad point on the tooth through a narrow sulcus. The right P4 deeply in-
vades the anterior face of the M3. The lower molars are not distinctive. The
large anterior mental foramen is anterad to the root of P3, the small posterior
mental foramen is below the anterior root of M3.
A comparison of limb elements with the Tonopah material can only be
made with the metatarsals and the radioulna. The Camp Creek metatarsals are
about 10% longer and the radioulna about 10% shorter. This may be a signifi-
cant variation or may be normal within this group, study of a large sample from
a single fauna will be required before a decison can be made. There is one Tono-
pah radioulna (CIT 2823), which is only 443 mm. long, which Henshaw
(1942) referred to this species.
Figure 7. Hesperocamelus stylodon Macdonald, LACM 4318, lower jaws with I2-M3,
occlusal view. X 3/7.
1966
Late Miocene Mammals from Nevada
13
He believed that there was only one camel in the Tonopah fauna. It is pos-
sible that this specimen represents a small individual, but, in light of the great
diversity of camels found in most Nevada Miocene and Pliocene faunas, it is
presumptuous to assume a fauna to contain but one species.
Table 2.
Measurements for Aepycamelus sp.
Tonopah fauna
LACM CIT CIT CIT
4095 2819 2826 2824
A-P diameter Canine
Diastema, Canine— Pi
A-P diameter Pi
Diastema, Pi.2
Alveolar length, P2-M3
A-P diameter P2
A-P diameter P3
A-P diameter P4
A-P diameter Mi
A-P diameter M2
A-P diameter M3
Alveolar length, P2.4
Alveolar length, Mr3
Length, metacarpals III-IV
Width at head
A-P diameter at head
Width at distal end
Length, metatarsals III-IV
Width at head
A-P diameter at head
Width at distal end
Ulnoradius, length to semilunar notch
Prox. phalanx length
Right
Left
12.1
11.0
24.1
27.3
10.3
10.6
23.8
146.6
149.2
144.6
13.5
14.0
11.6
16.0
16.4
15.9
19.6
20.3
16.3
24.6
25.8
25.1
32.2
32.6
31.4
43.2
43.9
43.2
45.0
46.1
42.3
99.2
101.1
101.1
460.1
54.2
43.8
66.3
459.0
458.0
410.0
53.3
54.1
49.8
47.3
49.6
45.4
68.5
65.9
55.5
513.0
514.0
85.4
573.0
It is often desirable to review the characteristics used to separate various
genera within a family. The accompanying table may be useful to the student
and the curator. A review of papers dealing with camels by W. D. Matthew,
J. T. Gregory, and the writer does produce a group of characteristics which
may be used in distinguishing four common late Miocene and early Pliocene
camelid genera— Procamelus, Pliauchenia, Aepycamelus, and Hesperocamelus.
The accompanying table presents these characteristics.
14
Contributions in Science
No. 92
1966
Late Miocene Mammals from Nevada
15
Camelidae, genus indeterminate
Referred specimens:
Locality
Specimens
AMNH
AMNH
45823.
Pair of jaws with dP2-M2 and an atlas.
Camp Creek
AMNH
45824.
Astragalus.
AMNH
45825.
Jaw fragment and proximal phalanx.
AMNH
45826.
Three proximal phalanges; distal ends of two
LACM
LACM
5380.
metapodials; left trapezoid, cuneiform scaph-
oid, and magnum; and a right lunar.
Distal end of metapodial and proximal end
1533
LACM
5378.
of a proximal phalanx.
Left pisaform.
LACM
5377.
Left scaphoid.
LACM
5376.
Right navicular.
LACM
5379.
Right astragalus.
1535
LACM
5440.
Left scaphoid.
LACM
5438.
Right lunar and cuboid.
LACM
5436.
Right astragalus.
LACM
5441.
Distal ends of two metapodials and two
LACM
5439.
broken astragali.
Distal end of a proximal phalanx.
LACM
5374.
Distal end of a metapodial.
LACM
5375.
Distal end of a metapodial.
LACM
5373.
Right navicular.
LACM
5372.
Left scaphoid.
Discussion: This list of fragmentary camelid remains once again points to
the amazing abundance of camels during the late Miocene and early Pliocene
times in Nevada— or to the facility with which portions of these animals are
preserved. Most of this material is not in any way referable to the generic
level although a few specimens suggest either Hesperocamelus or Aepy-
camelus.
Hesperocamelus may be represented by the following specimens: AMNH
45825, a proximal phalanx 79.5 mm. in length, AMNH 45823, a pair of jaws
and associated atlas. The jaws retain the dP2-4 and the M3 is not erupted. They
are somewhat smaller than known specimens of Hesperocamelus.
16
Contributions in Science
No. 92
Table of Measurements
Greatest antero-posterior diameter
Right
Left
dp2
9.1
9.6
pp3
13.45
13.65
pp4
25.20
27.7
Mj
23.5
m2
26.2
LACM 5441, the distal ends of two metapodials and two broken astragali are
of a size to be referable to Hesperocamelus.
LACM 5439, a distal end of a proximal phalanx is Hesperocamelus size.
LACM 5374 and 5375, distal ends of two metapodials could be from the same
individual, one being a metacarpal and the other a metatarsal of a Hespero-
camelus.
Aepycamelus may be represented by the following: AMNH 45825, a jaw
fragment without teeth but of a size with LACM 4095 which has been referred
to this genus.
The remainder of the specimens showing camelid affinities represents the
usual puzzling array indicating a great variation in size but no solid foundation
for reference to known forms.
Antilocapridae Gray, 1866
Merycodus Leidy, 1854
Merycodus Leidy, 1854:90.
?Merycodus, species indeterminate
Referred specimens: LACM 4256, a fragment of the labial wall of an
M1 or M2, LACM Locality 1286. LACM 5369, a jaw fragment with heavily
worn P3 -4, LACM Locality 1537. LACM 5443, an astragalus, distal end of a
metapodial, and a medial phalanx, LACM Locality 1535.
This collection of fragments may be tentatively assigned to the genus
Merycodus. The jaw fragment with a broken Py and a heavily worn P4 could
well have been collected with the Tonopah or Barstow faunas. The fragment
of the labial wall of an upper molar indicates a higher crowned tooth and a
tooth with a greater anteroposterior diameter than the locally available
material from these faunas, but it would be well within possible limits of
variation. The distal end of the metapodial is larger than any of the available
specimens from late Miocene faunas, but it is matched in size by some of the
very early Clarendondian specimens from the Tejon Hills fauna.
1966
Late Miocene Mammals from Nevada
17
Cervidae Gray, 1821
Cervidae, genus indeterminate
Referred specimen: LACM 5436, the proximal end of a proximal phalanx,
LACM locality 1535.
This specimen has been broken away from the shaft just anterad of the
epiphysial joint. The articular surface is complete and unworn or weathered.
Its transverse diameter is 14.5 mm. and the height is 16.7 mm. The conforma-
tion of the articular surface resembles that of Dromomeryx cf. borealis from
the Skull Springs fauna of Oregon. It is smaller than the average specimen
from this fauna, but there are samples from Oregon which are essentially
from the same sized individuals.
CONCLUSIONS
Although this fauna is small by Great Plains standards, it is better than
the average Great Basin fauna. It gives a fairly valid age determination to the
Raine Ranch formation, post-Sheep Creek and pre-Barstow, and adds an
additional unit to the Tertiary faunas of Nevada.
Literature Cited
Buwalda, J. P., and G. E. Lewis
1955. A new species of Merychippus. U.S. Geol. Surv. Prof, paper 264-G:
147-152, 5 figs.
Downs, Theodore
1956. The Mascall fauna from the Miocene of Oregon. Univ. Calif. Publ.
Geol. Sci., 31(5) : 199-354, 50 figs., 8 pis.
Henshaw, P. C. *
1942. A Tertiary mammalian fauna from the San Antonio Mountains near
Tonopah, Nevada. Carnegie Inst. Wash. Publ. 530:77-168, 7 figs., 11 pis.
Leidy, Joseph.
1858. Notice of remains of extinct Vertebrata, from the valley of the Niobrara
River, collected during the exploring expedition of 1857, in Nebraska,
under the command of Lieut. G. K. Warren, U.S. Top. Eng., by Dr. F. V.
Hayden. Proc. Acad. Nat. Sci. Phila., 1858, pp. 20-29.
Lovejoy, D. W.
1959. Overthrust Ordovician and the Nannie’s Peak intrusive, Lone Mountain,
Elko County, Nevada. Bull. Geol. Soc. Amer., 70:539-564, 3 figs., 1 pi.
18
Contributions in Science
No. 92
Macdonald, J. R.
1949. A new Clarendondian fauna from northeastern Nevada. Univ. Calif.
PubL, Bull. Dept. Geol. Sci., 28:173-194, 11 figs.
Merriam, J. C.
1915. New horses from the Miocene and Pliocene of California. Univ. of Calif.
Pubis., Bull. Dept. Geol. Sci., 9:49-58, 12 figs.
1919. Tertiary mammalian faunas of the Mojave Desert. Univ. Calif. Pubis.,
Bull. Dept. Geol. Sci., 9:49-58, 12 figs.
Osborn, H. F.
1918. Equidae of the Oligocene, Miocene and Pliocene of North America;
iconographic type revision. Mem. Amer. Mus. Nat. Hist., new ser.,
2:1-330, 173 figs., 44 pis.
Quinn, J. H.
1955. Miocene Equidae of the Texas Gulf Coastal Plain, Univ. Texas. Bur.
Econ. Geol., Publ. no. 5516, 102 pp., 5 figs., 14 pis., 25 tables.
Regnier, Jerome
1960. Cenozoic Geology in the vicinity of Carlin, Nevada. Bull. Geol. Soc.
Amer., 71 : 1 189-1210, 3 figs., 2 pis.
Scharf, D. W.
1935. A Miocene mammalian fauna from Sucker Creek, southeastern Oregon.
Carnegie Inst. Wash. Publ. 453:97-118, 11 figs., 2 pis.
Schultz, C. B., and C. H. Falkenbach
1940. Merycochoerinae, a new subfamily of oreodonts. Bull. Amer. Mus. Nat.
Hist., 77:213-306, 18 figs.
Sharp, R. P.
1939. The Miocene Humboldt formation in northeastern Nevada. J. Geology,
47:133-160, 9 figs.
Van Houten, F. B.
1956. Reconnaissance of Cenozoic sedimentary rocks of Nevada. Bull. Amer.
Assoc. Petrol. Geol., 40:2801-2825.
LOS
ANGELES
COUNTY
MUSEUM
CONTRIBUTIONS
IN SCIENCE
UMBER 93
April 4
A KEY TO THE SPECIES OF OPHIUROIDEA
(BRITTLE STARS) OF THE SANTA MONICA BAY
AND ADJACENT AREAS
By Richard A. Boolootian and David Leighton
Los Angeles County Museum of Natural History
Los Angeles, California 90007
Exposition Park
CONTRIBUTIONS IN SCIENCE is a series of miscellaneous technical papers
in the fields of Biology, Geology and Anthropology, published at irregular intervals
by the Los Angeles County Museum of Natural History. Issues are numbered sepa-
rately, and numbers run consecutively regardless of subject matter. Number 1 was
issued January 23, 1957. The series is available to scientific institutions on an ex-
change basis. Copies may also be purchased at a nominal price.
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or to papers dealing largely with specimens in the Museum’s collections. Manuscripts
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MANUSCRIPT FORM. — (1) The 1960 AIBS Style Manual for Biological
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PROOF. — Author will be sent galley proof which should be corrected and re-
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David K. Caldwell
Editor
A KEY TO THE SPECIES OF OPHIUROIDEA
(BRITTLE STARS) OF THE SANTA MONICA BAY
AND ADJACENT AREAS1
By Richard A. Boolootian2 and David Leighton3
Abstract: Thirty ophiuroid species occur off the coast of
Southern California. The bathymetric range, color in life, habitat,
and meristic characteristics are considered. A dichotomous key
is presented.
Southern California ophiuroids are now well catalogued, although no
key to the species existing in any geographically distinct region of the California
shore and the continental shelf between La Jolla and Monterey has been pre-
viously published.
The pioneer work in the field of Pacific North American ophiuroids was
done by Lyman ( 1861 ) , who listed ten species and later increased the figure to
sixteen. Nine species were added to the list by Clark (1911). Neilsen’s ( 1932)
resume of the material collected during the Mortensen Pacific Expedition of
1914-1916 has been invaluable in the composition of this key.
Excellent work has been done on the Japanese ophiuroids by Matsumoto
(1917); species occurring in the Nanaimo district were listed by Berkeley
( 1927) ; those found in the Philippine seas were presented by Koehler ( 1922) .
For those species occurring along the North American coast, Neilsen (1932)
prepared a key considering the entire area from the Strait of Georgia to the
Gulf of Panama, and Busch (1918, 1921) a key to the ophiuroids of Friday
Harbor, Washington. Barnard and Ziesenhenne (1961) discussed the ophiu-
roid communities of Southern California coastal bottoms. The only works
which are locally applicable are the keys of McClendon (1909) for the San
Diego region and May (1924) for Monterey Bay. McClendon’s key is the
only one useful to investigators in Southern California.
Through the work of the investigators noted above, there are now 40
recognized species of ophiuroids from the North American Pacific coast.
Thirty species of ophiuroids are included in this key, ten of which may be
collected intertidally.
Materials used in this study were obtained by employing SCUBA for the
sub tidal forms. Some of the intertidal species were collected by the authors;
others were provided by Fred Ziesenhenne of the Allan Hancock Foundation,
University of Southern California.
In this key an attempt has been made to utilize ophiuroid characters which
are least subject to variation and which can be observed externally with a hand
lSupported by National Science Foundation Grant G-9561.
2Department of Zoology, University of California, Los Angeles; and Research Associate in Marine
Zoology, Los Angeles County Museum of Natural History.
sScripps Institution of Oceanography, La Jolla, California.
1
2
Contributions in Science
No. 93
Figure 1. Ophioderma panamense, diagnostic parts
1. oral arm plate 7. genital slit
2. angle of mouth 8. side arm plate
3. madreporite 9. tentacle pore
4. apex of jaw 10. tentacle scale
5. oral papilla 11. interbrachial area of disc
6. oral shield 12. arm spine
1966
Brittle Stars of Southern California
3
Figure 2. Two-fifths of oral aspect of a diagrammatic disc to show diagnostic parts
1. teeth
2. angle of mouth
3. adoral plate
4. tentacle scale
5. tentacular pit
6. oral papilla
7. oral shield
8. genital slit
9. interradial portion of disc
10. arm spine
11. 1st oral arm plate
12. side arm plate
4
Contributions in Science
No. 93
lens, requiring no dissection of material. Disc-arm ratios, general shape, color,
and other potentially ambiguous characters have been avoided.
Oral papillation is a fundamental key character, but whether enlarged oral
tentacle scales should be included in the number of oral papillae per jaw in
all cases is questionable. Where these structures are obvious, they have been
included (see Ophionereis annulata ). Together with the key we include a table
indicating where the specimens may be found (Table 1), as well as a photo-
graph (Fig. 1) and a diagram showing general diagnostic features (Fig. 2).
An illustration showing the details of the oral papillae is included for each
species.
The key is in no way a natural one, though for the most part, related genera
fall closely together.
KEY
I. Both disc and arms covered by a leathery skin; aboral arm plates absent or
rudimentary; arms branched (Fig. 3) Gorgonocephalus eucnemis
II. Arms never covered by a thickened skin; aboral arm plates present; arms
never branched.
A. Aboral disc scaled, though scales may be discontinuous.
1. Oral papillae six or less than six per jaw.
a. Oral papillae two to four (rarely five) per jaw.
( 1 ) . Individuals often six-rayed; oral papillae blunt.
(a) . Radial shields small, never joining with mate; four smooth
spines on each side arm plate; two oral papillae per jaw (Fig. 4) .
Ophiactis simplex *
(b) . Radial shields large; mates joining distally; five (rarely six)
spines with fine serration on each side arm plate; four or five
oral papillae per jaw (Fig. 5) Ophiactis savignyi*
(2) . Individuals never six-rayed; oral papillae sharp, numbering two
or three per jaw; one apical or subapical and two (occasionally three)
distal oral papillae.
(a) . One tentacle scale; disc strongly scaled (Fig. 6)
Amphiura diastata
(b) . Two tentacle scales; disc occasionally not scaled centrally
(Fig. 7) Amphiura arcystata
b. Oral papillae six per jaw; three or occasionally four spines per side
arm plate.
(1). Two proximal pairs of oral papillae small; distal pair broad and
elongate.
(a). Interbrachial areas granular; radial shields separate or
meeting only distally (Fig. 8). ... A mphichondrius granulosus
1966
Brittle Stars of Southern California
5
(b). Interbrachial areas scaled; radial shields in solid contact.
i. Longest arm spines about IV2 times length of arm joint;
arms markedly long and narrow (Fig. 9)
Amphipholis pugetana *
ii. Longest arm spines about 1 arm joint in length; arms
relatively short (about four times the disc) (Fig. 10). ...
Amphipholis squamata*
(2) . Oral papillae all subequal in size and shape.
(a) . Some of the disc scales with free ends prolonged into fine
points.
i. Scales of aboral disc few and large (Fig. 11)
Amphiodia (Amphispina) digitata
ii. Scales of aboral disc numerous and small (Fig. 12). . .
Amphiodia (Amphispina) urtica
(b) . Disc scales never prolonged into fine points.
i. Disc with a rosette of large scales aborally; tentacle
scales (2) unequal in size; plates about mouth inflated
(Fig. 13) Amphiodia psara
ii. Disc with fine scales; tentacle scales (2) equal in size;
plates about mouth not inflated (Fig. 14)
Amphiodia occidentalis
2. Oral papillae more than six per jaw.
a. Eight oral papillae per jaw (rarely nine) .
(1) . Spines on disc partially covering scales; oral papillae spinose
and globose (Fig. 15) Amphiacantha amphacantha
(2) . No spines present on disc; most oral papillae heavy though a
few are terete. Two tentacle scales in angle of mouth often consid-
ered to be oral papillae ( 10) .
(a) . Tentacle scales in angle of mouth separate from true oral
papillae row; proximal oral papillae heavy and globose; other
oral papillae heavy but tapered (Fig. 16)
Amphioplus strongyloplax
(b) . Tentacle scales in angle of mouth closely adjacent to row
of true oral papillae; oral papillae tapered and not heavy (Fig.
17) Amphioplus hexacanthus
b. Nine or more than nine oral papillae per jaw.
(1). Oral papillae nine to ten; those in angle of mouth curved and
pointed (actually tentacle scales). Tentacle scales large and saucer
shaped; three arm spines on each side arm plate.
(a). Aboral arm plate large; accessory plates very small. Disc
with scattered large scales of lighter pigmentation; arms mot-
tled brown and cream (Fig. 18). . Ophionereis eurybrachyplax
6
Contributions in Science
No. 93
(b).Aboral arm plates equaled in size by accessory plates;
light spots scattered on disc incorporating several small scales;
arms banded (Fig. 19) Ophionereis annulata*
(2). Oral papillae more than ten per jaw; tentacle scales often more
than one, neither large nor saucer shaped.
(a) . Arm spines sharp, about one arm joint in length; small
notches in disc above arm base edged with small papillae;
symmetrical scale situated centrally on aboral disc (Fig. 20).
Ophiura lutkeni
(b) . Arm spines not sharp and considerably less than one arm
joint in length; disc notches and symmetrical scale absent; oral
papillae in even rov/s.
i. Oral papillae partially fused; tentacle pores only on first
three oral arm plates; aboral arm plates not divided (Fig.
21) Ophiomusium jolliensis
ii. Oral papillae not fused; aboral arm plates divided into
many smaller plates; arms flattened (Fig. 22)
Ophioplocus esmarki*
B. Scales or plates of aboral disc covered or partially obscured by superficial
structures.
1 . Disc covered by a thickened epidermis.
a. Velvet-like epidermis covering disc; oral papillae and arm spines
small and numerous; adults often over twelve inches in diameter (Fig.
23) Ophioderma panamense*
b. Smooth or parchment-like epidermis covering disc in interradial
areas; arm spines long, flattened, narrower at base than at end; tentacle
scales similar to arm spines and usually held in crossed position on oral
surface of arm (Fig. 24) Ophiopsila calif ornica
2. Disc covered with spines or short stumps.
a. Spines of arms held normally to arm axis (unless improperly pre-
served).
( 1 ) . Arm spines heavy and flattened; low rounded stumps cover disc;
dorsal-most arm spine very short; dental papillae numerous (Fig.
25) Ophiopteris papillosa*
(2) . Arm spines rather light and delicate; no oral papillae; disc
covered by short spines.
(a) . Arm and disc spines serrated; seven arm spines on each
side arm plate (Fig. 26) Ophiothrix spiculata*
(b) . Arm and disc spines rather smooth; five or six arm spines
on each side arm plate (Fig. 27) Ophiothrix rudis *
b. Arm spines form small angles with arm axis.
1966
Brittle Stars of Southern California
7
(1) .Arm spines short and blunt; disc fairly heavily covered with
branched spines; small supplementary plates partially surround
aboral arm plates (Fig. 28) Ophiopholis bakeri
(2) . Arm spines rather long and tapered; side arm plates nearly or
completely meeting above and below; granules cover most of disc.
(a) . Oral papillae twelve to fourteen per jaw; some fine scales
in evidence on disc.
i. Spines of considerable size scattered on aboral disc;
shorter stumps and granules cover most of balance of disc;
oral arm plates well separated by side arm plates; longest
arm spine about three arm joints in length (Fig. 29)
Ophiacantha phragma
ii. Small granules almost completely hiding scales of disc;
oral arm plates not widely separated by side arm plates;
longest arm spines about five arm joints in length (Fig. 30) .
Ophiacantha diplasia
(b) . Oral papillae seven to nine per jaw; short spines with fine
points cover disc.
i. Longest arm spines about two arm joints in length;
stumps on disc drawn out to fine (single) points; tentacle
scales conical (few scales may show on disc) (Fig. 31).
Ophiacantha normani
ii. Longest arm spines about four arm joints in length;
disc with short multi-fid spines; tentacle scales not conical;
arm spines serrated (Fig. 32). . Ophiacantha rhachophora
* Specimens collected intertidally
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Brittle Stars of Southern California
11
DEFINITIONS OF TERMS
Aboral: side opposite the mouth; the dorsal aspect of the animal.
Aboral arm plates: superficial plates covering the dorsal portion of each arm joint.
Aboral plates : shields or plates situated on either side of an oral shield.
Angle of mouth: the distal portion of the slit formed by approximation of any two
adjacent jaws.
Disc: the central body of an ophiuroid which is sharply marked off from the arms.
Distal: occupying a position away from the mouth or away from the center of the
disc.
Genital scales: scales, usually in orderly rows, bordering the genital slits.
Genital slits: slits located interbrachially and orally on the disc (on either side of
each arm base) indicating the position of the genital bursae.
Interbrachial areas: the oral disc lying between adjacent arms.
Jaws: five (or rarely six) triangular structures surrounding the mouth and usually
bearing a number of oral papillae laterally and a vertical row of teeth apically.
Oral: the ventral surface as opposed to the aboral or dorsal surface; implying direc-
tion toward the mouth or on the same surface as the mouth.
Oral arm plates: those plates situated on the ventral surface of the arm joint through
which pass the podia.
Oral papillae: modified spines usually found on the sides of each jaw and bordering
the angle of the mouth.
Oral shield: a plate, usually comparatively large, situated on the mid-interbrachial
line at the base of each jaw.
Podia: tube feet projecting through the tentacle pores of the oral arm plates.
Proximal: toward the oral-aboral axis; opposed to distal.
Radial shields: plates, often large, existing in pairs and located on or approaching
the radius of the aboral disc.
Radius: an imaginary line drawn from the center of the disc to any arm tip.
Side arm plates: those plates covering the lateral aspect of each arm joint and sup-
porting the arm spines.
Tentacle pores: a pair of openings in the oral arm plate through which pass the podia
or tentacles.
Tentacle scales: scales found bordering the tentacle pores which, in some species,
completely close the tentacle pore.
Tooth papillae: small papillae lying ventrally and about the teeth on the axis of the
jaw. (Found in relatively few of the species considered in this key.)
12
Contributions in Science
No. 93
Literature Cited
Barnard, J. L., and F. C. Ziesenhenne
1961. Ophiuroid communities of Southern California coastal bottoms. Pacific
Naturalist, 2:131-152.
Berkeley, Alfreda
1927. A preliminary list of the ophiurans of the Nanaimo District. Cont. to
Canadian Biol, and Fisheries, 3:319-322.
Busch, Mildred
1918. A key to the ophiuroids of Friday Harbor, Washington. Publ. Puget
Sound Biol. Sta., 2:17-44.
1921. Revised key to the echinoderms of Friday Harbor. Publ. Puget Sound
Biol. Sta., 3:65-77.
Clark, H. L.
1911. North Pacific ophiurans in the collection of the United States National
Museum. Bull. U. S. Natl. Mus., 75:1-302.
Koehler, R.
1922. Ophiurans of the Philippine Seas and adjacent waters. Bull. U. S.
Natl. Mus., 100(5) : 1-480, pis. 1-103.
Lyman, Theodore
1861. Descriptions of new Ophiuridae, belonging to the Smithsonian Institu-
tion and to the Museum of Comparative Zoology at Cambridge. Proc.
Boston Soc. Nat. Hist., 7:193-205, 252-262.
McClendon, J. F.
1909. The ophiurans of the San Diego region. Univ. Calif. Publ. Zool.,
6:33-64.
Matsumoto, H.
1917. A monograph of Japanese ophiuroidea, arranged according to a new
classification. J. Coll. Sci., Imp. Univ. Tokyo, 38(2) : 1-407, 7 pis.
May, R. M.
1924. Ophiurans of Monterey Bay. Proc. Calif. Acad. Sci., ser. 4, 13:261-303.
Nielsen, E.
1932. Papers from Dr. Th. Mortensen’s Pacific Expedition, 1914-16. LIX.
Ophiurans from the Gulf of Panama, California, and the Strait of
Georgia. Videnskabelige Meddelelser fra Dansk naturhistorisk Foren-
ing, 91:241-346.
1
1966
Brittle Stars of Southern California
13
Figure 3. Gorgonocephalus eucnemis. Figure 4. Ophiactis simplex.
Figure 5. Ophiactis savignyi.
Figure 6. Amphiura diastata.
14
Contributions in Science
No. 93
Figure 7. Amphiura arcystata. Figure 8. Amphichondrius granulosus.
Figure 9. Amphipholis pugetana. Figure 10. Amphipholis squamata.
1966
Brittle Stars of Southern California
15
Figure 11. Amphiodia digitata.
Figure 13. Amphiodia psara.
Figure 12. Amphiodia urtica.
Figure 14. Amphiodia occidentalis.
16
Contributions in Science
No. 93
Figure 15. Amphiacantha amphacantha. Figure 16. Amphioplus strongyloplax.
Figure 17. Amphioplus hexacanthus. Figure 18. Ophionereis eurybrachyplax.
1966
Brittle Stars of Southern California
17
Figure 19. Ophionereis annulata.
Figure 20. Ophiura lutkeni.
Figure 21. Ophiomusium jolliensis. Figure 22. Ophioplocus esmarki.
18
Contributions in Science
No. 93
Figure 23. Ophioderma panamense.
Figure 24. Ophiopsila californica.
Figure 25. Ophiopteris papillosa.
Figure 26. Ophiothrix spiculata.
1966
Brittle Stars of Southern California
19
Figure 27 . Ophiothrix rudis.
Figure 28. Ophiopholis bakeri.
Figure 29. Ophiacantha phragma. Figure 30. Ophiacantha diplasia.
20
Contributions in Science
No. 93
Figure 31. Ophiacantha normani. Figure 32. Ophiacantha rhachophora.
angeles CONTRIBUTIONS
= IN SCIENCE
mber 94 April 4
Vjl
PLIOCENE BIRDS FROM CHIHUAHUA, MEXICO
By
Hildegarde Howard
Los Angeles County Museum of Natural History • Exposition Park
Los Angeles, California 90007
CONTRIBUTIONS IN SCIENCE is a series of miscellaneous technical papers
in the fields of Biology, Geology and Anthropology, published at irregular intervals
by the Los Angeles County Museum of Natural History. Issues are numbered sepa-
rately, and numbers run consecutively regardless of subject matter. Number 1 was
issued January 23, 1957. The series is available to scientific institutions on an ex-
change basis. Copies may also be purchased at a nominal price.
INSTRUCTIONS FOR AUTHORS
Manuscripts for the LOS ANGELES COUNTY MUSEUM CONTRIBU-
TIONS IN SCIENCE may be in any field of Life or Earth Sciences. Acceptance of
papers will be determined by the amount and character of new information and the
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or to papers dealing largely with specimens in the Museum’s collections. Manuscripts
must conform to CONTRIBUTIONS style and will be examined for suitability by
an Editorial Committee. They may also be subject to critical review by competent
specialists.
MANUSCRIPT FORM.— (1) The 1960 AIBS Style Manual for Biological
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of literature citation must conform to CONTRIBUTIONS style — see number 90 and
later issues. Spell out in full the title of non-English serials and places of publication.
(7) A factual summary is recommended for longer papers. (8) A brief abstract must
be included for all papers. This will be published at the head of each paper.
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Editor
PLIOCENE BIRDS FROM CHIHUAHUA, MEXICO
By Hildegarde Howard1
Abstract: Seven birds from the Yepomera fauna of the Rio
Papigochic valley, Mexico are discussed, only one of which had
been previously recorded from the area. All appear to be extinct,
and one is described as a new genus and species.
The Pliocene deposits exposed in the valley of the Rio Papigochic, western
Chihuahua, Mexico, have received considerable attention from geologists and
paleontologists. A discussion of the deposits, with map, list of mammals
recorded, and large bibliography has been presented by Lance (1950).
The area yielding the most fossils centers around the small towns of
Rincon de la Concha and Yepomera. Although several collecting localities are
involved, the mammalian fauna is more or less uniform (Lance, 1950:7) and
is known as the Yepomera Fauna (formerly Rincon Fauna). The age is con-
sidered to be middle Pliocene (Hemphillian) .
The California Institute of Technology (CIT) collected extensively in the
Rincon- Yepomera area from at least ten separate fossiliferous localities. Bird
bones were recovered at three of these: Arroyo de los Burros (CIT locality
276), Arroyo de las Barrancas Blancas (CIT locality 286) and Arroyo de los
Ponos (CIT locality 289). All California Institute of Technology material is
now part of the collections of the Los Angeles County Museum (LACM).
The only avian species so far recorded from the Yepomera Fauna is a
small flamingo, Phoenicopterus stocki, described from locality 289, with a
total of nine bones referred, including the type (Miller, 1944). Nine more
bird bones from locality 289, two from 276, and three from 286 are included
in the collections. The list of avian species as now recognized includes seven
species, as follows:
Avian Species in the Yepomera Fauna Number of
Locality 276 specimens
Phoenicopterus stocki (flamingo) 1
Erolia (?), sp. (small shorebird) 1
Locality 286
Wasonaka yepomerae, n. gen., n. sp. of duck 3
Locality 289
Phoenicopterus stocki 12
Eremochen cf. russelli (goose) 1
Oxyura, sp. (duck) 3
Anas bunkeri (teal) 1
Mimidae (?) (thrasher-like bird) 1
"23
1Research Associate in Vertebrate Paleontology, Los Angeles County Museum of
Natural History.
1
2
Contributions in Science
No. 94
1966
Pliocene Birds from Chihuahua, Mexico
3
With the exception of the one passerine, all of the avian species are
aquatic, and fit well into the picture of ephemeral lakes or marshes such as
suggested by Lance (1950:8). The representation is too small to be of any
significance as an age indicator. Probably all of the species are extinct, al-
though the shorebird and the passerine cannot be definitely determined. The
two extinct forms that have been previously recorded, Eremochen russelli and
Anas bunkeri are both typically Pliocene, the former lower Pliocene, the
latter middle Pliocene into late Blancan (early Pleistocene).
DESCRIPTION OF MATERIAL
Phoenicopterus stocki
Except for the type tibiotarsus, the specimens previously referred to this
small species (Miller, 1944) were not individually listed. They have since
been catalogued, and are as follows: right and left distal ends of tibiotarsus,
LACM 4624 and 4626, proximal end of tibiotarsus figured by Miller
(1944:78), LACM 4623; two left distal ends of humeri discussed by Miller
(1944:80), LACM 4629 and 4630; proximal fragments of left ulna, LACM
4627, and left carpometacarpus, LACM 4628; and distal fragment of left
tarsometatarsus, LACM 4625. To these may now be added a fragment of
scapula (LACM 9731), and proximal and distal ends of radius (LACM
9732-9733) from the type locality (loc. 289), and a right distal end of
humerus (LACM 4616) from locality 276, all of which are smaller than
comparable elements of six Recent specimens of P. ruber. The humerus
(4616) falls between the two previously recorded specimens of this element
of P. stocki in size.
Measurements compared with Recent specimens of P. ruber :
Humerus, breadth distal end, P. ruber, 21.6-24.9 mm.
LACM 4630, 19.2 mm.
LACM 4616, 20.0 mm.
LACM 4629, 21.9 mm.
Ulna, breadth proximal end, P. ruber, 14.8-16.3 mm.
LACM 4527, 13.6 mm.
Radius, breadth distal end, P. ruber, 9.7-10.6 mm.
LACM 9733, 8.7 mm.
Radius, minimum and maximum dimensions of proximal end, P. ruber 6.1
mm. x 8. 1 mm. — 6.8 mm. x 8.6 mm.
4
Figure 1. A — E, Wasonaka yepomerae, n. gen., n. sp.: A-B, type humerus, LACM
4620, anconal and palmar views; C, D, E, paratype ulna, LACM 4619, external,
palmar and internal views. F — G, Anas bunkeri Wetmore, referred coracoid, LACM
4621, anterior and posterior views; H — I, Wasonaka yepomerae, paratype furcula,
LACM 4618, posterior and anterior views; J, Eremochen cf. russelli Brodkorb,
referred scapula, LACM 9734, dorsal view. x 1.
Photos by George Brauer.
4
Contributions in Science
No. 94
LACM 9732, 5.9 mm x 7.3 mm.
Carpometacarpus, breadth proximal trochlea, P. ruber, 7. 7-8. 7 mm.
LACM 4628, 7.4 mm.
Scapula, breadth from glenoid facet to shaft posterior to acromion, P. ruber,
1 1.0-12.6 mm.
LACM 9731, 10.4 mm.
Anseriformes
Anserinae
Eremoehen cf. russelli
Fig. l.J
A scapula (LACM 9734) from locality 289, with well-marked forward
projection of the acromion, resembles this element in the geese except for the
large dorsal pneumatic fossa adjacent to the coracoidal facet. In size the
specimen approximates scapulae of Recent A user hyperboreus. The coracoidal
facet is, however, smaller, and in place of the dorsal fossa there is a very
slight depression. Although there is great variability in this element among
Recent genera of geese, in none is the pneumatic fossa so subordinated.
Eremoehen russelli, from the lower Pliocene of Malheur County, Oregon,
is based on a proximal end of humerus as type, with scapula, carpometacarpus
and tibiotarsus referred (Brodkorb, 1961:175-176). The scapula has been
loaned for this study through the courtesy of Dr. J. A. Shotwell of the
Museum of Natural History, University of Oregon. As in the Mexican
scapula, this element of Eremoehen is nonpneumatic in the area of the cora-
coidal facet; there is even less indication of a depression than in the Mexican
fossil. In size, angularity of the anterior tip of the glenoid facet, and develop-
ment of the muscle scar posterior to the acromion, ventrally, the Oregon and
Mexican scapulae are markedly alike. The coracoidal facet appears slightly
larger in the former, and, possibly, the acromion is more laterally and dorsally
developed (unfortunately the tip of the acromion is broken away in the
Oregon scapula so, its proximal extent cannot be determined). Considering
the great variability that may be found in this element among individuals of a
species in living geese, the notable similarities between the Mexican scapula
and that of Eremoehen russelli, leave no alternative but to allocate the Mexican
fossil to the genus Eremoehen. In view of the deviations in some characters,
and the slight age difference, the species assignment is tentative.
Anatinae
Three species of ducks are included in the Yepomera avifauna. The one
species recovered from locality 286 is represented by a furcula, humerus and
ulna, probably all of one individual. The bones suggest a duck of about the
size of a mallard (Anus platyrhynchos) but with more slender wings. Rela-
tionship is perhaps closer to the perching ducks (tribe Cairinini) than to the
1966
Pliocene Birds from Chihuahua, Mexico
5
dabblers (Anatini). Woolfenden (1961:109) found the osteological charac-
ters of these tribes so similar that he recommended grouping them together.
However, at least between Anas (including species formerly allocated to
Chaulelasmus, Spatula, Mareca, Nettion and Querquedula) , and the cairinine
genera Aix, Sarkidiornis and Cairina, there appear to be a few recognizable
differences in the humerus and furcula. The fossil humerus combines many
of the characters found in Anas with others that seem to be more typical of
Aix or Sarkidiornis. The furcula is even more similar to that of the perchers,
although, at the same time being quite distinctive. The new genus and species
herein described is, therefore, tentatively allocated to the Cairinini. For con-
venience of identification, however, Anas platyrhynchos is used as a basis of
comparison in the detailed description.
Wasonaka, new genus
Type: Wasonaka yepomerae, new species.
Generic diagnosis: Humerus with attachment of external head of triceps
muscle depressed and distinctly bordered below head by curved line terminat-
ing at median edge of long, oval pectoral scar; pectoral scar appressed to shaft
at proximal edge of deltoid crest; deltoid crest outwardly flared, and external
surface excavated below heavy proximal border; on palmar surface, bicipital
furrow markedly depressed externally at base of prominent proximal border
of deltoid crest; distal edge of bicipital crest well above level of termination
of deltoid crest; distally, impression of brachialis anticus muscle a small oval;
entepicondylar process prominent and tending to overhang attachment of
pronator longus muscle. Furcula relatively straight and V-shaped, with practi-
cally no posterior flexure of symphysis; symphysis thickened anteroposteriorly,
lacking lines or depression anteriorly, bearing blunt furcular process posterior-
ly; dorsal surface of symphysis visible in posterior view, forming angular
junction with posterior surface; coracoidal tuberosity of clavicle distinct, but
small and papilla-like.
The generic name is derived from wasona-ka, meaning “duck” in the lan-
guage of the Tarahumar Indians of Chihuahua, Mexico.
Wasonaka yepomerae, new species
Fig. 1, A-E and H-I
Type: Right humerus complete except for broken edges of bicipital and
deltoid crests, internal tuberosity and anconal surface of external side of
distal end; LACM 4620: collected by California Institute of Technology field
party in 1946.
Locality and age: LACM (CIT) locality 286: Arroyo de las Barrancas
Blancas, !4 mile east of town of Yepomera, state of Chihuahua, Mexico;
middle Pliocene (Hemphillian) .
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Contributions in Science
No. 94
Paratypes: Furcula (LACM 4618) lacking tips of both clavicles; and
left ulna, complete (LACM 4619); both bones found associated with type
humerus at locality 286.
Diagnosis: See generic diagnosis.
Detailed description: Humerus (fig. 1, A-B) resembling that of Anas
platyrhynchos in (1) length, (2) distinctness and extent of attachment of
external head of triceps muscle, (3) long, oval pectoral scar appressed to
shaft, (4) small, oval impression of brachialis anticus muscle, (5) angular
apex of palmar surface of shaft above distal condyles, terminating at median
edge of impression of brachialis anticus and about on a longitudinal line with
center of external condyle, (6) attachment of anterior articular ligament
facing slightly distally, and contour angular in internal profile; distinguished
from A. platyrhynchos by (1) shaft more slender and more curved,
(2) anconal contour of head more evenly rounded, with less overhang over
capital groove, (3) groove narrower, (4) deltoid crest flared and connecting
with shaft (at its distal tip) at an abrupt angle, (5) external surface of deltoid
crest excavated, with (6) heavy proximal border undercut on palmar side by
deep bicipital furrow, (7) bicipital crest shorter relative to length of deltoid
crest, (8) intermuscular line distal to bicipital crest at extreme edge of shaft,
(9) entepicondylar prominence more pronounced and tending to overhang
attachment of pronator longus muscle, (10) internal condyle more round,
less oval, and tending to constrict intercondylar groove toward anconal side.
Furcula (fig. 1, H-I ) V-shaped as in some species of Anas. Distinguished
from this element in Anas as follows: coracoidal tuberosity smaller and
papilla-like; symphysis thickened and less flexed posteriorly; junction of dorsal
and posterior surfaces angular and marked by lines on posterior surface that
separate at the midpoint and continue downward to merge with the blunt
furcular process; anterior surface of symphysis flatter and unmarked by lines
or depression.
Ulna (fig. 1, C-E) longer and more slender than in Anas platyrhynchos ,
with relatively narrower impression of brachialis anticus muscle (see Table 1) ;
external ligamental attachment notably produced onto palmar surface at base
of olecranon; external cotyla extending from pitlike depression adjacent to
external ligamental attachment and terminating in long, narrow7 lip appressed
to palmar side of shaft; bicipital attachment a well-marked papilla; inter-
cotylar area compressed near olecranon, reflecting similar constricting of
intercondylar groove of humerus; distally, external, condyle less prominent
than in Anas, both in depth and height.
Measurements: Humerus, greatest length, 92.6 mm.; breadth proximally
from internal tuberosity to bicipital crest, 19.7 mm.; greatest breadth distal
end, 14.2 mm.; least transverse breadth of shaft, 6.4 mm. Furcula, height of
symphysis measured immediately adjacent (but not through) furcular process,
4.8 mm.; anteroposterior depth of symphysis (measured at same place), 3.4
mm. Ulna, greatest length, 86. 5 mm.; length to internal cotyla, 79.8 mm.;
1966
Pliocene Birds from Chihuahua, Mexico
7
breadth across proximal cotylae, 9.7 mm.; breadth distal trochlea, 5.9 mm.;
depth external condyle, 8.9 mm.; height external condyle, 9.2 mm.
Discussion: In the anseriform humerus, a well-flared deltoid crest with
excavated external surface and prominent proximal border undercut on palmar
side by deep bicipital furrow, is usually accompanied by a short, raised pectoral
scar, as for example, in the geese and the tadornines. The long pectoral scar
appressed to the shaft accompanies a smoothly rounded external surface of
deltoid crest and shallow bicipital furrow in Anas. Among the anatid humeri
examined, those of Aix (tribe Cairinini) have a combination of appressed long
pectoral scar, and characters of the deltoid crest approaching the condition in
the fossil though the excavation of the crest is less marked. Sarkidiornis (also
of the tribe Cairinini) has a well excavated external surface of the deltoid
crest, but, in this genus, the pectoral scar is short, and there is no marked
depression of the bicipital furrow.
The furcula is more subject to individual variation than is the humerus,
and is, therefore, less reliable as a generic marker. The anteroposteriorly
thickened symphysis appears in several anseriform genera, but is most charac-
teristic of the geese and tadornines. The fossil furcula is distinguished from
both of these groups by the well marked coracoidal tuberosity. Thickening of
the symphysis also occurs among the Cairinini, and, in Sarkidiornis, is accom-
panied by an angular junction of dorsal and posterior surfaces and separated
lines leading to the furcular process as in the fossil bone. However, the whole
symphyseal area is more swollen in the Recent specimen examined. In less
marked degree a tendency to thickening of symphysis and double lines merg-
ing with the furcular process occurs in some individuals of the genus Aix.
The small, papilla-like coracoidal tuberosities and blunt furcular process of
the fossil are also more like the conditions found in the cairinine genera
than in the Anatini. However, the cairinines examined have a more U-
shaped furcula, with the symphyseal area narrower dorsoventrally, and more
rounded. The flat anterior surface of symphysis in the fossil, and comparative
lack of posterior flexure have not been observed in any of the Recent ducks.
If, as is believed, the ulna and humerus of Wasonaka yepomerae came
from the same individual, the ratio of length of the two elements is most
closely approximated among the tadornines (see Table 1). Compared with
Anas platyrhynchos, the fossil ulna is approximately 5 mm. longer than the
maximum of six Recent specimens, while the fossil humerus is slightly smaller
than the average of the Recent form. In Cairina and Aix, the ulna is relatively
short as in Anas; the ulna of Sarkidiornis is not available for comparison.
Anas bunkeri
Fig. 1, F-G
A nearly complete coracoid (LACM 4621 from locality 289), lacking
only the sternocoracoidal process, is teal-like in general contours and size, but
8
Contributions in Science
No. 94
is stockier than in Anas crecca, A. cyanoptera, or A. discors, with shorter,
broader upper end (see Table 2). As the character of heaviness is the chief
diagnostic feature of A. bunkeri, described (Wetmore, 1944:92) from a car-
pometacarpus in the upper Pliocene of Kansas, it seems proper to allocate this
coracoid to that species.
Other characters that distinguish the Mexican coracoid from that of
Recent teals are: head lower, and brachial tuberosity lower relative to level of
top of glenoid facet; coracohumeral surface more depressed, and bordering
Table 1
Proportions of Wasonaka yepomerae and Anas platyrhynchos Compared
(Ratios in per cent)
Furcular symphysis:
Ratio of anteroposterior
depth to height
Wasonaka
yepomerae
71.0
Anas platyrhynchos
(6 specimens)
max. mean min.
59.2 54.9 49.2
Humerus
Ratio of least breadth
of shaft to length of element
6.9
7.5
7.3
7.0
Ratio of length of bicipital
crest to length of deltoid
crest (both measured to tip of head)
74.0
81.2
79.7
78.4
Ulnaa
Ratio of breadth across
proximal cotylae to length
12.1
14.1
13.6
13.3
Ratio of depth of external
condyle to length
11.1
12.4
12.1
11.6
Ratio of height of external
condyle to length
11.5
13.3
12.9
12.0
Ratio of breadth-to-length
of surface for attachment
of brachialis anticus muscleb
19.5
34.3
31.4
28.6
Ulna/Humerus
Ratio of length of ulna to
length of humerus0
86.2
79.9
78.7
77.8
aLength of ulna measured from distal-most extent of external condyle to palmar edge
of internal cotyla.
bThis ratio in Tadorna is 20.9-23.1
cThis ratio in Tadorna is 86.4-88.6
1966
Pliocene Birds from Chihuahua, Mexico
9
Table 2
Measurements and Proportions of Teal Coracoids, Fossil and Recent
(Measurements in millimeters; ratios in per cent)
Anas bunkeri Anas crecca Anas cyanoptera
(referred) ( 1 1 specimens) and A. discors3-
LACM 4621 USNM 12833 max. mean min. max. mean min.
a. Length to
internal
angle
32.3
35.1
33.1
31.0
34.5
33.8
32.0
b. Distance
from head to
underside of
scapular
facet
10.3
10.8
11.7
11.0
10.2
11.7
11.2
10.8
c. Breadth below
furcular facet,
across triosseal
canal
5.0
5.2
5.3
4.8
4.5
5.2
4.9
4.7
d. Breadth of
neck measured
on anterior
face
4.7
4.6
5.0
4.4
4.2
4.9
4.4
4.2
e. Least breadth
of shaft below
procoracoid
3.3
3.4
3.8
3.3
3.0
3.7
3.4
3.2
f. Depth of
shaft below
procoracoid
3.3
3.2
3.0
2.8
2.7
3.1
2.8
2.6
g. Distance from
top of glenoid
facet to internal
edge of furcular
facet
5.6
5.3?
6.9
6.2
5.7
6.7
6.4
6.3
Ratio of
measurement c
to measurement
b
48.5
48.2
45.1
44.1
42.6
45.4
44.1
42.4
Ratio of
measurement d
to measurement
g
84.0
86.7
78.0
71.5
65.5
74.3
69.2
65.5
aMeasurements on two available specimens of Anas discors agreed so closely with
those of four specimens of A. cyanoptera that the two species are grouped together.
10
Contributions in Science
No. 94
contours more curved (anteriorly concave, posteriorly convex); glenoid facet
depressed at center so that posterior edge appears to protrude more
prominently.
The coracoid of A. bunkeri was not described from the type locality, but
has since been recorded from the middle Pliocene of Oregon (Brodkorb,
1958:252) in association with carpometacarpi that compare favorably in all
respects with the type of the species. Unfortunately this coracoid lacks both
ends, and only a minimum breadth of shaft is recorded. Brodkorb ( 1958:253)
lists this dimension as 3.7 mm., a breadth .3 mm. greater than that of the
Mexican fossil. However, both specimens fall within the size range of this
dimension as measured in a series of coracoids of Recent A. crecca. As noted
above, the heaviness of the Mexican bone is notable particularly in the upper
end.
In a recent listing of the localities from which A. bunkeri is known
(Brodkorb, 1964:225), the late Blancan (early Pleistocene) Hagerman Lake
beds of Idaho are included. In 1965 correspondence with the present writer,
Brodkorb states that the listing is tentative and is based on a coracoid (U.S.
Natl. Mus. 12833) assigned by Wetmore (1933:1 1) to “ Querquedula , sp!’
Through the courtesy of Dr. Lewis Gazin, Curator of Vertebrate Paleon-
tology, United States National Museum, I have had the opportunity to examine
this bone. The specimen includes the upper % of the element, but unfortunately
the surface of the furcular facet is eroded, and the anterior edge is broken
away. Nevertheless, in all characters that are preserved, this specimen and the
coracoid from Mexico agree, eg., the upper portion is broad, the head and
brachial tuberosity are seemingly low, the coracohumeral surface is depressed
and curved, and the glenoid facet is depressed.
A number of other teals have been described from the late Tertiary. With
the exception of A. eppelsheimensis, from the lower Pliocene of Germany,
none have the characteristic stocky proportions of A. bunkeri, and the fossil
here at hand. The German specimens, fragments of coracoid, humerus and
wing phalanx, have never been compared in detail with the North American
teals, and there has been no opportunity to examine the material now. How-
ever, the description of the coracoid of A. eppelsheimensis (Lambrecht,
1933:362) characterizes the glenoid facet as being semicircular in form, in
contrast to the outwardly projected contour found in A. crecca. As there is
some individual variation in this contour within A. crecca, I cannot be certain
of the diagnostic value of this character without seeing the specimen. It does,
at least, suggest distinction from the Mexican coracoid, in which the glenoid
facet is outwardly projected.
Oxyura, sp.
Right and left proximal ends of femora (LACM 9735 and 9736) from
locality 289 resemble Oxyura jamaicensis in having a large head prominently
1966
Pliocene Birds from Chihuahua, Mexico
11
protruding from the shaft, with slightly depressed space below on the internal
face of the shaft. The proximal tip of the trochanter is broken on both speci-
mens, but the trochanter appears to be blunt anteriorly as in O. jamaicensis.
The chief difference to be observed between the fossil fragments and the
Recent specimens is the more oval shaft of the former. A proximal half of
ulna (LACM 4631), also from locality 289, resembles this element in O.
jamaicensis in the deeply excavated anconal edge of the attachment of the
brachialis anticus muscle, and the general depression of the entire attachment,
resulting in a flattened internal surface of the shaft. The fossil differs from
O. jamaicensis, however, in having a clearly defined intercotylar ridge; in the
Recent species the two cotylae blend into one another without a ridge. In
breadth and depth of proximal end, the fossil falls within the size range of
O. jamaicensis, but appears to be slightly stouter in diameters of the shaft. The
shaft measurements agree closely with those of the ulna from the middle
Pleistocene of Vallecito Creek, California, referred to O. bessomi (Howard,
1963:13); unfortunately, however, the proximal end is not preserved in the
latter bone. In view of the fragmentary condition of the femora, and the
impossibility of making direct comparison with the geologically younger, O.
bessomi, the Mexican bones are referred only generically. The possibility of
relationship between the California and Mexican birds should be kept in
mind should further material become available at either locality.
Charadriiformes
Erolia (?) sp.
A fragment of distal end of ulna (LACM 4617) from locality 276 agrees
in size and general characters with Recent specimens of this element of the
Least Sandpiper, Erolia minutilla. Definite identification is impossible from
this fragment, and it is highly unlikely that the Recent species is represented
in this Pliocene deposit.
Passeriformes
Mimidae ?
An incomplete humerus, lacking proximal extremity (LACM 4622),
from locality 289, suggests the thrashers in contours, and is only slightly
larger than Recent specimens of Toxostoma redivivum from California.
Literature Cited
Brodkorb, Pierce
1958. Birds from the middle Pliocene of McKay, Oregon. Condor, 60:252-255.
1961. Birds from the Pliocene of Juntura, Oregon. Quart. J. Florida Acad.
Sci., 24(3): 169-184.
1964. Catalogue of fossil birds. Pt. 2 (Anseriformes through Galliformes).
Bull. Florida State Mus., Biol. Sci., 8(3) : 195-335.
12
Contributions in Science
No. 94
Howard, Hildegarde
1963. Fossil birds from the Anza-Borrego Desert. Los Angeles County Mus.,
Contrib. Sci., 73 : 1-33.
Lambrecht, Kalman
1933. Handbuch der Palaeornithologie. Berlin: Gebruder Borntraeger,
xix+1024 pp.
Lance, John F.
1950. Paleontologia y estratigrafia del Plioceno de Yepomera, estado de
Chihuahua. Pt. 1: Equidas, excepto Neohipparion. Universidad
Nacional Autonoma Mexico, Instituto Geologia, Bull. 54, viii+81 pp.
Miller, Loye
1944. A Pliocene flamingo from Mexico. Wilson Bull. 56(2):77-82.
Wetmore, Alexander
1944. Remains of birds from the Rexroad fauna of the upper Pliocene of
Kansas. Univ. Kansas Sci. Bull., 30, pt. 1(9): 89-105.
Woolfenden, Glenn E.
1961. Postcranial osteology of the waterfowl. Bull. Florida State Mus., Biol.
Sci., 6(1) : 1-129.
LOS
ANGELES
COUNTY
MUSEUM
CONTRIBUTIONS
IN SCIENCE
April 4
OBSERVATIONS ON THE BEHAVIOR OF WILD AND
CAPTIVE FALSE KILLER WHALES, WITH NOTES ON
ASSOCIATED BEHAVIOR OF OTHER GENERA OF
CAPTIVE DELPHINIDS
By David H. Brown, David K. Caldwell, and
Melba C. Caldwell
Los Angeles County Museum of Natural History • Exposition Park
Los Angeles, California 90007
OBSERVATIONS ON THE BEHAVIOR OF WILD AND
CAPTIVE FALSE KILLER WHALES, WITH NOTES ON
ASSOCIATED BEHAVIOR OF OTHER GENERA OF
CAPTIVE DELPHINIDS1
By David H. Brown2, David K. Caldwell3, and
Melba C. Caldwell4
Abstract: Behavioral data are presented for a captive sub-
adult female false killer whale, Pseudorca crassidens, from off
southern California. Intergeneric behavior is discussed that oc-
curred between this animal and a captive female Pacific bottle-
nose dolphin, Tursiops gilli, a captive female Pacific common
dolphin, Delphinus bairdi, several captive male and female east-
ern Pacific pilot whales, Globicephala scammoni, and two captive
female Pacific striped dolphins, Lagenorhynchus obliquidens.
The stillbirth of a dead fetus by the common dolphin and the
reactions of the other captive delphinids to it are described. Feed-
ing habits of a captive male Pseudorca in Hawaii are noted, as
well as comments on its intergeneric relationship with a captive
male Stenella cf. roseiventris. Evidence for observational learn-
ing in Pseudorca , Tursiops truncatus and Lagenorhynchus obliqui-
dens is discussed. Ingestion of foreign objects by the Pseudorca is
noted. Known behavior by this form in the wild is considered as
it seems related to the observed captive behavior. Growth and
serologic data for the captive Pseudorca are presented, along
with growth data for a captive Globicephala scammoni male. Ap-
parent records of Pseudorca at sea in the eastern and northeastern
Gulf of Mexico are included, as well as strandings of this form
in Australia. Various kinds of behavior related to that observed
for the Pseudorca are discussed for Tursiops truncatus. Erysipelas
infection and its treatment in captive cetaceans is described.
Introduction
A recent summary (Mitchell, 1965) of eight eastern North Pacific
records (14 specimens) of the false killer whale, Pseudorca crassidens
(Owen), suggests that this species is not as uncommon there as was once
believed. Mitchell based his summary on earlier literature records, on newly-
1Partial support for certain phases of this study was received by the Caldwells through
grants from the National Institute of Mental Health (MH-07509-01 ) and the Na-
tional Science Foundation (GB-1189).
2Curator of Mammals, Marineland of the Pacific, Palos Verdes Estates, California.
3Curator of Ichthyology, Los Angeles County Museum of Natural History; Also Re-
search Associate, Florida State Museum, and Collaborator in Ichthyology, Institute
of Jamaica.
4Research Associate, Los Angeles County Museum of Natural History; Also Staff Re-
search Associate, Allan Hancock Foundation, University of Southern California.
2
1966
Behavior of Wild and Captive Killer Whales
3
obtained skulls from western North American beaches and on sight records
of animals at sea attributed to this species. Included in that summary was the
captive animal which forms the primary basis for the present report. Despite
this increasing evidence for the frequent occurrence of this species, few data
have been provided concerning its behavior in the wild and nothing has been
published heretofore on the details of captive behavior.
Since early 1954, Marineland of the Pacific collectors, Captain Frank
Brocato and his assistant Frank Calandrino, have had the opportunity of
observing from the collecting vessel Geronimo the movements and behavior
of delphinids in the coastal waters of southern California. They first reported
sighting Pseudorca on December 1, 1959 (also see Norris and Prescott, 1961:
335). On this occasion approximately 300 animals were seen three miles
northwest of the west end of Santa Catalina Island, California.
More recently, on October 10, 1963, Brocato and Calandrino, encoun-
tered a second school of false killer whales some four miles southwest of
Long Point lighthouse, Palos Verdes Peninsula, near Los Angeles, California.
On this day Geronimo was following a southwesterly course; the sea was
moderately rough and a wind of 12 to 15 knots was blowing in a southwesterly
direction. Calandrino, who was at the wheel, noticed a school of animals ap-
proaching from the south. These rapidly gained on Geronimo and were
identified as false killer whales. Approximately 300 animals were in a diffuse
school which consisted of numerous small groups of two to six individuals.
The school was spread over an area approximately Vi mile wide and two
miles long. Brocato and Calandrino estimated the whales’ swimming speed
to be at least 10 knots. As the animals passed Geronimo they were observed
to slap their tail flukes forcefully on the water’s surface. Many were vocal-
izing at this time; the sounds emitted by animals from between 150 to 200
yards away were clearly heard. Both Brocato and Calandrino described
these sounds as piercing, harsh and quite consistent (see Schevill and Watkins,
1962: 13). The size of the animals varied from the calves of approximately
five feet to adult animals, some of which were estimated to be 18 feet long.
Shortly after passing Geronimo , several of these groups encountered a
school of bonito, Sarda lineolata (Girard), and commenced to feed upon
these fish. This attracted other groups that also stopped to feed. The collectors
were most impressed by the power and speed of these delphinids whose rushes
at the fish not infrequently caused them to lunge their bodies, as far as the
pectoral flippers, out of the water. On several occasions individuals were
observed grasping the large powerful bonito in their jaws. The Geronimo was
able to approach a feeding group of five or six animals and collector Calan-
drino was thus presented with the opportunity of snaring one of these. The
fellow members of this group stood by the ensnared animal; however, the
other whales continued to behave and feed in a normal manner. Many vocali-
zations were heard at this time that were described as piercing whistles which
seemed to be of a higher frequency than those heard to emanate from other
delphinids.
4
Contributions in Science
No. 95
The first capture was attempted on an animal approximately 13 feet long,
but this individual threw off the snare. Some 30 minutes after the unsuccessful
attempt, a second animal was taken. The other whales in the group were seen
to rub their bodies on the line securing the ensnared animal. When alongside
Geronimo’s hull they abandoned the captive, which showed little tendency to
fight against the line. When lifted aboard, the false killer whale was found to
be a female that measured exactly 1 1 feet, 3 inches, from the tip of the snout
to the median notch of the tail fluke. When lying on the stretcher on Geronimo
she snapped her jaws on several occasions when the collectors ventured
too close.
The false killer whale was released upon its arrival at Marineland into
a holding tank occupied by three female eastern Pacific pilot whales, Globi-
cephala scammoni Cope. The animal began immediately to swim around the
periphery of the tank at great speed, whistling constantly. The individual
phonations lasted approximately two seconds. The animal’s calls were loud
and audible at a considerable distance from the tank. The following morning
the whale had slowed down, but continued, however, to swim in a circular
motion around the enclosure and appeared to shun completely the company
of the Globicephala which at this time maintained a stationary position at
the surface in the center of the tank.
Later in the morning, while throwing squid to these animals, the attendant
was amazed to see the Pseudorca commence to feed. It did this quite vora-
ciously and accepted a considerable quantity of food throughout the day. The
following morning the false killer whale swam to the feeding platform and
accepted mackerel directly from the attendant’s hand.
On October 14, after draining the tank to a depth of three feet, the whale
was examined and a blood sample taken (see Appendix I). The animal
accepted food at this time and did not appear alarmed by the procedures.
In the ensuing weeks the false killer whale became very tame and
quickly learned to leap free of the water to receive portions of her food
(Fig. 1).
No aggressive behavior was directed towards the pilot whales. The smaller
of this trio would, however, attack and hotly pursue the Pseudorca around
the tank.
This lack of aggression by the false killer whale was also observed by
Mr. Chris Varez ( pers . conversation, 1965) for a male that he once handled
at Sea Life Park in Hawaii. Varez noted that this animal was very mild in
its relationships with other delphinids captive with it. Like the Marineland
animal, it also readily learned to take food, although it would become startled
at first if Varez touched it during this learning process. However, Varez was
impressed with the fact that even though the Pseudorca would become startled
and swim away on such an occasion, it would appear to reach out with its
tail flukes as it swam by.
The false killer whale was moved to the circular oceanarium tank on
November 4. The animal appeared at ease during these proceedings and lay
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5
Figure 1. Captive female false killer whale jumping completely clear of the water on
cue. The general outlines of the body are clearly shown. Photograph by Cliff Brown,
Marineland of the Pacific.
quietly in a stretcher while the transfer was in progress. Upon release, she
quickly swam to a platform and accepted food.
Marineland’s circular tank is 80 feet in diameter and 19 feet deep. Glass
windows placed in three levels of corridors permit observation of the animals
exhibited within its depths. From this underwater observation point, we
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observed that our sub-adult female possessed double twin mammary slits
(i.e., one pair of slits on each side of the vaginal opening). However, Comrie
and Adam (1938: 521f.) have indicated that, in a fetal specimen they examined,
one slit of each pair seemed to house a functional nipple and the other of
each pair they termed an accessory groove.
Social Behavior of Pseudorca with other Captive Delphinids
Prior to November, 1963, the Marineland exhibit consisted of one male
and one female pilot whale and two female Pacific striped dolphins, Lagen-
orhynchus obliquidens Gill. The active false killer whale (Fig. 2) tended to
ignore the lethargic pilot whales and sought the company of the active dolphins.
The three animals thereafter were seen rubbing their bodies together and the
larger animal frequently “mouthed” the bodies of her small companions. The
whale often pursued the dolphins around the tank. This playful activity
usually ended in the three swimming closely together, the smaller dolphins
coasting alongside the false killer whale.
Figure 2. Captive female false killer whale underwater. Photograph by Cliff Brown,
Marineland of the Pacific.
A female Pacific bottlenose dolphin, Tursiops gilli Dali5, placed into the
tank on December 10, 1963, became a close companion of the false killer
whale. Immediately following her introduction, the dolphin was closely
examined by the larger animal and subjected to the gentle “mouthing”
described above.
5Daugherty (1965: 43) followed some recent authors in using the combination T.
truncatus gilli for this form. However, we follow more common published usage in
giving the form named gilli full specific recognition.
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Behavior of Wild and Captive Killer Whales
7
Chris Varez ( pers . consersation, 1965) stated that the male Hawaiian
Pseudorca that he worked with at Sea Life Park soon became close companions
with a small male spinner dolphin, Stenella cf. roseiventris (Wagner)6, and
often carried it on his snout even though the almost-white (at capture; it
later darkened in color) Stenella was in apparent perfect health.
Behavior of Pseudorca During Birth of a Common Dolphin
On December 13, 1963, a female Pacific common dolphin, Delphinus
bairdi Dali7, arrived at Marineland. Shortly before being placed into a
quarantine tank, the dolphin showed the symptoms of shock. Following
the intramuscular injection of a tranquilizer drug, the animal became soporific
and could not remain at the surface without assistance. Help was rendered
by one of Marineland’s divers who entered the tank and supported the little
dolphin in his arms. Upon release, the effects of the drug remained evident
and the animal drifted at the surface and made little attempt to swim. Silver
smelt thrown into the tank elicited immediate response and the tranquilized
newcomer consumed a quantity of this fish. The dolphin continued to feed
and on January 9 she was transferred to the Circular Tank.
The common dolphin is difficult to maintain in a captive environment.
This species appears to be peculiarly emotional and particularly sensitive to
the competitive feeding behavior normally demonstrated by larger, more
aggressive, forms (Brown and Norris, 1956: 318). This specimen, however,
appeared to adapt rapidly to an enclosure shared with delphinids of four
other genera.
Dolphins in the latter stages of pregnancy normally display a pronounced
distention of the inguino-abdominal region. The small common dolphin
failed to show these signs. It was, therefore, a surprise when, at approximately
11:50 AM on February 15, observers saw a small tail protruding from her
birth canal. The birth progressed very rapidly and by 12:05 PM the entire
posterior portion of the fetus had been expelled. The umbilical cord, which
seemed stretched and taut, was clearly visible.
The striped dolphins and false killer whale followed the laboring female
(Fig. 3). The dolphins showed particular interest and nosed the female’s
abdominal region on several occasions (Fig. 4).
GIn the use of this name for the Hawaiian animal, we follow the suggestion of F. C.
Fraser {pers. comms. from R. J. Harrison to Brown, 1965, and from Fraser to D. K.
Caldwell, 1965) following his examination of material of this species furnished him
by Brown.
7F. C. Fraser, British Museum (Natural History), regards the Pacific common dol-
phin under study here as probably conspecific with the Atlantic form, D. delphis
Linnaeus, {pers. comm., 1964, to Brown after making a comparison of skulls from
the Atlantic and northeastern Pacific). Daugherty (1965: 26) used the trinomial
combination D. delphis bairdi. For the present, however, we follow usual published
usage in our application of full and separate specific rank to the form of Delphinus
here discussed.
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Figure 3. Captive female Pacific common dolphin in labor (fetus partially extruded)
followed immediately below and to the right by two female Pacific striped dolphins
and below them by the female false killer whale. The tail of the fetus obscures the
head of an eastern Pacific pilot whale circling in the background. Photograph by Cliff
Brown, Marineland of the Pacific.
The dorsal fin of the calf appeared to obstruct its further passage. In
normal births the dorsal fin folds at its base either to the right or left, but
in this case it remained erect and caught internally at the apex of the
vaginal introitus (Fig. 5).
At 12:15 PM one of the striped dolphins grasped the fetal tail flukes in
its mouth and withdrew the infant from the parental birth canal. A discharge
of amniotic fluid and a little blood followed the delivery (Fig. 6).
The infant was stillborn, and delayed expulsion at a critical phase of
parturition was no doubt incriminated in this fetal death.
McBride and Kritzler (1951: 253) attributed difficulties in the birth of
an Atlantic spotted dolphin, Stenella plagiodon (Cope), to the left pectoral
flipper impeding passage of the calf. It is interesting to note, however, that the
maternal exertions witnessed by these authors were not observed in the present
case in question.
Our common dolphin, attended by the striped dolphins, carried her dead
infant's body to the surface. These efforts were, however, terminated by the
male pilot whale, who seized the body by its head (Fig. 7). The pilot whale
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Behavior of Wild and Captive Killer Whales
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Figure 4. Captive female Pacific common dolphin in labor with fetus partially ex-
pelled. She is followed to the left by the female false killer whale and to the lower
right by two female Pacific striped dolphins. Photograph by Cliff Brown, Marineland
of the Pacific.
devoured the small cadaver, entire, after carrying it to and from the surface
for 38 minutes (see Appendix II).
The common dolphin at first seemed little affected by the intervention of
the pilot whale, but appeared greatly distressed by his ingestion of the cadaver.
Whistling constantly, she moved rapidly around the tank, swimming in an
erratic manner, apparently searching for her calf.
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Figure 5. Captive female false killer whale following female Pacific common dolphin
with protruding fetus. The dorsal fin of the fetus is erect (only the base can be seen)
and hooked internally at the apex of the vaginal opening. A reflection on the viewing
window has been retouched so that it is not so obvious as it appears just before the
snout of the false killer whale. Photograph by Cliff Brown, Marineland of the Pacific.
The animal quickly resumed a more normal swimming pattern, in the
company of the striped dolphins, but she continued to vocalize intermittently
for several hours.
Since 1:00 PM, continuous uterine contractions had caused a three-inch
length of the umbilical cord to move in and out of the female’s urogenital
opening. At 4:06 PM, the common dolphin sought the company of the
female false killer whale. She was observed at this time to deliberately avoid
the company of the striped dolphins and begin to swim on the west side of
the tank quite close to the surface. The false killer whale swam to the little
dolphin and, after an apparent deliberate examination of her genital area,
gently grasped the umbilical remnant in her mouth, and with a lateral move-
ment of her head withdrew this tissue some six inches from the common
dolphin’s body. The dolphin rolled on her back and broke away from the
larger animal, but then returned and again waited for the false killer whale.
Once more, the whale seized the placenta and repeated the behavior previously
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Behavior of Wild and Captive Killer Whales
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Figure 6. Following the delivery of her dead fetus (seen just below and to the left of
the large female false killer whale), a cloud of amniotic fluid and a little blood is
discharged by the now-upside-down female Pacific common dolphin (center). An
interested female Pacific striped dolphin is seen at the lower right, while in front of
the false killer whale another female Pacific striped dolphin nuzzles the dead fetus.
Photograph by Cliff Brown, Marineland of the Pacific.
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Figure 7. Captive male eastern Pacific pilot whale carrying newly-stillborn Pacific
common dolphin fetus just prior to swallowing it. Photograph by Cliff Brown, Ma-
rineland of the Pacific.
described and withdrew the membrane another three inches. The common
dolphin during these periods was observed to actively flex her body and
appeared to try to assist the false killer whale in its attempts to remove the
afterbirth. At the third attempt, the female false killer whale was successful
and withdrew the entire placental membrane from the smaller animal. This
was released and immediately both animals resumed normal activity in the
tank. The free placental membrane was closely examined by the striped
dolphins, but was swept away and down the drain before it could be recovered
by the Marineland staff or before the dolphins could further investigate it.
Sexual Behavior
Intergeneric sexual behavior has been observed between the female
Pseudorca and the large male Globicephala (the same animal discussed in
Appendix II) held captive with her.
On April 27, 1965, at 1:00 PM iust after the male pilot whale had been
fed, the false killer whale was observed lying on her side at the surface. The
male slowly approached and rubbed his bulbous cephalic melon against her
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Behavior of Wild and Captive Killer Whales
13
tail flukes for some 5 to 8 minutes (Fig. 8). On another occasion, it had
been noted that this same male pilot whale and a smaller female of the same
species violently bumped melons, head on, in their precopulatory sexual
behavior (Brown, 1962: 60).
Figure 8. Diagrammatic representation of position of captive male eastern Pacific pilot
whale while rubbing his cephalic melon on the tail flukes of female false killer whale
during initiation of precopulatory sexual behavior. Sketch by Donald Hackett, Ma-
rineland of the Pacific.
After rubbing his melon against the tail flukes of the Pseudorca, the
pilot whale had an erection. The Pseudorca then remained on her side at the
surface while the pilot whale swam alongside her, belly to belly and head
to force her way between the Pseudorca and the male pilot whale, rubbing
both of the large animals in the process. She was not successful in this attempt
to separate the mating pair.
At 7:30 AM on the third day, April 29, the same preliminary behavior
was again observed. However, on this occasion intromission occurred (Fig. 9).
During this behavior on the third day, a small female pilot whale attempted
tank wall. Similar behavior, also ending in the failure of intromission, was
observed on the following morning at 9:00 AM.
to head. Although attempted, intromission failed— possibly because the pair
was disturbed when the currents in the tank caused the animals to drift into the
Tavolga and Essapian (1957: 14) noted that, in pairs of captive Tursiops
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truncatus (Montagu)8 in Florida, the male took the more active role in
initiating precopulatory behavior. This was also the case in the sexual inter-
action between the Pseudorca and the Globicephala. Here the male appeared
to make the initial advances and no attempts by the female Pseudorca to
Figure 9. Diagrammatic representation of positions of captive female false killer
whale, on her side at top, and upside-down male eastern Pacific pilot whale, during
intromission. Sketch by Donald Hackett, Marineland of the Pacific.
solicit his attentions were observed. These findings, and those of Tavolga
and Essapian, are contrary to those observed previously tor Globicephala-
Globicephala and Lagenorhynchus-Tursiops sexual pairs at Marineland of
the Pacific. Instead, the females generally seemed to be responsible for the
initiation of precopulatory behavior (Brown, 1962: 61). We have also recently
observed this same female sexual aggression in mating pairs of T. truncatus
at Marineland.
Homosexual Behavior
The female Pseudorca also was involved in homosexual behavior. Shortly
after she was introduced into the Circular Tank with the Pseudorca, the female
Tursiops gilli and the false killer whale established the close relationship
8Also see footnote 5. If one follows some recent authors in giving the form named
gilli only subspecific rank, then this Atlantic form would be called T. truncatus
truncatus.
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Behavior of Wild and Captive Killer Whales
15
noted above. Soon after this relationship was established, homosexual behavior
was observed between the two.
In this behavior one female rubbed the urogenital aperture of the other
with her snout (jaws closed) and often even inserted the tip of the snout
(jaws closed) or lower jaw (mouth open) into the vulva of the passive
participant. This position was frequently maintained even as the two animals
swam around in the tank at considerable speed. Immediately after such an
interaction, the two participants exchanged positions and roles and the behavior
was repeated. Such behavior was observed on many occasions.
This same behavior was also exhibited by the Pseudorca with one of
the small female Globicephala, with each taking both roles.
Again, the same behavior has been observed between the same captive
female Delphinus and the same captive Tursiops gilli noted above. We have
also observed it as it took place recently at Marineland between two captive
female T. truncatus and two captive female Lagenorhynchus obliquidens, and
Brown (1962: 62) reported similar behavior between a captive female
L. obliquidens and a female Globicephala scammoni.
Within minutes, the same two female T. truncatus that we observed
engaged in homosexual behavior were also engaged, before and after, in
sexual behavior with a male of the same species.
Protective Behavior
On January 6, 1965, it became necessary to drain the Circular Tank for
the purpose of giving the occupants their bi-annual erysipelas vaccination
(see Appendix III). The water level had reached the desired depth of three
feet by 7:45 PM. It was then dark, and it was necessary to illuminate the work
area with flood lights placed around the top of the tank.
After administering a prophylactic injection to a pilot whale, two of the
men helping in the tank, trainer Ray Cribbs and diver Richard Blacker,
effected the capture of the common dolphin. The small dolphin immediately
commenced to emit a series of high-pitched whistles. The false killer whale,
apparently attracted by these vocalizations, inserted her head between the
man holding the hinder end of the Delphinus and the animal’s body. It then
gently, but nevertheless very deliberately, proceeded to push its Companion
out of its captor’s arms. Both Cribbs and Blacker later stated it was impossible
to hold the Delphinus at the time. The false killer whale made no attempt to
bite, and in fact failed to direct any aggressive behavior at either of the
men involved. Upon effecting the dolphin’s release, both it and the Pseudorca
swam together for a short time. Shortly thereafter, the common dolphin was
captured once more, and the injection made quickly before the false killer
whale could again intervene.
On March 20, 1965, the female Delphinus refused to accept food. Emesis
also occurred on several occasions during this and the following day. It was
decided to remove her from the exhibit, and in the evening the Circular Tank
was again drained to a depth of approximately three feet.
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Upon the common dolphin’s being secured by trainer Joe Beazie, the
Pseudorca again approached and intervened. The dolphin was whistling at
this time. The whale quickly effected the dolphin’s release by pushing her out
of the man’s arms. The next attempt was made by both trainers Cribbs and
Beazie and they were again obliged to release the animal owing to the inter-
vention of the false killer whale. At this time the false killer whale grasped
and gently pressed one of Cribb’s legs in her mouth. A third attempt made by
Beazie elicited the same behavior; i.e., the whale seized his leg and relinquished
its hold only upon the dolphin’s release. Immediately after this, the false killer
whale carried the Delphinus on its back for several seconds. It was then
decided to drain more water out of the tank until a depth of 18 inches at the
sides was reached. Again the animal was captured and this time removed.
During this last attempt, the false killer whale swam to the shallow edge of
the tank and stranded herself in her effort to come to the aid of the common
dolphin.
On the day following the removal of the Delphinus (see Appendix IV),
the Pseudorca continued to behave and feed in a normal manner.
Observational Learning
The successful maintenance of the Pseudorca provided many opportuni-
ties to observe her behavior under prolonged captive conditions. As noted
above, this animal rapidly adapted to the captive environment and consistently
proved more precocious than other delphinids exhibited at Marineland. She
also adapted to new situations and new objects in her tank more rapidly than
we have observed other delphinids to do (e.g., Tursiops truncatus, T. gilli,
Lagenorhynchus obliquidens, Globicephala scammoni, Stenella plagiodon,
S. cf. roseiventris and Delphinus bairdi). In conjunction with this behavior,
it was found that the false killer whale appeared to be extremely adept at
learning by observation (also see Appendix V). Before her introduction into
the displays, the learning by one animal of unnatural conditioned behavior
by observation of another animal had not been encountered in Marineland’s
cetacean colonies.
However, the Pseudorca, after being introduced into the Circular Tank,
in this manner learned several of the trained pilot whales’ tricks. Some of this
learned behavior was later reinforced, but initially it was not even encouraged.
All of the tricks were learned by the Pseudorca during the first year of her
captivity and included: (1) “Shaking hands’’ i.e., lying on the side at the
surface of the water and presenting a flipper, out of water, to be “shaken” by
the trainer; (2) “Dance’’ i.e., extending the upper part of the body out of
the water in a perpendicular position and revolving in place; (3) “Sing’’ i.e.,
vocalize through the blowhole with that opening out of water; (4) Leap to
grasp a large paddle in the mouth and fall back into the water in order to
trigger a camera that photographs a human patron.
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Behavior of Wild and Captive Killer Whales
17
Ingestion of Foreign Objects
The pathologic consequences of foreign-body ingestion are a constant
threat to survival of captive delphinids (Brown, 1960: 345; Brown, et al.,
1960; Caldwell, Caldwell and Siebenaler, 1965: 6). Dr. Masayuki Nakajima,
Curator of the Enoshima Marineland, Futisawa, Japan, recently wrote Brown
( pers . comm., October 7, 1965) that the ingestion of pieces of rope, cloth,
stones and balls have been incriminated in the deaths of 14 dolphins of
unstated species held at that establishment. At Marineland of the Pacific the
swallowing of foreign objects has resulted in the loss of 8 trained animals,
and is the greatest single cause of cetacean mortality yet encountered at the
oceanarium.
The false killer whale seems peculiarly prone to playing with objects of
this kind. Fortunately, however, when observed indulging this habit she will
usually swim to the feeding platform when summoned and permit removal of
the object from her mouth (Fig. 10). Combs, pens, plastic toys, coins, and a
variety of items have been recovered in this manner.
In the evening of June 1, 1965, two metal bottle caps were dropped into
the tank where both were seized almost immediately by the false killer whale.
Figure 10. Captive female false killer whale voluntarily allowing an attendant to
reach into her mouth to retrieve a foreign object which she has picked up in the tank.
Photograph by Cliff Brown, Marineland of the Pacific.
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Chief diver Jake Jacobs witnessed this event and, despite his endeavors to
effect their recovery, both caps were swallowed by the whale. The adminis-
tration of two quarts of mineral oil the following day induced emesis. A
number of items were regurgitated. However, these settled to the floor of
the tank and migrated into the drain before identification could be made.
A plastic toy and cigar holder floated, however, and these were netted and
removed from the water before they could be swallowed again by the whale.
The use of mineral oil as an emetic in delphinids was described by Brown
(1960: 345) after the egestion of a rubber inner tube by a captive Globi-
cephala scammoni. Since then mineral oil has been used successfully at Marine-
land of the Pacific in several similar cases, and its use apparently saved the
life of a captive Tursiops truncatus that had swallowed a large amount of
plastic material (Caldwell, Caldwell and Siebenaler, 1965). In September,
1965, a trained Tursiops sp. owned by Mr. Jack Evans of Tweedheads Aqua-
rium in Australia, swallowed a metal whistle used in training the animal. Two
quarts of mineral oil were given by intubation. Regurgitation occurred and
the whistle was recovered 24 hours after this treatment.
On July 8, the Marineland false killer whale showed little interest in its
food and avoided contact with the other animals. Its behavior did not change
during the following two days.
On July 11, the whale became inappetent and drifted at the surface in
a listless manner. The tank was drained to a three-foot level at 8:45 PM.
The animal proved unusually difficult to restrain and, although several attempts
were made, efforts to take a blood sample failed. The whale was, therefore,
released after the intra-muscular administration of a wide-spectrum antibiotic
and vitamin Bx.
The next day the whale was appetent once more. Emesis was again
induced but additional foreign bodies were not seen.
In the following two weeks the false killer whale gradually returned to
its usual feeding and behavior pattern, and at this writing appears normal
in every way.
The accidental dropping of inedible material is a perpetual menace to the
health of this interesting animal, and its continued survival is largely due to
the vigilance and concern of the Marineland training and diving staff.
Growth
On November 4, 1963, when she was first moved to the Circular Tank,
the Pseud orca weighed 825 pounds and her snout to caudal-notch length was
reconfirmed at 1 1 feet, 3 inches.
On August 31, 1965, when the water in the Circular Tank was lowered
for other routine purposes, the false killer whale was again measured and
weighed for the first time since that date in 1963. It was found that she had
increased in length to 12 feet, 5 inches, and in weight to 1100 pounds.
An accurate record of her food consumption had been kept during this
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Behavior of Wild and Captive Killer Whales
19
22-month interval, and it was found that a total of 30,650 pounds of fish and
squid had been required to make these increases in this very active animal.
Although he was too large to weigh with the facilities available, by
comparison the large male pilot whale, involved in various incidents of behavior
noted herein, was 17 feet, 3 inches in snout to caudal-notch length when first
measured on January 21, 1959, and had increased to a length of 19 feet, 2
inches on August 31, 1965. Brown (1962: 63) earlier noted that a female pilot
whale had increased 23 inches in length over a period of three years and two
months in captivity at Marineland.
Wild Behavior Related to the Captive Observations
Published reports on observations of large herds at sea and on mass
strandings provide abundant evidence that Pseudorca is a very social form
(e.g. : Harmer, 1931; Fraser, 1936, 1937: 296-298, 1946: 40; Peacock, 1936;
Kellogg, 1940: 84, 89; Mitchell, 1965; Appendix VI herein). It is probably
due, at least in large part, to this social behavior among its fellows in the
wild that the captive animal discussed above has proven so adaptable in its
relationships with its tank mates, consisting at times of representatives of as
many as four other delphinid genera.
Furthermore, the false killer whale appears to exhibit a remarkable
lack of fear toward new and strange situations and hence seems to be more
quickly and readily trainable in captivity than many other cetaceans (most
notably Tursiops truncatus ) which have been studied. The rapid acclimation
of our captive to taking food from the hand of a trainer has already been
discussed above. That this reduced fear behavior also regularly extends to the
wild is suggested by the following field observations on cetaceans made in
Florida, and which seem, with little doubt, to be attributable to the false
killer whale.
In the hope of obtaining field observations on cetaceans, during the
summers of 1964 and 1965 the Caldwells interviewed a number of profes-
sional sport-fishing-boat captains operating in the northeastern Gulf of Mexico
in the region from shore to some 50 miles offshore between Pensacola and
Panama City, Florida. During the course of the conversations, there were
persistent reports of a “large black porpoise” which the captains called “black-
fish!’ The captains reported that the “blackfish” were most often seen singly
or in pairs, but that sometimes they were seen in herds of up to an estimated
100 animals and were said to occur in waters of 30 fathoms or greater, which
in this region would mean some 20 miles or more offshore. The “blackfish”
were said to be about twice the size of the common offshore spotted dolphin
of the region, Stenella plagiodon, or thus an average length of some 15 feet.
It was said to have a “snout’’ a full set of obvious teeth in both jaws, a dorsal
fin more curved than that of the spotted dolphin, and to be overall black in
color. A universal behavioral comment was that the “blackfish” is a notorious
fish-stealer and that on many occasions large game fish of different kinds had
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been stolen from fishermen’s lines before they could be landed. On one
occasion, such a fish was dropped by the “blackfish” and on being gaffed
from the fishing boat proved to be a large snapper ( Lutjanus ) weighing some
16 or 17 pounds. Because of this habit of fish stealing, the “blackfish” have
been observed in more detail by the fishermen than most cetaceans that they
observe and hence the greater amount of lay information available concerning
the animal. Animals with similar appearance, behavior and ecological distri-
bution have also been informally reported to us by fishermen working off
the general region of St. Petersburg, Florida, in the eastern Gulf of Mexico.
In trying to identify these animals, the most obvious suggestion would be to
call them the pilot whale {Globicephala) , which seems to fit the description
in size and color and in the Gulf of Mexico as elsewhere is often termed
“blackfish!’ However, several of the captains interviewed were familiar with
Globicephala at sea and in captivity and stated that their “blackfish” was
definitely not the same because it possessed a noticeable “snout” and lacked
the large bulbous forehead of Globicephala. The full set of teeth reported for
the unidentified “blackfish” eliminates any of the beaked whales of the genera
Ziphius and Mesoplodon which might be of about the right size and color.
The genera Kogia and Grampus can be eliminated on general morphology and
tooth description and somewhat on the basis of size. Such delphinid genera as
Orcinus, Tursiops, Stenella, Steno, Phocoena and Delphinus that are known
from the Western Atlantic can be eliminated for various reasons of size, color
or familiarity by the boat captains. The same may be said for the large toothed
whale, Physeter. By elimination, no other likely cetacean seems to remain to
fit the description of the “blackfish” except the false killer whale, Pseudorca.
Furthermore, and on the more positive scale, this species is known previously
from the Gulf of Mexico (Bullis and Moore, 1956; and Fig. 11 herein), it
is the proper size and color, has a full set of very obvious teeth, has a noticeably-
curved dorsal fin (Fig. 1), could be said (especially by a layman) to possess
a noticeable “snout” (though not a beak), lives offshore (which is typical of
Pseudorca , according to Bullis and Moore, 1956: 5) and is known to feed
at times on large fish (see Scheffer and Slipp, 1948: 289; Bullis and Moore,
1956: 3; Daugherty, 1965: 38; the feeding notes given above as related to
the capture of our California specimen; Appendix VII herein). Finally,
behavioral notes supplied by Donnelley ( 1937) , and repeated by Moore ( 1953 :
141), for an animal observed off southeastern Florida, are surprisingly similar
in context to the reports of fish-stealing given the Caldwells for the Gulf of
Mexico “blackfish!’ In this instance, a lone cetacean, with some certainty
identified from photographs as Pseudorca (an identification with which we
concur after seeing the photographs), took a 2 Vi -foot bonita (probably
Sarda ) bait dragged before it with such force and tenacity that the attached
line soon broke.
The captains interviewed by the Caldwells reported that despite their
willingness to steal fish, the “blackfish” still are often very wary and will not
come too close to the boats. This wariness probably is learned behavior due
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Behavior of Wild and Captive Killer Whales
21
Figure 11. Adult female false killer whale from the central Gulf of Mexico. See
Bullis and Moore (1956) for details concerning this specimen. U.S. Bureau of
Commercial Fisheries photograph courtesy of Harvey R. Bullis, Jr.
at least in part to the frequent gunfire that is directed toward these animals
because of their larcenous practices. One captain noted that once when a
“blackfish” was wounded in such a manner, a second “blackfish” with it
immediately departed the scene. We suspect that the injured animal in this
22
Contributions in Science
No. 95
case was a male, and that the animal with it was a female. Such a lack of
aiding behavior exhibited toward a distressed adult male, even though a
comrade, seems almost typical of cetacean behavior under such circumstances
(Caldwell and Caldwell, 1966), and contrasted with the positive interest
shown toward the animal, a female, by its schoolmates during the initial
phase of her capture, as described above, off California. We suspect that the
wounded Pseudorca reportedly abandoned by its schoolmates during capture
off Los Angeles (Norris and Prescott, 1961: 335) likewise was a male.
ACKNOWLEDGMENTS
Several members of Marineland’s staff contributed their observations,
both under captive and wild conditions, to this report. We would like to thank
Captain Frank Brocato and his assistant, Frank Calandrino, for their con-
tinued help and accurate observations made at sea. We also wish to thank
J. Courtland Beazie, Bruce Parks, Richard Blacker and Lawrence Clark for
observations reported here. The interest and advice of Drs. M. E. Webber,
Richard Hubbard and James C. Roberts, Jr., and Mr. Fred Newman, are also
much appreciated. Particular thanks are also due Drs. Rankin W. McIntyre
and John Simpson of the Los Angeles County Livestock Department for their
skillful postmortem examinations of the pilot whale and common dolphin, and
for their continued interest in the health problems of marine mammals. Mr.
Jack Evans, owner of Tweedheads Aquarium, Coolangatta, Queensland, Aus-
tralia, kindly allowed us to use some of his data on erysipelas infection in
Australian Tursiops and on strandings of Pseudorca on those shores. We
would also like to thank Mrs. Eleanor Zetterberg for her help in the prepa-
ration of an early draft of the manuscript, and Mary V. Butler of the Los
Angeles County Museum of Natural History for help with preparation of
one of the illustrations.
Literature Cited
Brown, David H.
1960. Behavior of a captive Pacific pilot whale. J. Mammal., 41(3) : 342-349.
1962. Further observations on the pilot whale in captivity. Zoologica, 47 (1) :
59-64, 2 pis.
Brown, David H., Rankin W. McIntyre, C. A. Delli Quadri, and Robert J. Schroeder
1960. Health problems of captive dolphins and seals. J. Amer. Veterinary
Med. Assn., 137(9) :534-538.
Brown, David H., and Kenneth S. Norris
1956. Observations of captive and wild cetaceans. J. Mammal., 37(3) :31 1-326.
Bullis, Harvey R., Jr., and Joseph C. Moore
1956. Two occurrences of false killer whales, with a summary of American
records. Amer. Mus. Novitates, 1756:1-5.
Caldwell, David K., and David H. Brown
1964. Tooth wear as a correlate of described feeding behavior by the killer
whale, with notes on a captive specimen. Bull. So. Calif. Acad. Sci.,
63(3) : 128-140.
1966
Behavior of Wild and Captive Killer Whales
23
Caldwell, Melba C., David H. Brown, and David K. Caldwell
1963. Intergeneric behavior by a captive Pacific pilot whale. Los Angeles
County Mus., Cont. in Sci., 70:1-12.
Caldwell, Melba C., and David K. Caldwell
1966. Epimeletic (care-giving) behavior in Cetacea. In K. S. Norris, ed.,
Whales, dolphins and porpoises. Proc. 1st Internatl. Symposium on
Cetacean Res., Univ. Calif. Press.
Caldwell, Melba C., David K. Caldwell, and J. B. Siebenaler
1965. Observations on captive and wild Atlantic bottlenosed dolphins, Tur-
siops truncatus, in the northeastern Gulf of Mexico. Los Angeles County
Mus., Cont. in Sci., 91 : 1-10.
Comrie, L. C., and Ann B. Adam
1938. The female reproductive system and corpora lutea of the false killer
whale, Pseudorca crassidens Owen. Trans. Roy. Soc. Edinburgh, 59 (pt.
2, no. 19) :521-531, 1 pi.
Daugherty, Anita E.
1965. Marine mammals of California. Sacramento: Calif. Dept. Fish and
Game, 87 p.
Donnelley, Thorne
1937. Hooking a killer whale. Pleasure, 1(1) :40-41.
Fraser, F. C.
1936. Recent strandings of the false killer whale, Pseudorca crassidens. Scot-
tish Nat., 220:105-114.
1937. Whales and dolphins. In J. R. Norman and F. C. Fraser, Giant fishes,
whales and dolphins. London: Putnam, pp. 201-349.
1946. Report on Cetacea stranded on the British coasts from 1933 to 1937.
Rept. No. 12. London: Brit. Mus. (Nat. Hist.), 56 p., 7 maps.
Harmer, Sidney F.
1931. The false killer dolphin. Nature, 127(3193) :60.
Hester, F. J., J. R. Hunter, and R. R. Whitney
1963. Jumping and spinning behavior in the spinner porpoise. J. Mammal.,
44(4) :586-588.
Kellogg, Remington
1940. Whales, giants of the sea. Natl. Geogr. Mag., 67(1) : 35-90.
McBride, Arthur F., and Henry Kritzler
1951. Observations on pregnancy, parturition, and post-natal behavior in
the bottlenose dolphin. J. Mammal., 32(3) :25 1-266.
McKenna, Owen
1965. Whale size mystery. Australasian Post, for Aug. 12, p. 9.
Mitchell, Edward
1965. Evidence for mass strandings of the false killer whale (Pseudorca
crassidens) in the eastern North Pacific Ocean. Norsk Hvalfangst-
Tidende (Norwegian Whaling Gazette), 54(8) : 172-177.
Moore, Joseph C.
1953. Distribution of marine mammals to Florida waters. Amer. Midland
Nat., 49(1) : 117-158.
Norris, Kenneth S., and John H. Prescott
1961. Observations on Pacific cetaceans of Californian and Mexican waters.
Univ. Calif. Publ. Zool., 63(4) :291-402, pis. 27-41.
Peacock, A. D.
1936. The false killer whale stranded in the Tay Estuary. Scottish Nat., 220:
93-104.
24
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Scheffer, Victor B., and John W. Slipp
1948. The whales and dolphins of Washington State with a key to the cetaceans
of the west coast of North America. Amer. Midland Nat., 39(2) : 257-
337.
Schevill, William E., and William A. Watkins
1962. Whale and porpoise voices. A phonograph record. Woods Hole, Mas-
sachusetts: Woods Hole Oceanogr. Inst., 24-page booklet and phono-
graph record.
Siebold, H. R., and James E. Neal
1956. Erysipelothrix septicemia in the porpoise. J. Amer. Veterinary Med.
Assn., 128(11) :537-539.
Tavolga, Margaret C., and Frank S. Essapian
1957. The behavior of the bottle-nosed dolphin (Tursiops truncatus ): mating,
pregnancy, parturition and mother-infant behavior. Zoologica, 42(1):
Appendix I
Blood specimens are obtained from all newly-collected specimens of
Cetacea at Marineland of the Pacific. Certain of the Marineland trained
dolphins are also routinely subjected to serologic examination.
The animal involved in such an examination is exposed to a minimum of
stress. Capillary bleeding is induced by a small surface incision in the trailing
edge of the dorsal fin or tail flukes. Glass microtubes coated with ammonium
heparate are used to collect blood samples. These microtubes are 150 mm.
in length and hold the volume necessary for a complete blood count. Serologic
examination has proved to be a useful device in both the diagnosis of disease
and its prevention. The following are the complete blood counts for the false
killer whale as taken on October 14, 1963, and on August 31, 1965:
1963
1965
Red blood count
4,500,000
5,200,000
White blood count
6,600
5,600
Hemoglobin
13.9 gm. or 90%
15.7 gm. or 101%
Hemacrit
45%
52%
Differential :
Segs
55
48
Stabs
2
0
Small lymphocytes
37
48
Monocytes
2
1
Eosinophils
4
3
Basophils
0
0
Appendix II
The large male pilot whale was captured and introduced into the Circular
Tank on January 1, 1959 (Brown, 1962), where, after 10 months he was
1966
Behavior of Wild and Captive Killer Whales
25
conditioned to perform in the trained whale display. Prior to the incident
described in the present paper, this male pilot whale had demonstrated unusual
behavior. Two such cases, in 1960 and 1962, have already been reported
elsewhere (Brown, 1962: 62f.; Caldwell, Brown and Caldwell, 1963).
This same animal again commenced to behave in a psychotic manner in
July, 1962, and aggressive asocial activity vied with sporadic anorexia and
depression. The female pilot whales captive with him became increasingly
subject to apparently unprovoked attack despite long periods before when
they had lived in apparent perfect harmony. As the weeks passed these attacks
occurred with increasing frequency, particularly in the late afternoon and
evening hours.
A major change became evident in the male pilot whale following the
oral administration of Sparine (promazine hydrochloride). In the evening
of August 16, 1962, after 7 days of therapy and improved behavior, he
viciously attacked and killed his smallest female companion. The 780-pound
female was thrown clear of the water surface by the violence of the assault.
Pathologic examination of this animal revealed a fractured right ramus of the
mandible and extensive bruising in the left ventral thoracic area. The pleural
cavity contained approximately 14 liters of blood and an irregular tear some
30 mm. in length was found in the right ventricle of the heart.
On the evening of August 17, 1962, in an attempt to prevent further
fatalities, the pilot whales were stranded on the floor of their tank. It was
felt that a common stress conjointly shared might re-establish the strong social
relationship normally so evident in this gregarious species. Throughout the
stranding, the three animals lay closely together, and vocalized continuously.
After 40 minutes the tank was refilled and to this writing further aggressive
behavior has not been directed by the male pilot whale towards his companions
and no further direct medication has been required to tranquilize him.
In the summer of 1963, this large male once more became partially
inappetent. On August 27 he refused nourishment for 7 days and ate only
intermittently during the weeks that followed.
No evidence of infection or disease could be detected, and his fasting
continued despite the administration of appetite-stimulating drugs. Inanition
became pronounced and by the end of September the animal apparently had
lost at least 500 pounds of body weight.
On October 10, 1963, the pilot whale exhibited a dramatic response after
Niamid (nialamide), a psycho-therapeutic (anti-depressant) drug, had been
administered by intubation. He became appetent once more and continued
this improvement during and at the conclusion of three months of Niamid
therapy.
It is not inconceivable that the aberrant activity following the dolphin
birth was allied to the abnormal behavior previously exhibited.
This animal now behaves in an ordinary manner. The whale is fed to
repletion each day and, since his illness, has been excluded from further
participation in the feeding shows.
26
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No. 95
It is felt the psychotic behavior demonstrated by this large whale was
closely associated with environmental stress and normal activity repression.
Since his retirement from active participation in the trained routines, the
animal again indulges in play and reproductive activity, which drives were
not expressed after his integration into the trained whale show.
Appendix III
Infection with Erysipelothrix rhusiopathiae was first incriminated in the
loss of several captive dolphins, Tursiops truncatus and Stenella plagiodon, at
Marine Studios in Florida (Siebold and Neal, 1956). Erysipelas has since
been incriminated in the deaths of delphinids held in other oceanaria in the
United States. In adition, in Australia, Jack Evans reported (pers. comm.
to Brown) that two deaths suspected to be the result of erysipelas septicemia
had occurred in Tursiops that he had supplied to the Taronga Park Zoological
Gardens, Sydney. Mr. Evans reported further that the same pathogens had
been recovered in Australia as in the United States.
Deaths have occurred from the chronic and acute forms of infection.
The onset of symptoms in the chronic form of this disease is heralded by
elevated body temperature, partial or complete inappetence and the eruption
of cutaneous lesions which generally are elevated and sharply outlined. These
can develop on any part of the infected animal.
Cases of chronic infection, if promptly treated, can be controlled readily
by the administration of penicillin. E. rhusiopathiae is a gram negative organ-
ism and is normally extremely sensitive to this antibiotic. If the animal is
treated in time, the skin of the affected area and the adipose tissue beneath
sloughs and the denuded area is gradually invaded by healthy tissue.
The acute form is difficult both to diagnose and treat. The onset of
symptoms is rapid, and these usually consist of inappetence, high fever and
recumbent behavior. The disease generally terminates fatally soon after these
symptoms become apparent.
Two pilot whales succumbed from acute erysipelas shortly after their ar-
rival at Marineland of the Pacific. In both cases E. rhusiopathiae was isolated
from the spleen and liver. Confirmation of this diagnosis was obtained by pass-
ing an inoculum of the isolate into mice, with pathology resulting.
Following these losses, which occurred in 1964, all of Marineland’s del-
phinids have been inoculated with erysipelas bacterin. This bacterin is prepared
for the immunization of swine and turkeys by subcutaneous administration.
The bacterin is administered to delphinids by injection into the muscle of the
caudal peduncle. No anaphylactoid reactions have resulted from this treatment
to date, and other effects have been limited to a transitory stiffness in the area
of injection.
Since embarking on this program of preventive medicine, no further losses
from acute erysipelas septicemia have occurred. Several mild cases of the
chronic form have responded rapidly to the oral administration of penicillin.
1966
Behavior of Wild and Captive Killer Whales
27
Appendix IV
After removal from the Circular Tank, the female common dolphin was
placed into a holding pool and treated for gastric impaction. Her condition,
however, continued to regress and on April 15, 1965, the animal was found
dead on the floor of her pool.
Pathologic investigation showed intussusception to be the cause of death.
This condition was found in the small intestine 12 inches below the pylorus. In
this area the submucosae showed varying degrees of necrosis, edema and con-
gestion.
Intussusception, or the evagination or telescoping of the intestine, is com-
monly caused by irregular or excessive peristaltic movements. Enteritis, intes-
tinal parasites and major dietary changes are frequently incriminated in the
excitation of this condition. No parasites or gastroenteric pathogens were re-
covered from this animal. However, the variety of fish on which she had sub-
sisted for many months became unavailable and a change of food became
necessary shortly before the initial symptoms of illness were recorded.
Appendix V
Caldwell, Caldwell and Siebenaler (1965: 4) described the phenomenon
of observational learning in captive bottlenose dolphins, Tursiops truncatus.
More recently, at Marineland of the Pacific, a female T. truncatus, some 6 feet
in length and originally captured some months before at New Smyrna Beach,
Florida, learned to spin by observing the natural behavior of a 65-inch female
spinning dolphin, Stenella cf. roseiventris (Wagner), from Hawaii. The spin-
ning by the Tursiops was observed only minutes after she was placed for the
first time in a show tank with the Stenella. The Tursiops spinning leap (Fig. 12)
was made almost immediately after the Stenella had made her spin upon a
previously-reinforced cue. Although a different species of Stenella apparently
was involved, the spinning behavior was described and illustrated by Hester,
Hunter and Whitney ( 1963). The Marineland Stenella did not leap as high as
the illustrated dolphin, but the horizontal form of the leap and spin on the long
axis of the body was essentially the same.
At this writing an attempt is being made to reinforce the spin by the
Tursiops so that the animal can be integrated into the dolphin show. However,
before such reinforcement was begun she was clearly learning to make the
spinning motion without human instruction. Although not polished, the spin
out of water consisted of almost 1 Vi complete revolutions, or about he action
illustrated by the first 9 to 1 1 frames beginning on the right in the figure cited
above. We have neither seen ourselves nor heard reports of a spinning leap by
an Atlantic Tursiops made under wild or unconditioned captive conditions.
Newly captive Pacific striped dolphins ( Lagenorhynchus obliquidens) will
learn to leap in an arc as high as 15 feet from the surface of the water, copying
the leaps of specimens of this species already established in the Marineland
display. Such high leaping is reinforced at Marineland, but leaping in a similar
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Contributions in Science
No. 95
Figure 12. Captive female Atlantic bottlenose dolphin during her spinning leap
described in text. Photograph by Cliff Brown, Marineland of the Pacific.
manner is natural to this species and these active dolphins can in fact frequently
be observed behaving in this manner at sea. However, the high leaps in unison
and on cue at Marineland seem at least in large part to be a result of observa-
tional learning by new animals from old residents.
1966
Behavior of Wild and Captive Killer Whales
29
Learning by observation may have been demonstrated by a captive male
T. truncatus at Marineland of the Pacific in which he observed adult females
of this species break up their food before eating it and then apparently attempt-
ed to do this for himself (see Norris and Prescott, 1961 : 312).
Appendix VI
Mr. Jack Evans has provided a stranding record, supported by photographs
(Fig. 13), that clearly is the false killer whale. Although an exact date is not
available, Mr. Evans reported that several of these animals stranded “several
years ago” on a beach in northern New South Wales approximately 300 miles
south of Coolangatta, Queensland, Australia. One of the photographs that he
provided shows heavy wear on the tips of all of the teeth of the left mandibular
row of one of the whales. The wear shown is reminiscent of that illustrated by
Caldwell and Brown ( 1964: 133) for the ferocious killer whale, Orcinus orca
(Linnaeus), although it is less extreme in the case of the illustrated Pseud orca.
Such apical wear in a false killer whale may only be of the same sort often
found in older individuals of other delphinids, ones that generally only swallow
their food entire, and not the rather complicated tooth wear found in Orcinus.
However, its presence, coupled with the known feeding habits of Pseudorca as
described herein (Appendix VII), should be noted as an invitation for careful
study of the wear on teeth of this latter species.
Ten mass strandings of small whales on Australian shores during the past
17 years were listed (five with photographs) in a recent popular newspaper
article (McKenna, 1965). Two of these strandings were, with little doubt, of
Pseudorca. Where data were given, including information taken from the pho-
tographs, at least four animals stranded together in each case, and as many as
about 80 (another news release on this stranding gave a figure of 100). Many
lay opinions, mostly untenable, were proposed to explain the strandings. How-
ever, it is perhaps significant to note that McKenna reported that all of the
strandings occurred on gradually sloping beaches, on broad shallow flats or in
shallow bays and estuaries.
Of the ten reports, we believe that the stranding of some 80 (100) animals
on a beach fully exposed to the sea, near the tip of Flinders Island, in Bass
Strait, sometime in the first half of 1965, consisted of Pseudorca. The animals,
in low-level photographic aerial view, were dark in color, had appropriate body
proportions, in several instances had the typical form of the pectoral flipper that
is long and sickle-shaped (see Figs. 2, 5, 6 and 1 1 herein), and ranged, accord-
ing to McKenna, from 10 to 20 feet in length (the latter figure is probably
somewhat excessive).
A stranding of at least 25 animals (the photograph was cropped in such a
way as to suggest that there were many more) near Burnie, Tasmania, in 1948,
also appeared to be of Pseudorca although our determination in this case was
made with somewhat less confidence than the previous one. The animals were
dark in color, had the strongly-hooked dorsal fin typical of Pseudorca (see Fig.
30
Contributions in Science
No. 95
Figure 13. Upper and lower: stranded false killer whales on beach in northern New
South Wales approximately 300 miles south of Coolangatta, Queensland, Australia.
Note worn tips of mandibular teeth below swollen tongue in lower animal, apparently
a male. Photographs courtesy of Jack Evans, Tweedheads Aquarium, Coolangatta.
1966
Behavior of Wild and Captive Killer Whales
31
1 herein), and, as judged from human figures associated with the animals, were
of an appropriate size. The pectoral flipper of one whale appeared to have the
typical Pseudorca sickle-shape, but this same animal seemed to have a much
more prominent chevron of light pigment on the ventral side between and
anterior to the insertions of the pectoral flippers than the rather faint light
chevron of the Marineland captive. However, this difference could have been
a photographic artifact due to glare on the wet animal.
The McKenna photographs of strandings in 1956 in northern New South
Wales (at least 40 animals); at Bremer Bay, Western Australia, in 1960 (at
least 4 individuals); and at Ninety Mile Beach, Victoria, in 1961 (at least 27
animals), all could be of Pseudorca. The animals were dark in color and ap-
parently of an appropriate body configuration. However, there were no suitable
scale guides with which to estimate their size, and characters typical of Pseu-
dorca were not obvious as had been the case in the two other strandings just
discussed.
According to McKenna, 32 “black” whales were reportedly seen “jostling
one another” to leave the water to strand at Wreck Bay, near Nowra, New
South Wales, in 1963.
Four other strandings reported by McKenna are even less certain with re-
gards to specific identification: 13 whales swam ashore at Newcastle, New
South Wales, in 1962, and three mass strandings of whales “in big numbers”
took place in a recent but unstated 5-year period at Doubtful Island near
Albany, southwest Western Australia.
Only two of the ten Australian strandings listed by McKenna had enough
data associated with them to make a relatively positive determination of the
kind of animal involved. However, we include all of them here not only because
the data with them suggest that some or all could have been Pseudorca, but also
because we believe that it is important to note the relatively frequent reports of
such mass cetacean strandings along the shores of the southern half of Australia.
Appendix VII
In April, 1964, Mr. Georges Gilbert told Brown of an interesting observa-
tion on cooperative feeding behavior which had involved Pseudorca in the wild
at sea off the Kona coast of Hawaii, late in 1963.
Gilbert stated that he watched a large female false killer whale as she cap-
tured a very large mahimahi, or dolphin-fish (Coryphaena) , and then stop and
hold it in her mouth in order to allow her accompanying young to feed on it
by tearing large chunks of flesh from the fish.
On another occasion in Hawaii, Brown witnessed an autopsy of a large
male Pseudorca shortly after it was captured. The stomach of this animal con-
tained the remains of a large Coryphaena.
In October, 1965, Mr. Chris Varez told us of his experiences with Pseu-
dorca feeding behavior, also in Hawaii. He noted that in late 1963 or early
1964 he had often encountered this species in schools of some 10 to 50 animals
32
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No. 95
off the Kona coast. On one such occasion in particular he had been impressed
by the dramatic capture of a large 18- to 20-pound Coryphaena by a false killer
whale. The whale captured the Coryphaena and then extended the upper part
of its body out of the water with the large fish draped between its jaws, blood
from the fish streaming down the black sides of the whale. Varez then observed
the Pseudorca bite off and consume small pieces of the fish.
These observations, along with those cited in the main body of the text
above, clearly indicate that Pseudorca is a frequent predator on large pelagic
fishes.
Varez also noted that a captive male Pseudorca he had worked with at Sea
Life Park in Hawaii tended to break up the large fish it was fed before actually
eating them, as the wild one had done, although the captive seemed to prefer
having the large fish initially in order to follow this feeding pattern. Varez
stated that the false killer whale would accept a whole large fish, such as a
Coryphaena, and then shake it vigorously until the head broke off and most of
the entrails spilled out and also broke away. Then the whale would eat the flesh
in smaller pieces, but usually would leave the skin uneaten. It would also leave
uneaten the skin of large filets of marlin and other large fishes given it, eating
only the flesh. Apparently this behavior became an esthetic problem to the
establishment because the unsightly skin that remained would be allowed by the
whale to drift to the bottom of the tank. We know of no other observations on
delphinids so deliberately breaking up their food with the exception of two
rather old female Tursiops truncatus which often did so at Marineland of the
Pacific (also see Norris and Prescott, 1961: 312).
Despite the behavior of this false killer whale after receiving its food, Varez
was impressed with how gentle the whale was when it took the food from Varez’
hand. This behavior is like that of our animal at Marineland, and, also like ours,
the Hawiian captive Pseudorca would eat dead squid and small fish. However,
the Hawaiian annual later refused the squid and seemed to prefer fish (the
larger the fish, the better it seemed to be to his liking) .
Finally, Varez noted that the Hawaiian Pseudorca often carried a food fish
for as long as 3 or 4 hours before eating it, although he obviously intended to
do so. The Marineland false killer whale has also been observed to save the last
fish offered it at the end of a whale show and to carry it for a long time, chewing
on it and often dropping it near a dolphin tank-mate before grabbing it back as
if to tease the other animal.
CONTRIBUTIONS
IN SCIENCE
LOS
ANGELES
COUNTY
MUSEUM
Simber 96
07.9S
L L%&
A NEW PEROMYSCUS FROM THE LATE PLEISTOCENE OF
ANACAPA ISLAND, CALIFORNIA, WITH NOTES ON
VARIATION IN PEROMYSCUS NESODYTES WILSON
April 4
:
By John A. White
,
Los Angeles County Museum of Natural History • Exposition Park
Los Angeles, California 90007
A NEW PEROMYSCUS FROM THE LATE PLEISTOCENE OF
ANACAPA ISLAND, CALIFORNIA, WITH NOTES ON
VARIATION IN PEROMYSCUS NESODYTES WILSON
By John A. White1
Abstract: A new fossil rodent, Peromyscus anyapahensis,
is described from one of the California offshore Channel Islands.
The new form is compared to closely related forms.
Recently, through the efforts of Edward D. Mitchell, Jr., Jeri Lipps, Mi-
chael K. Hammer and James Valentine, specimens of a distinctive cricetine ro-
dent were collected in terrace deposits of the late Pleistocene of West Anacapa
Island. These specimens are referred to the genus Peromyscus because the cheek
teeth are brachyodont, the cusps are alternately arranged, the Mi are markedly
reduced, and the coronoid process of the mandible is only slightly developed.
These characters distinguish Peromyscus from most cricetine genera. The cheek
teeth in Reithrodontomys closely resemble those in Peromyscus. However, the
smallest of the Anacapa Island specimens is considerably larger than any known
specimen of Reithrodontomys creper, the largest-sized species of that genus
(Hooper, 1952: 175). Furthermore, in R. creper and allied species, accessory
cusps and lophs are invariably present (Hooper, 1952: 177-183), whereas in
the Anacapa Island species no such cusps and lophs are present.
Wilson (1936) described Peromyscus nesodytes from the Santa Rosa Is-
land Pleistocene of California. The latter species is characterized by” ... its
larger size, which is greater than in any living species of Peromyscus native to
the United States. . .” (Wilson, 1936: 408). Although clearly related to P.
nesodytes the Anacapa Island specimens are not referrable to that species and
differ to such a degree from mainland species that they should be placed in a
new species.
Peromyscus anyapahensis,2 new species
Figures 3 and 4
Holotype: Los Angeles County Museum of Natural History, Vertebrate Paleon-
tology no. 9205; left mandible with complete dentition and with angular process
and tip of coronoid process missing.
Locality : West Anacapa Island; Los Angeles County Museum of Natural His-
tory, Vertebrate Paleontology locality number 1764.
Age and stratigraphic position: Late Pleistocene terrace on the north side of the
island, 25 feet above sea level (See Lipps, 1964) .
iDepartment of Biology, California State College at Long Beach, and Research As-
sociate, Los Angeles County Museum of Natural History.
2Based on Anyapah, the Chumash word for Anacapa.
2
1966
New Fossil Rodent from California
3
Referred specimens: 8 mandibles with more or less complete dentitions and one
partial left palate with dentition, the labial half of M1 missing.
Diagnosis: Larger than in any living species except species of the subgenus
Megadontomys, Peromyscus zarhynchus, and P. nesodytes. Lower cheek teeth
similar to those of P. eremicus and P. calif ornicus but of marked larger size.
Significantly smaller than P. nesodytes and distinct from P. zarhynchus, and
species of P. ( Megadontomys ) in lacking accessory cusps and lophs in the
molars.
Description: Large size (Fig. 1); mental foramen anterior and slightly ventrad
of the anterior extension of masseteric scar; capsular process of lower incisor
ventrad of coronoid process and with ridge extending posteriad to lower one-
third of condyloid process; mandibular foramen dorsad to posterior projection
of cheek tooth row and approximately one-half the distance between posterior
surface of condyloid process and posterior edge of M3.
Anterior median cusp of Mi with a slightly developed furrow or groove on
the anterior face causing it to be slightly bipartite. A V-shaped valley exists
between the anterior median cusp and anterolingual cusp of M,. There are no
accessory cusps or lophs on the cheek teeth. The cusps on the cheek teeth tend
to remain recognizably high even when worn.
Comparisons with other species: The mandible in P. anyapahensis is signifi-
cantly shorter than in P. nesodytes (Fig. 1). The cheek teeth are smaller and
the anterior-median cusp in is divided by the presence of a groove that ex-
tends a shorter distance down the front face of tooth than in P. nesodytes. The
mental foramen is situated in front of the apex of the masseteric scar and not
variably placed as it is in P. nesodytes.
P. anyapahensis differs from both P. ( Megadontomys ) and P. ( Peromys-
cus) zarhynchus in the absence of extra cusps and lophs which are conspicuous
in the latter groups.
P. antiquus Kellog is as large in size as P. nesodytes, as noted by Wilson
(1936) but differs from both P. anyapahensis and P. nesodytes in having a rel-
atively larger M3 (Fig. 2) and in the relatively shorter anterior lobe of Mj.
The absence of extra cusps and lophs in the molars of P. anyapahensis
seems to favor the assignment of this species to the subgenus Haplomylomys.
Wilson ( 1936) and Hooper ( 1957) agree that the presence or absence of extra
cusps and lophs in Peromyscus is quite variable and should be used with caution
in characterizing subgenera. However, it is thought that since in both the above
island species no trace of such structures can be seen, their assignment to Hap-
lomylomys seems reasonable, especially since all other large-sized species have
these structures.
Materials
Most of the comparative materials used in this study are listed herein by
species and/ or subspecies together with region of collection, source institution
Figure! Bar diagrams modified from Hubbs and Hubbs (1953) showing variation in the alveolar length of the mandibular cheek
teeth. The numbers in parentheses indicate size of each respective sample. Measurements are in millimeters.
1966
New Fossil Rodent from California
5
AL
WM7
WMj
WMJ
F EDA C B
Figure 2. Ratio diagrams modified from Simpson et al. ( 1960) comparing mandibles
in Peromyscus anyapahensis new species (A) with P. nesodytes (B), P. thomasi
(C), P. zarhynchus (D), P. antiquus (E), and P. calif drnicus parasiticus (F). AL
= alveolar length of mandibular cheek teeth. The logs of the means of the dimen-
sions in P. anyapahensis are assumed to be zero, while the differences between the
log of the mean in the latter species (standard) and species being compared are
plotted to the positive ( + ) or negative ( — ) sides of the zero line.
(abbreviated as indicated under acknowledgments) and numbers of specimens.
Peromyscus nesodytes Wilson
Santa Rosa Island, California, SBM (7),L.A.M. (1).
P. thomasi Merriam
Guerrero, Mexico, MVZ (5).
P. zarhynchus Merriam
Chiapas, Mexico, KUM (19).
P. antiquus Kellog
Thousand Creek beds, Nevada, UCP ( 1 ) .
P. calif drnicus parasiticus (Baird)
San Francisco Bay Area, California, KUM (14), LACM (8).
Peromyscus nesodytes Wilson
Seven mandibles of this species are in the collections of the Santa Barbara
Museum of Natural History and are briefly discussed here since the species was
based originally on a single specimen.
The newly-available specimens confirm the “specific characters” proposed
by Wilson (1936). The position of the mental foramen, however, varies from
a lateral to a nearly dorsal position on the mandibles, probably reflecting the
6
Contributions in Science
No. 96
Figure 3. Peromyscus anyapahensis new species. Lingual (A) and buccal (C) views
of left mandible (approximately x 4.6), and occlusal view (B) of lower cheek teeth
(greatly enlarged) of holotype (LACM 9205).
usual high variability of species (Table 1 ) . Although present, the groove on the
anterior face of M) is not so strongly expressed as was inferred from the study
of the type specimen.
Remarks: The presence of large-sized Peromyscus on both Anacapa and Santa
Rosa Islands possibly indicates an adaptive trend toward a Neotoma-like habi-
tus. The fact that among several thousand specimens of rodents from Santa
1966
New Fossil Rodent from California
7
Figure 4. Peromyscus anyapahensis new species. Occlusal view (A) of RMi
(LACM 4588), and occlusal view (B) of left maxillary cheek teeth (LACM 4879).
Both A and B above are greatly enlarged and not to the same scale.
Table 1
Statistical data relating to four measurements on specimens of Peromyscus
anyapahensis new species, and P. nesodytes. AL indicates alveolar length of
mandibular cheek teeth; N — number of individuals in sample; X = arithmetic
mean; o- — standard deviation; V = coefficient of variability. All measure-
ments estimated to the nearest 0.01 mm.
Peromyscus anyapahensis new species
N
X
Max.
Min.
<T
V
AL
9
5.51
5.71
5.32
.1425
22.58
WAT
8
1.41
1.52
1.31
.0784
5.56
wm2
7
1.53
1.60
1.47
.0486
3.16
wm3
6
1.18
1.22
1.13
.0334
2.83
Peromyscus
nesodytes
AL
7
5.95
6.15
5.68
.1614
27.10
WMj
5
1.44
1.58
1.32
.1162
8.19
wm2
2
1.61
1.66
1.55
-
-
wm3
2
1.19
1.19
1.19
-
-
Rosa Island (Orr, 1962:419) no Neotoma bones were found, tends to sub-
stantiate the absence of Neotoma from the Pleistocene of Santa Rosa, and by
inference, from Anacapa Island as well. The numerous specimens of rodents
8
Contributions in Science
No. 96
from Santa Rosa Island consisted principally of Peromyscus, most of which
were small in size and resembling P. maniculatus. It seems probable that the
latter specimens are P. maniculatus streatori that inhabit the island. A specimen
of P. nesodytes was found in association with the pigmy mammoth ( Mammu -
thus [ Archidiscodon ] exilis),
P. anyapahensis seems to be intermediate, in size and in dental characters,
between P. nesodytes from Santa Rosa Island and P. eremicus and P. calif or-
nicus from the Recent of the mainland. This suggests that Santa Rosa Island
was separated from the mainland before Anacapa Island and that both islands
were connected to the mainland to the east.
Subsequent to 1936, biological surveys of the Channel Islands have been
conducted, especially of the living biota. The results of these surveys indicate
rather clearly that neither P. anyapahensis nor P, nesodytes have survivors on
Anacapa and Santa Rosa Islands, respectively.
Acknowledgments
For helpful suggestions, I wish to thank Seth Benson, C. A. MacLaughlin,
E. D. Mitchell, Jr., and R, A. Stirton. Theodore Downs and J. R. Macdonald
kindly read and criticized the manuscript. For permission to examine compara-
tive materials in their care I gratefully acknowledge Seth Benson, Museum of
Vertebrate Zoology (MVZ) University of California, Berkeley; J. Knox Jones,
Jr., Museum of Natural History, University of Kansas (KUM); J. R. Mac-
donald and C. A. MacLaughlin, Los Angeles County Museum of Natural His-
tory (LACM) ; Phil On*, Santa Barbara Museum of Natural History (SBM);
and R. A. Stirton, Museum of Paleontology (UCP), University of California,
Berkeley. Pamela Immel made the drawings, and I drafted the other figures.
Literature Cited
Hooper, Emmet T.
1952. A systematic review of the harvest mice (genus Reithrodontomys) of
Latin America. Misc. Publ. Mus. Zoo!., Univ. Michigan, 77:1-225.
1957. Dental patterns in mice of the genus Peromyscus . Misc. Publ. Mus. ZooL,
Univ. Michigan, 99:1-59.
Hubbs, Carl L. and Clark Hubbs
1953. An improved graphical analysis and comparison of series of samples.
Systematic ZooL, 2 (2) : 49-57.
Lipps, Jere II.
1964. Late Pleistocene history of West Anacapa Island, California. Bull. Geol.
Soc. Amen, 75:1169-1176.
Orr, P. C.
1962. The Arlington Spring site, Santa Rosa Island, California. Amer. Antiq-
uity, 27 (3) : 4 1 8-4 19.
Simpson, G. G., Anne Roe and R. C. Lewontin
1960. Quantitative Zoology; rev. ed.,: Harcourt, Brace and Co., 440 p., 64
text figs.
Wilson, Robert W.
1936. A new Pleistocene deer-mouse from Santa Rosa Island, California. J.
Mammal, 17 (4):408~410.
LOS
ANGELES
COUNTY
MUSEUM
CONTRIBUTIONS
¥i7h*< IN SCIENCE
] JMBER 97
May 5, 1966
A NEW SPECIES OF HETERANTHIDIUM FROM CALIFORNIA
(HYMENOPTERA: MEGACHILIDAE )
By Roy R. Snelling
Los Angeles County Museum of Natural History • Exposition Park
Los Angeles, California 90007
CONTRIBUTIONS IN SCIENCE is a series of miscellaneous technical papers
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David K. Caldwell
Editor
A NEW SPECIES OF HETERANTHIDIUM FROM CALIFORNIA
(HYMENOPTERA: MEG ACHILIDAE )
By Roy R. Snelling1
Abstract: Heteranthidium autumnale, a member of the ze-
bratum complex, is described from a single pair collected near
Desert Hot Springs, California. The complex is reviewed; H. tim-
berlakei is elevated to species level and H. subtimberlakei is syn-
onymized with it. Under H. zebratum are the synonyms H. z.
chippewaense, H. z. mississippi and H. cockerelli. A key to United
States species of Heteranthidium is given.
The following species is described at this time in order that the name may
be available for inclusion in a taxonomic study of the California Anthidiinae by
A. A. Grigarick and L. A. Stange, as a part of the California Insect Survey
series.
Heteranthidium autumnale, new species
Figure 1
This species may be separated from its congeners in both sexes by the fol-
lowing combination of characters: anterior coxae with distinct, sharp spines;
pronotal lobes with strong, high, sharp carina; mesopleurae immaculate. The
female is further characterized by the evenly convex, non-denticulate, anterior
clypeal margin.
Male: Length, 14 mm.; of forewing, 10 mm. Black, with yellowish white
maculae as follows: mandibles, except apical margin and teeth; labrum; cly-
peus, except reddish, translucent apical margin; roughly stirrup-shaped supra-
clypeal mark; lateral face marks, ending narrowly at level of lower margin of
anterior ocellus; underside of scape; stripe along lower, outer margin of eye for
about one-fourth eye length; irregular stripe behind upper one-fourth of eye;
narrow, irregular, medially interrupted stripe across top of vertex; minute spot
on anterior margin of pronotal lobe; short, broad stripe on mesoscutum adja-
cent to tegulae; comma-shaped mark on tegulae; minute spot on axillae; pair of
submedian spots on mesoscutellum; underside of anterior and middle femora,
two spots at apex of hind femora; basally and apically broadened stripes on all
tibiae; all basitarsi; spot on second segment of hind tarsi; medially attenuated
subapical bands on tergites I-V, those of I and II anteriorly emarginate, those
of III-V with free, or nearly free, lateral black marks; tergite VI largely yellow,
except extreme base, apical rim and pair of small, lateral, brownish spots; ter-
gite VII with a pair of preapical spots; sternites II and III with small lateral
spots. The following are ferruginous: legs, except for maculae noted above, the
ferruginous becoming very dark on the femora posteriorly; tarsal claws, tips
darker; all sternites entirely, except as noted above; extreme lateral margins of
tergites. Wings brownish-hyaline, veins and stigma dark brown.
1Preparator in Entomology, Los Angeles County Museum of Natural History.
1
2
Contributions in Science
No. 97
Figure 1. Heteranthidium autumnale Snelling, new species. A.-D., male sternites VIII,
VII, VI, V, respectively, vestiture removed from one side; E., male genitalia, left half
ventral aspect, right half dorsal aspect; F., male face; G., female face. F. and G.
drawn by Mrs. Evie Templeton, Los Angeles County Museum of Natural History.
1966
New California Bee
3
Head broader than long; facial breadth at level of clypeal base 0.86 times
distance from anterior ocellus to apex of clypeus; first flagellar segment slightly
longer than second; distance between posterior ocelli 1.2 times distance between
these ocelli and eyes, 0.9 times distance between ocelli and posterior margin of
vertex. Mandibles tridentate. Apical clypeal margin nearly straight, with three
low, rounded denticles medially, distance between denticles exceeding their
height; disc, when viewed from above, more strongly convex than in related
species, punctures coarse, deep, contiguous, integument shining between punc-
tures; punctures of remainder of head mostly a little finer, more irregular, more
dense.
Carina of pronotal lobes high, height equal to about one-third times an
ocellar diameter; punctures of mesocutum, axillae and mesoscutellum deep,
smaller than those of clypeus, very dense, so that surface appears subopaque,
granulose; upper half of anterior face of mesepisternum distinctly, rather close-
ly punctate, a few smaller punctures present, lower half shinier, very sparsely
punctate; lateral faces with dense, coarse punctures, integument shining; tegulae
with fine, dense punctures; basal area of propodeum densely, rugosely punctate;
integument minutely tessellate; discal and lateral areas of propodeum with
punctures a little finer and less dense. Spine at apex of anterior coxae about as
long as basal width; middle tibiae broad, a little more than twice as long as
broad; middle basitarsi about five times longer than wide, equal in length to
hind basitarsi, which are about four-fifths as long as the hind tibiae.
Abdominal tergites with narrow, impunctate apical margins, the discal
punctures finer than those of mesoscutum, separated by about one-half a punc-
ture diameter; pygidial carina, in profile, distinctly concave; apical margin of
pygidium distinctly convergent toward the apical process formed by the median
carina; sternite III with a pair of low, medio-posteriorly directed blunt tuber-
cules on each side of mid line, near apical margin; sternites V-VIII hidden.
Female: Length, 14 mm.; of forewing, 9 mm. Black, with pale yellow mac-
ulae as follows: obscure preapical blotch on outer side of mandibles; clypeus,
except apical margin and sutural infuscations; triangular lateral face marks
ending slightly below level of anterior ocellus; short stripe behind upper third
of eye; irregular, but rather broad, stripe across summit of vertex; obscure spot
on outer, anterior corner of pronotal lobes; anterior two-fifths of tegulae; broad
stripe along sides of mesoscutum; large spots on axillae and mesoscutellum;
spot at base, and another, obscure, spot about three-fourths along length of the
anterior tibiae; basal spot on middle and hind tibiae; narrow stripe on basal
three-fifths of hind basitarsi; medially narrowed, subapical bands of tergites
I-V; tergite I with completely enclosed black spot on each side; tergites II-IV
with anterior black emarginations of the bands on each side; tergite VI with a
roughly butterfly-shaped macula. Legs, except maculae noted above; piceous,
the apical tarsal segments and tarsal claws ferruginous; tibial spurs very dilute
ferruginous.
Head broader than long; facial breadth at level of clypeal base 0.9 times
4
Contributions in Science
No. 97
distance from anterior ocellus to apex of clypeus; distance between posterior
ocelli 1.1 times distance between ocelli and eyes, 0.9 times distance from ocelli
to posterior margin of vertex. Apical clypeal margin gently, evenly convex,
without median denticles, discal punctures coarse, shallow, contiguous; supra-
clypeal and baso-medial area of clypeus rather strongly arched; disc of clypeus,
in profile, nearly flat. Punctures of frons variable, but generally finer and more
dense than those of clypeus, especially laterally, immediately below anterior
ocellus they are rather coarse, with flattened interspaces; punctures of vertex
uniform, dense, finer than those of clypeus.
Thoracic punctation similar to that of male. Middle basitarsi about three
times longer than broad, three-fourths as long as hind basitarsi, the latter a little
over twice as long as broad, about four-fifths as long as their femora.
Abdominal punctation very similar to that of male; tergite VI, very
densely, finely, punctate, surface opaque; scopa ferruginous medially, becom-
ing paler basad and laterad.
Holotype male and allotype female from 3 miles northeast of Desert Hot
Springs, Riverside County, California, 8 November, 1963, collected by *R. Mc-
Diarmid. Mr. McDiarmid has informed me ( personal comm.) that these speci-
mens were flying around an unknown composite which was most likely a species
of Chrysothamnus. The types are deposited in the collections of the Los Ange-
les County Museum of Natural History.
This species belongs to the H. zebratum (Cresson) complex, in the pres-
ent taxonomic treatment of which there is much confusion. Since my in-
terpretation of the various taxa involved differs from previous ones, it is advis-
able to review the development of the old arrangement. The first species to be
described was H. zebratum, described by Cresson (1872) from both sexes,
based on material from Texas and Colorado. Titus ( 1902) redescribed the male
as Protanthidium cockerelli. When he erected the genus Heteranthidium, Cock-
erell ( 1904) was not aware that this species was congeneric with the type of the
genus, H. dorsale (Lepeletier) . In 1910, Graenicher described Anthidium
( Protanthidium ) chippewaense from material of both sexes taken in Wisconsin.
When Schwarz (1926) reviewed the genus, he included the above names
and presented a key to the species as he understood them at that time. He sub-
sequently (1928) described two additional members of this complex from Cali-
fornia, H. timberlakei and H. subtimberlakei. Michener (1947) added H. z .
mississippi from material collected in Mississippi. In the Catalog of Nearctic
Hymenoptera Michener (1951) arranged the series as follows:
H. zebratum chippewaense (Graenicher). Wisconsin
H. zebratum mississippi Michener. Mississippi
H. zebratum subtimberlakei Schwarz. California
H. zebratum timberlakei Schwarz. California
H. zebratum zebratum (Cresson). South Dakota, Nebraska, Colorado,
New Mexico, Texas.
synonym: Protanthidium cockerelli Titus.
1966
New California Bee
5
Mitchell (1962) was able to show that H. z. chippewaense, H. z. mississippi
and H. z. zebratum belong to a single, highly variable species, and accordingly
synonymized the former two with the latter.
Since Dr. Stange has advised me {in lift.) that H. z. subtimberlakei is a
synonym of H. z. timberlakei, there is no necessity for further reference to the
former name. I here elevate H. timberlakei to species status, arranging the
members of this complex as follows:
H. autumnale Snelling. Colorado Desert, California.
H. timberlakei Schwarz. San Diego County to Trinity County, California,
synonym: H. subtimberlakei Schwarz.
H. zebratum (Cresson). Rocky Mountains, east to Michigan and Missis-
sippi.
synonyms: H. z. mississippi Michener, H. z. chippewaense (Graenicher) ,
H. cockerelli (Titus).
Both H. autumnale and H. timberlakei differ from H. zebratum in a num-
ber of important features. The former two species both possess well-developed
spines on the anterior coxae; these are lacking in H. zebratum. The pronotal
lobes of H. zebratum are characteristically without a trace of a carina at the
juncture of the anterior and dorsal surfaces. In H. autumnale and H. timber-
lakei these surfaces meet at a distinct angle (evenly rounded in H. zebratum) ;
in H. timberlakei a fine, but distinct carina is present at this juncture, while in
H. autumnale the carina is very high and strong, almost lamellate in appear-
ance. In the latter two species, the basic outline of the apical clypeal margin is
evenly convex; in H. zebratum the apex is slightly angulate, with a median
convexity. In the males there are slight, but consistent differences in the apical
sternites and genitalia. Finally, it should be pointed out that there is no evidence
to show that the California forms intergrade with H. zebratum. The present
writer inclines to the view that there is a complex of several closely related,
allopatric species.
The following key should assist in the recognition of the various species
currently known to occur in the United States. Two species, H. cordaticeps
Michener and H. fontemvitae Schwarz are placed in the key on the basis of
descriptions alone; the male of H. cordaticeps is unknown at the present.
KEY TO UNITED STATES SPECIES OF HETERANTHID1UM
Males
1. Thorax to some degree maculate; abdominal tergites with bands subapical
in position 2
Thorax entirely black, immaculate; abdominal tergites with narrow, ivory-
colored, apical bands H. ridingsi (Cresson)
2. Third abdominal sternite without spines, at most with a pair of blunt, flat-
tened tubercles on either side of midline; pygidium not distinctly tridentate
6
Contributions in Science
No. 97
at apex 3
Third abdominal sternite with a pair of short, dark, slender spines; py-
gidium distinctly tridentate apically H. fontemvitae Schwarz
3. Pronotal tubercles rounded above, without a trace of a carina 4
Pronotal tubercles with at least a fine carina above, frequently sharply
crested 5
4. Anterior coxae with short, blunt, flattened apical tubercles; pygidial apex
emarginate; mesoscutum usually with L-shaped marks
H. occidentale (Cresson)
Anterior coxae usually unarmed, at most with a very low, conical tubercle;
pygidial apex pointed or bluntly rounded, not medially emarginate; meso-
scutum usually immaculate, rarely with short lateral stripes near tegulae . .
H. zebratum (Cresson)
5. Outer hind tibial spur thick, strongly curved near apex; apical process of
first sternite high, longitudinally impressed, apical margin concave
H. dor sale (Lepeletier)
Outer hind tibial spur slender, straight; first sternite without prominent
apical elevation 6
6. Anterior face of mesepisternum strongly shining, very sparsely punctate;
anterior margin of clypeus concave between two prominent teeth on each
side of midline; apical margin of pygidium with sinuate, transparent apical
lip H. larreae (Cockerell)
Anterior face of mesepisternum strongly roughened and/or punctate, at
least on upper half; anterior margin of clypeus broadly convex, if slightly
concave medially, then flanking teeth reduced to minute denticles; apical
margin of pygidium variable 7
7. Hind basitarsi unusually short, about one-third as long as hind tibiae,
shorter than following tarsal segments combined; apical margin of seventh
tergite very slightly convex, appearing almost straight
H. crassipes (Cresson)
Hind basitarsi well over half as long as hind tibiae, distinctly longer than
following segments combined; apical margin of seventh tergite distinctly
convex, usually slightly produced medially 8
8. Pygidial carina variable; clypeus sparsely pubescent 9
Pygidial carina, in profile, concave; clypeus densely pubescent
H. bequaerti Schwarz
9. Pygidial carina, in profile, nearly straight; apical margin of pygidium even-
ly convex, the median carina forming a slightly projecting nipple; pronotal
carina low and fine, but distinct; apical margin of clypeus with four to six
strong teeth, the clypeal disc strongly shining between the coarse, rather
shallow punctures H. timberlakei Schwarz
Pygidial carina, in profile, distinctly concave; apical margin of pygidium
distinctly convergent toward the prominent apical process formed by the
1966
New California Bee
7
apex of the median carina; pronotal carina high and sharp; apical margin
of clypeus with teeth low and rounded, the clypeal disc moderately shining
between coarse, deep punctures H. autumnale Snelling
Females
1 . Vertex with broad, deep median indentation, so that head appears some-
what heart-shaped; integument reddish-brown, with reddish maculae; pro-
notal carina strong H. cordaticeps Michener
Vertex without indentation, head not at all cordate; integument largely
black, with variable maculae; pronotal carina present or absent 2
2. Head, thorax and legs immaculate, abdominal tergites with pale, narrow,
apical fasciae; pronotal lobes non-carinate H. ridingsi (Cresson)
Head and thorax maculate; abdominal bands subapical in position, usually
distinctly yellowish; if pale whitish, then broad; pronotal lobes with or
without carina 3
3. Pronotal lobes non-carinate, usually rounded above (the summit of the
anterior face is sometimes distinctly angulate in H. occidentale, but a carina
is not present) . 4
Summit of anterior face of pronotal lobes sharply angulate and marked by
a distinct, though sometimes fine, carina 5
4. Anterior coxae without apical spine, rarely with a very small conical tu-
bercle; sixth tergite, in profile, very strongly convex; pronotal lobes always
rounded, without angulation at summit of anterior face
H. zebratum (Cresson)
Anterior coxae with distinct, flattened apical spine; sixth tergite, in profile
gently, gradually convex; pronotal lobe sometimes strongly angulate at
summit of anterior face, never evenly and smoothly rounded as above ....
H. occidentale (Cresson)
5. Clypeus black, immaculate, no supraclypeal spot; clypeus decidedly con-
cave in profile toward apex, apex with four distinct, close-set teeth
H. bequaerti Schwarz
Clypeus yellow, except apical margin, supraclypeal spot usually present;
clypeus usually evenly convex in profile, sometimes slightly flattened to-
ward apex 6
6. Mesoscutum with a pair of elongate median stripes; tibiae wholly yellow
externally 7
Mesoscutum without elongate median stripes; tibiae usually with yellow
stripes externally 8
7. Outer spur of hind tibiae thick, abruptly bent near apex; abdominal
punctation dense, strong; propodeum entirely black
H. dor sale (Lepeletier)
Outer spur of hind tibiae slender, slightly curved apically; abdominal punc-
tures sparse; propodeum maculate H. larreae (Cockerell)
8
Contributions in Science
No. 97
8. Mesopleurae immaculate; earina of pronotal lobes very strong; anterior
clypeal margin evenly rounded, without denticles; spines of anterior coxae
long, slender, sharply pointed H. autumnale Snelling
Mesopleurae distinctly maculate; earina of pronotal lobe usually low, fine;
anterior clypeal margin with distinct denticles; anterior coxal spines rather
broad, flattened 9
9. Abdomen semi-opaque, with dense, distinct punctation; mandibles yellow,
except along apical edge; tibiae red, with yellow maculae; abdominal bands
anteriorly emarginate on each side of midline . . . H. fontemvitae Schwarz
Abdomen shining between large, moderately dense punctures; mandibles
variable; tibiae black with yellow maculae; abdominal bands not anteriorly
emarginate 10
10. Mandibles and sixth tergite black, immaculate; earina of pronotal lobes
strong, sharp; hind basitarsi almost as broad as their tibiae, about twice as
long as broad; erect pubescence of mesoscutal disc little, if any, longer than
diameter of anterior ocellus H. crassipes (Cresson)
Mandibles and sixth tergite maculate; earina of pronotal lobes low, fine;
hind basitarsi distinctly narrower than their tibiae, about 2.5 times longer
than broad; erect pubescence of mesoscutal disc much longer than diame-
ter of anterior ocellus H. timberlakei Schwarz
Literature Cited
Cockerell, T. D. A.
1904. New genera of bees. Ent. News, 15 : 292.
Cresson, E. T.
1872. Hymenoptera Texana. Trans. Amer. Ent. Soc., 4: 153-292.
Graenicher, S.
1910. Wisconsin bees — new and little known species. Canad. Ent., 42: 101-
104, 157-160.
Michener, C. D.
1947. Bees of a limited area in southern Mississippi. Amer. Midi. Nat., 38:
443-455.
1951. in Muesebeck, et al„ Hymenoptera of America North of Mexico. Synop-
tic Catalog. U.S. Dept. Agricul. Monog. 2, 1420 p.
Mitchell, T. B.
1962. Bees of the Eastern United States. II. North Carolina Agricul. Exper.
Sta., Tech. Bull. 152, 1-577.
Schwarz, H. F.
1926. North American bees of the genus Heteranthidium. Amer. Mus. Novi-
tates, 218: 1-16.
1928. Bees of the subfamily Anthidiinae, including some new species and va-
rieties, and some new locality records. J. New York Ent. Soc., 36: 369-
418.
Titus, E. S. G.
1902. Three new Anthidiinae from Colorado. Ent. News, 13: 169-171.
LOS
ANGELES
COUNTY
MUSEUM
IlJMBER 98
CONTRIBUTIONS
tl'lU IN SCIENCE
May 5, 1966
STUDIES ON NORTH AMERICAN BEES
OF THE GENUS HYLAEUS
1. DISTRIBUTION OF THE WESTERN SPECIES
OF THE SUBGENUS PROSOPIS WITH DESCRIPTIONS
OF NEW FORMS (HYMENOPTERA: COLLETIDAE)
By Roy R. Snelling
Los Angeles County Museum of Natural History • Exposition Park
Los Angeles, California 90007
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STUDIES ON NORTH AMERICAN BEES
OF THE GENUS HYLAEUS.
1. DISTRIBUTION OF THE WESTERN SPECIES
OF THE SUBGENUS PROSOPIS WITH DESCRIPTIONS
OF NEW FORMS (HYMENOPTERA: COLLETIDAE)
By Roy R. Snelling1
Abstract: In this paper representatives of the subgenus
Prosopis are considered. A key to the species occurring in the
western United States is given. The following new synonyms are
made: H. dunningi and H. albertensis — H. affinis; H. universita-
tis = H. r. rugulosus; The following are assigned subspecific rank
under H. rugulosus : H. coquilletti, H. episcopalis, H. giffardiellus.
Two new forms are described: H. r. metzi (cismontane southern
California) and H. insol itus (southern Arizona to western Texas
and northern Mexico). H. transvittatus, described from central
Mexico, is recorded from the United States (Arizona) for the first
time.
This paper is the first of a continuing series devoted to this genus, and is
concerned with the representatives of the subgenus Prosopis. The members of
this subgenus are probably the most common and widely distributed of the
genus. Accordingly, there has been considerable confusion of the forms occur-
ring in the western United States, and the bibliographies of the various forms
are rather complicated.
During the course of this study considerable material from the western
states has been examined, and I wish to express my gratitude to each of the
following for making available to me the specimens in his care: H. Dietrich,
Cornell University (CU); H. E. Evans, Museum of Comparative Zoology
(MCZ); A. R. Gittins, University of Idaho (UI); P. D. Hurd, Jr., California
Insect Survey, University of California at Berkeley (CIS); K. V. Krombein,
United States National Museum (USNM); W. E. LaBerge, University of Ne-
braska (UN) ; U. N. Lanham, University of Colorado (UC) ; H. B. Leech, Cal-
ifornia Academy of Sciences (CAS); A. T. McClay, University of California
at Davis (UCD) ; C. D. Michener, University of Kansas (UK) ; H. E. Milliron,
Canadian National Collection (CNC); J. G. Rozen, Jr., American Museum of
Natural History (AMNH); F. G. Werner, University of Arizona (UA). Spec-
imens in the collections of the Los Angeles County Museum of Natural History
are indicated by (LACM).
In order to facilitate the identification of the several species of this sub-
genus found in the western states, I present the following key. Members of this
1Preparator in Entomology, Los Angeles County Museum of Natural History.
1
saiutsuiv
kM#hm
2
Contributions in Science
No. 98
subgenus may be recognized by the characters indicated in a paper to be pub-
lished elsewhere.
KEY TO WESTERN SPECIES OF SUBGENUS PROSOPIS
1. Antennae thirteen-segmented; abdomen with seven visible segments; cly-
peus wholly yellow; males 2
Antennae twelve-segmented; abdomen with six visible segments; clypeus
black, sometimes yellow maculate; females 6
2. Integument of propodeal triangle densely punctate, opaque, longitudinal
rugulae obsolescent; punctures of propodeal sides large, distinct, inter-
spaces less than a puncture diameter; facial quadrangle 1.8 times longer
than broad at level of clypeal base; mandible black
transvittatus (Cockerell)
Integument of propodeal triangle not densely punctate, shining between
the well-developed rugulae; punctures of propodeal sides generally small,
indistinct, integument rather strongly shining or else conspicuously rough-
ened; facial quadrangle variable; mandibles at least partly yellow 3
3. Underside of antennal scape with conspicuous yellow macula and lateral
face marks ending broadly about halfway between level of antennal sockets
and tops of eyes; propodeal sides conspicuously roughened between indis-
tinct punctures; facial quadrangle 1.8- 1.9 times as long as broad at level
of clypeal base affinis (F. Smith)
Underside of antennal scape as a rule inconspicuously maculate; if with
distinct macula, then lateral face marks ending near tops of eyes; propo-
deal sides usually with perceptible punctures, somewhat shining; facial
quadrangle variable 4
4. Rugulae of propodeal triangle not extending to base of declivity; facial
quadrangle 1.5 times as long as broad at level of clypeal base; thoracic
punctures large, distinct, well separated insolitus, new species
Rugulae of propodeal triangle extending to base of declivity; facial quad-
rangle variable, but at least 1.6 times as long as broad at clypeal base; tho-
racic punctures small, frequently with their margins obscure, often crowd-
ed, especially on mesoscutum 5
5. Facial quadrangle 1.6- 1.8 times longer than broad at clypeal base; frons
opaque, with punctures obscured by strongly roughened integument; lat-
eral face marks usually ending at level of upper margin of antennal sock-
ets; punctures of propodeal sides usually obscured by roughened, dull in-
tegument modestus citrinifrons (Cockerell)
Facial quadrangle 1. 9-2.0 times longer than broad at clypeal base; frons
somewhat shining, distinctly punctate; lateral face marks variable; punc-
tures of propodeal sides usually distinct, the integument somewhat shining
rugulosus (Cockerell)
1966
Bees of the Genus Hylaeus
3
6. Rugulae of propodeal triangle obsolescent apically; integument of entire
thorax dull, interspaces between punctures very densely tessellate; clypeus
with transverse apical whitish macula transvittatus (Cockerell)
Rugulae of propodeal triangle usually extending to base of declivity; if
obsolescent apically, integument of sides of thorax distinctly shining be-
tween punctures; clypeal maculae variable 7
7. Clypeus with broad, median, longitudinal macula; facial foveae ending
about midway between eyes and ocelli; facial quadrangle 1. 1-1.2 times
longer than broad at clypeal base insolitus, new species
Clypeus usually immaculate, sometimes with preapical spot; facial foveae
ending much nearer eyes than ocelli; facial quadrangle at least 1.4 times
longer than broad at clypeal base 8
8. Facial quadrangle 1.4 times longer than broad at clypeal base; frons dense-
ly tessellate and slightly roughened so that the dense punctures are ob-
scured, integument very dull, appearing granulose; lateral face marks,
when present, linear, separated from clypeal margin, except below, ending
below level of upper margin of antennal sockets
modestus citrinifrons (Cockerell)
Facial quadrangle 1.5 times as long as broad at clypeal base, or longer;
frons tessellate but somewhat shining, integument not conspicuously
roughened, punctures distinct; lateral face marks variable, but usually
broadly contiguous with lateral clypeal margin 9
9. Lateral face marks triangular, ending at or below level of upper margin of
antennal sockets, completely filling space between eye margin and lateral
margin of clypeus; propodeal sides dull, rather coarsely roughened between
obscure punctures; preapical area of clypeus black .... affinis (F. Smith)
Lateral face marks linear, usually extending well above level of upper mar-
gin of antennal sockets; if shorter, contiguous with lateral clypeal margin
at lower end only; propodeal sides variable, dull to moderately shining,
punctures distinct or obscured by fine roughening; preapical area of clype-
us suffused with reddish, or distinctly maculate . . . rugulosus (Cockerell)
Hylaeus ( Prosopis ) affinis (F. Smith)
Figures 1A, 2A
Prosopis affinis Smith, 1853. Cat. Hym. Brit. Mus., 1:24. 9, not 8 .
Prosopis ziziae Robertson, 1896. Canad. Ent., 28:136. $ 8. Metz, 1911.
Trans. Amer. Ent. Soc., 37:130.
Prosopis ziziae, race dunningi Cockerell, 1898. Ent., 31:188. 8. Metz,
1911. Op. cit. : 130 NEW SYNONYM.
Hylaeus dunningi, Cockerell and Sumner, 1931, Amer. Mus. Nov., 490:7.
Hylaeus albertensis Cockerell, 1937. Canad. Ent., 69:126. 8 9. NEW
4
Contributions in Science
No. 98
E F
Figure 1. Hylaeus ( Prosopis ) spp., faces of males: A, H. affinis (F. Smith); B, H.
modestus citrinifrons (Cockerell); C, H. r. coquilletti (Cockerell); D, H. r. episco-
palis (Cockerell); E, H. r. giffardiellus Cockerell; F, H. r. rugulosus (Cockerell).
Illustrations by Miss Dorothy Kresch, Los Angeles County Museum of Natural
History.
1966
Bees of the Genus Hylaeus
5
SYNONYM.
Metz recognized Cockerell’s name, H. dunningi, as a variety of this species
which he called P. ziziae. Twenty years later Cockerell and Sumner (1931:7)
accorded specific standing to the name without giving any reason for so doing.
Cockerell ( 1937: 126) again renamed the species, as H. albertensis, apparently
based upon a single pair from Edmonton, Alberta. Examination of series from
the western United States and Canada shows that the supposed separative char-
acters (all based upon markings) will not hold true, and it therefore seems in
order to reduce both of Cockerell’s names to synonyms of H. affinis.
New distribution data for the species are as follows: United States: Idaho :
Canyon Co.: 1 8,3 9$, Melba, 8 mi. N., VI-18-1957 (H. W. Homan) , carrot
flowers; 1 9, same data except VI-28-1957; 1 9 , Wilder, VI-27-1945 (W. E.
Shull; all UI), on peas. Washington : Garfield Co.: 2 8 8, Pullman, VII-4-1957
(W. H. Lange; UCD). Canada: Manitoba : 3 8 8, Shilo, 5 mi. SW., VI-16-
1958 (C. D. F. Miller) ;1 8,1 9 , Brandon, 6 mi. NW, VII-1-1958 (C. D. F.
Miller); 2 8 8, 2 9 9, Virden, VII-8-1953 (Brooks & Kelton); 1 8 Carberry,
VI-18-1953 (Brooks & Kelton) ; 1 8 , same locality and collectors, VI-23- 195 3;
1 8 , Horton, VII-25-1953 (Brooks & Kelton, all CNC). Saskatchewan : 1 8 ,
Chaplin, VI-22-1959 (A. R. Gittins; UI); 1 8, Parkbeg, VI-22-1959 (A. R.
Gittins; UI), on Melilotus\ 1 8 , St. Victor (49°20'-105o54'), VI-27-1955 (J.
R. Vockeroth) ; 2 8 8, White Fox, VII-8-1950 (L. A. Konotopetz) ; 1 9 , White
Fox, VII-10-1944 (O. Peck). British Columbia : 1 8 , Fairview, V-19-1919 (E.
R. Buckell, all CNC).
Hylaeus ( Prosopis ) modes tus citrinifrons (Cockerell)
Figures IB, 2B
Prosapis citrinifrons Cockerell, 1896. Psyche, 7 (supl.):27. 8 .
Prosopis modestus, Metz, 1911. Trans. Amer. Ent. Soc., 37:121-129. (in
part).
Hylaeus modestus citrinifrons, Cockerell and Sumner, 1931. Amer. Mus.
Nov., 490:6. 9. (in part).
When Cockerell established this form, as a full species, he used the absence
of maculae on the pronotal collar as a primary distinctive character. Metz
(1911:127) considered the form unworthy of recognition and synonymized it
under H. modestus Say; he evidently examined only a few males from Colorado
in reaching this conclusion. The name was revived in the present combination,
by Cockerell and Sumner (1930:6), but no new characters were cited.
Although this bee is superficially similar in size and markings to H. r.
rugulosus (Cockerell), it differs in the distinctly broader face, a character
which is consistent for both sexes. Further, females of this form have the punc-
tures of the frons so dense that the integument is opaque in appearance; in
females of H. rugulosus the punctures are close, but with shining interspaces.
In the males of H. m. citrinifrons this character is somewhat more variable, and
6
Contributions in Science
No. 98
E F
Figure 2. Hylaeus ( Prosopis ) spp., faces of females: A, H. affinis (F. Smith); B, H.
modestus citrinifrons (Cockerell); C, H. r. coquilletti (Cockerell); D, H. r. episco-
palis (Cockerell); E, H. r. giffardiellus Cockerell; F, H. r. rugulosus (Cockerell).
Illustrations by Miss Dorothy Kresch, Los Angeles County Museum of Natural
History.
1966
Bees of the Genus Hylaeus
7
hence less useful diagnostically. Cockerell relied principally upon the non-
maculate pronotal collar in the recognition of this form, a feature which is too
variable to be of any use. Material now before me includes specimens of both
sexes, many of which have well-developed maculae on the pronotal collar; the
presence or absence of these maculae apparently cannot be correlated with dis-
tribution, although individuals from more northern regions, or higher eleva-
tions, generally tend to be somewhat darker. The western populations do differ,
however, from those of the midwestern and eastern United States in the re-
duced maculae, duller and less distinctly punctured frons and duller mesopleu-
rae.
As currently understood, the range of this form extends from the southern
Rocky Mountains in New Mexico, northward through Colorado, Wyoming,
Montana and Alberta, thence westward to the Pacific Coast. Southern exten-
sions follow the Cascade, Sierra Nevada and Pacific Coast Ranges into Cali-
fornia. In the latter Range, H. m. citr ini frons occurs at least as far south as
Los Angeles County. In eastern Colorado, Wyoming and Montana intergrada-
tion with the nominate form presumably occurs, but too little material is avail-
able from this area to verify this assumption.
Hylaeus ( Prosopis ) rugulosus (Cockerell)
This is the common western species which has long been known as H. epis-
copalis (Cockerell). However, an examination of considerable material from
the Rocky Mountain region and the western states indicates that H. episcopalis
is best regarded as a subspecies of H. rugulosus. Both forms occur over large
areas of the western states. The nominate form has a more northern distribu-
tion, but follows the Rocky Mountains south to New Mexico, and the Cascades
and Sierra Nevada into montane central California, where it is common in the
Transition and Boreal Zones. The periphery of its range broadly overlaps that
of H. r. episcopalis and specimens from these overlapping areas are difficult to
determine to subspecies.
The situation is further complicated by the presence of three additional
forms which are structurally very similar to H. rugulosus. Because all of these
are considered to intergrade with H. r. rugulosus and H. r. episcopalis they are
here regarded as subspecies. One of these, H. r. giffardiellus Cockerell, presents
relatively few problems since its range is confined to the Sacramento and San
Joaquin Valleys of California, and is thus the most precinctive of the five sub-
species currently recognized. In keeping with this restricted habitat, this sub-
species intergrades only with the subspecies H. r. episcopalis in the surrounding
foothill areas.
Another subspecies, H. r. coquilletti (Cockerell), occurs throughout the
Great Basin Region, south to Baja California, Mexico, thence eastward to
western Texas. This form is a conspicuous member of the Lower Sonoran bee
fauna of the southwestern area and is perhaps more nearly deserving of specific
rank than any of the other forms.
8
Contributions in Science
No. 98
In the area around Flagstaff, Coconino County, Arizona, a form is found
which is quite difficult to assign to any of the currently recognized subspecies.
Although these bees probably represent an aberrant population of H. r. coquil-
letti, they are not readily separable from H. r. giffardiellus. However, the great
spatial barrier separating the two populations makes the latter assignment most
unattractive. For the present, therefore, they are left unassigned until such a
time as the entire complex can be critically examined.
A key to the currently recognized forms of H. rugulosus is presented be-
low. In the following discussion of distributions, no California records are in-
cluded since these will be given, in full, in a forthcoming treatment of the Cali-
fornia species of this genus.
KEY TO SUBSPECIES OF HYLAEUS RUGULOSUS (COCKERELL)
1. Antennae twelve-segmented; abdomen with six visible segments; females
2
Antennae thirteen-segmented; abdomen with seven visible segments; males
6
2. Lateral face marks extensive, lower marks filling lateral area, upper marks
extending beyond upper margin of antennal sockets 3
Lateral face marks more restricted, rarely, if ever, filling lower lateral areas;
upper marks terminating at or below upper margin of antennal sockets . . 5
3. Pubescence of tergites III and IV entirely pale, that of III no sparser on disc
than laterally; clypeus frequently with preapical whitish macula; posterior
basitarsi usually with small basal spot; deserts and Great Basin
; r. coquilletti (Cockerell)
Pubescence of tergites III and IV largely or entirely dark fuscous; clypeus
black apically, with preapical reddish area or with yellow macula; posterior
basitarsi entirely black 4
4. Maculae bright lemon-yellow; preapical area of clypeus sometimes with
yellow macula, more frequently with transverse reddish area; upper lateral
marks extending well above upper margin of antennal sockets; pronotal col-
lar largely yellow; anterior tibiae with pale stripe on basal half; pubescence
of tergite III largely or entirely appressed, sparser on disc than on sides; San
Joaquin and Sacramento Valleys of California . . . r. giffardiellus Cockerell
Maculae cream color or yellowish; preapical area of clypeus usually black;
upper lateral marks extending little above margin of antennal sockets; pro-
notal collar no more than 50% pale above; anterior tibiae with small basal
spot; tergite III with pubescence largely erect, as dense discally as laterally;
cismontane southern California r. metzi, new subspecies
5. Pronotal collar black or with small lateral maculae shorter than distance be-
tween them; tubercules no more than one-third pale; erect pubescence of
tergite III fuscous; maculae distinctly yellowish . . r. rugulosus (Cockerell)
1966
Bees of the Genus Hylaeus
9
Pronotal collar with lateral maculae longer than the distance between them;
tubercules usually at least 50% pale; erect pubescence of tergite III variable,
but usually pale; maculae whitish or cream-color . r. episcopalis (Cockerell)
6. Lateral face marks extending three-fourths or less of eye length, not widen-
ed above antennal sockets; maculae distinctly yellowish; maculae of pronotal
collar and lobes frequently reduced or absent; hind tibial stripe extending
one-half or less the length of the tibia 7
Lateral face marks extending four-fifths or more of eye length, conspicuous-
ly widened above antennal sockets; maculae cream-colored; maculae of
pronotal collar and lobes conspicuous, latter including three-fourths or more
of lobes; hind tibial stripe extending three-fourths or more the length of the
tibiae r. coquilletti (Cockerell)
7. Pronotal maculae reduced, distance between collar maculae exceeding one-
half the lengths of the maculae, pronotal lobes less than 50% pale; erect
hairs of apical tergites black or fuscous, sparser discally than laterally; apical
margin of second little, if at all, reflexed r. rugulosus (Cockerell)
Pronotal maculae usually separated by less than half their lengths; pronotal
lobes more than 50% pale; erect hairs of apical tergites frequently pale,
often as dense discally as laterally; apical margin of second tergite usually
somewhat reflexed 8
8. Lateral face marks bright lemon-yellow, completely filling space between
eyes and clypeus, extending upward about three-fourths length of the eyes;
clypeus frequently with preapical yellow spot; basal half of front tibiae
maculate r. giffardiellus Cockerell
Lateral face marks pale yellowish, frequently not filling space between eyes
and clypeus, usually ending little above level of upper margins of antennal
sockets; clypeus without preapical yellow spot, suffused with reddish pre-
apically; no more than basal one-third of front tibiae maculate 9
9. Punctures of second tergite coarse, close, usually separated by less than a
puncture diameter; erect pubescence of tergites four to six as dense medially
as laterally; apical margin of second tergite rather strongly reflexed
r. metzi, new subspecies
Punctures of second tergite rather fine, separated by more than a puncture
diameter; erect pubescence of tergites four to six sparser medially than later-
ally; apical margin of second tergite only slightly reflexed
r. episcopalis (Cockerell)
Hylaeus rugulosus rugulosus (Cockerell), new status
Figures IF, 2F
Prosapis rugulosus Cockerell, 1896. Psyche, 7 (supl.): 28. $.
Prosapis rugulosus var. fallax Cockerell, 1896. Op. cit .: 28. $ .
Prosopis universitatis Cockerell, 1906. Ann. Mag. Nat. Hist., (7), 17:225.
10
Contributions in Science
No. 98
$ . NEW SYNONYM.
Prosopis episcopalis, Metz, 1911. Trans. Amer. Ent. Soc., 37:131-133.
(in part).
Prosopis episcopalis var. subtristis, Metz, 1911. Op. cit.: 133. (in part).
Prosopis rudbeckiae, Metz, 1911. Op. cit.: 111-112. $ . (in part).
Hylaeus verticalis, Cockerell and Sumner, 1931. Amer, Mus. Nov., 490:3.
$ . (in part).
Hylaeus episcopalis, Cockerell and Sumner, 1931. Op. cit.'J . (in part).
In this form the female clypeus is entirely black, the lateral face marks are
frequently greatly reduced, never extending above the lower margin of the an-
tennal sockets, the pronotal collar is entirely black or with no more than one-
third of its dorsal surface yellow, and the front tibiae are entirely black or with
a very small basal spot. The male has the pronotal collar as in the female, the
lateral face marks ending at or slightly above the upper margin of the antennal
sockets and the scape entirely black or slightly suffused with reddish on the
underside. Both sexes have the wings distinctly and strongly suffused with
brownish.
The nominate form is common throughout the northern portions of the
range of the species, and in the southern areas is found in the Transition and
Boreal Zones. Much of its distribution in the Transition overlaps that of the
following subspecies and considerable intergradation occurs.
I have examined the type of H. universitatis Cockerell, which is now in the
Timberlake collection at Riverside, California. The type, a male collected at
Boulder, Colorado, by W. P. Cockerell on June 11, 1905, is not separable from
typical H. rugulosus, and I have no hesitation in reducing it into the synonymy
of that species.
Numerous specimens have been examined from the following Provinces
and States: Canada: British Columbia (Carbonate, Howser, Field, Revelstoke;
CNC, CU). United States: Oregon (Eagle Ridge, Griffin Cr., Tumalo Res.;
UCD), Idaho (Sandpoint, Orofino, Moscow Mtn. ; UI), Nevada (Ormsby Co.,
Carson City; CU, USNM, AMNH), California, Colorado (Peaceful V., Lari-
mer Co., Boulder, Steamboat Spgs.; UC, CAS, USNM, AMNH), New Mexico
(Sapella Can., Beulah; AMNH).
Hylaeus rugulosus episcopalis (Cockerell), new status.
Figures ID, 2D
Prosapis episcopalis Cockerell, 1896. Psyche, 7 (supl.) : 29-30. $ .
Prosopis episcopalis, Metz, 1911. Trans. Amer. Ent. Soc., 37: 131-133.
(in part).
Hylaeus modestus, Cockerell and Sumner, 1931. Amer. Mus. Nov., 490:6.
$ . (in part) .
Hylaeus episcopalis, Cockerell and Sumner, 1931. Op. cit.'J. (in part).
1966
Bees of the Genus Hylaeus
11
This form is more extensively maculated than the above. The female has the
lateral face marks entire, completely filling the space between the eye and cly-
peus and ending at or slightly above the upper margin of the antennal sockets,
the preapical area of the clypeus is slightly suffused with reddish, at least one-
half of the dorsal surface of the pronotal collar is maculate, and the basal one-
fourth to one-third of the front tibiae is yellow on the outer side. In the male
the lateral face marks extend about half of the distance along the eye between
the antennal socket and the upper end of the eye, and the pronotal collar is
maculate as in the female.
The major part of the range of this form lies in the Transition Zone, except
in the mountain ranges of California, Colorado and New Mexico where it be-
comes more abundant in the Upper Sonoran. Specimens have been seen from
the following states: Washington (Seattle, UC), Oregon (Warm Spr., Browns-
boro, Griffin Cr. in Jackson Co.; CAS, UCD), Idaho (Craters of the Moon,
Bear L.;UCD, UI), Utah (Mollies Nipple in Utah Co.; CIS), Wyoming (Stew-
art R. Sta.; AMNH), Colorado (Electra L., Boulder, Lake George, Cederedge,
McCoy, Bluebell Can.; AMNH, USNM, CIS, UC), New Mexico (Raton Pass;
AMNH) and California.
Hylaeus rugulosus giffardiellus Cockerell, new status.
Figures IE, 2E
Hylaeus giffardiellus Cockerell, 1925. Proc. Calif. Acad. Sci., (4), 14:
186. $.
This is the most highly precinctive form of the species, being restricted to
the Lower Sonoran Zone in the Sacramento and San Joaquin Valleys of Cali-
fornia. It differs from H. r. episcopalis, with which it intergrades in the margin-
al foothills, by having the lateral face marks extending well above the upper
margin of the antennal sockets; the apical one-third to one-half of the clypeus
strongly marked with reddish; clypeus frequently with small yellow preapical
spot; at least three-fourths of the dorsal surface of the pronotum yellow; the
stripe on the front tibiae extending one-half the length of the tibiae. In the male
the face marks are more extensive, the pronotal collar is as in the female, and
the scape is conspicuously maculate. In both sexes the maculae are bright
lemon-yellow.
Hylaeus rugulosus coquilletti (Cockerell), new status.
Figures 1C, 2C
Prosapis coquilletti Cockerell, 1896. Psyche, 7 (supl.) : 439. $ .
Prosopis episcopalis var. coquilletti, Metz, 1911. Trans. Amer. Ent. Soc.,
37:133.
Hylaeus r. coquilletti is the most highly maculate form of the species and
certainly the most distinctive. Superficially it bears a strong resemblance to the
12
Contributions in Science
No. 98
preceding but differs at once in the paler maculae which are cream-color rather
than bright lemon-yellow. The female frequently has a pale preapical spot on
the clypeus, the posterior basitarsi usually have a small basal macula (entirely
black in the other forms), and the pubescence of the third and fourth tergites
is entirely pale, that of the third no sparser on the disc than laterally. In the
male the lateral face marks extend nearly to the tops of the eyes and the ab-
dominal pubescence is pale.
This form occurs throughout the Lower and Upper Sonoran desert areas,
from Idaho south to Baja California and Sonora, and thence east to western
Texas. Although no specimens have been seen from Oregon and Washington it
is possible that this subspecies may be found in the more arid eastern portions
of these states. Specimens have been examined from the following localities:
United States: Idaho (Juliaetta, 4 mi. W., Emery Can., 12 mi. SE. of Oakley;
UI), California, Nevada (Pyramid, Pyramid L., Eastgate, 1 mi. W., Round
Mtn., Hazen, 2.5 and 3 mi. W., Sutcliffe, Dayton, Lyon Co.; CAS, CIS, CU,
AMNH, USNM, UCD, LACM), Arizona (Tucson, Castle Dome Mts., Palm
Can., San Carlos Lake, Quartzite, 10 mi. E., Whiteriver, 4 mi. N., McNary, 1 1
mi. E., Wikieup, 16 mi. N., Mammoth, Ray, 5 mi. S., Portal, 8 mi. NE., Safford,
18 mi. E., Duncan, 16 mi. W., Papago Reservation, Organ Pipe National Mon-
ument; CAS, CIS, AMNH, USNM, MCZ, UA, UC, UK, UCD, LACM), New
Mexico (Las Cruces; UK), Texas (Rio Grande City, 20 mi. SE.; AMNH).
Mexico: Sonora (Sonoyta, 50 mi. W.; UCD), Baja California (Agua Verde
Bay; CAS).
Hylaeus (Prosopis) rugulosus metzi, new subspecies
Figures 3A, B
This is the form common in cismontane southern California which has
long been known as H. episcopalis (Cockerell), a form now recognized as a
subspecies of H. rugulosus (Cockerell). The two are similar in distribution of
maculae, but differ in certain details of pubescence and sculpture.
This subspecies is named for Charles W. Metz who published a revision of
the Nearctic species of this genus in 1911, and whose work has been a constant
reference for the present study.
Female: Structurally similar to nominate and other forms. Integument black,
the following pale yellowish: paraocular areas, extending up along inner orbits
ending a little above level of upper margin of antennal sockets; stripe of vari-
able size, interrupted medially, on pronotal collar; pronotal tubercules largely;
spot on tegulae; basal spot on fore and middle tibiae; basal one-fifth to one-
fourth of hind tibiae. Pubescence of head, thorax and legs pale; of tergites I-III
pale; of tergite IV, pale, with intermixed fuscous hairs; of tergites V and VI,
largely fuscous. Erect pubescence of tergites III-VI longer and notably denser
than in other subspecies. Punctures of mesopleurae a little coarser and less con-
tiguous than in H. r. rugulosus, integument less densely tessellate than in H. r.
1966
Bees of the Genus Hylaeus
13
Figure 3. Hylaeus ( Prosopis ) spp. A, face of H. r. metzi Snelling, male; B, face of
H. r. metzi, female; C, face of H. insolitus Snelling, male; D, face of H. insolitus, fe-
male; E, H. insolitus, ventrite VIII of male; F, H. insolitus, ventrite IX of male. A-D
by Miss Dorothy Kresch, Los Angeles County Museum of Natural History.
14
Contributions in Science
No. 98
episcopalis. Punctures of tergite I sparser and finer than those of tergite II; of
tergite II separated by a puncture diameter or less; tergites III and IV with
large, poorly defined subcontiguous punctures.
Male : Structurally similar to nominate and other forms. Integument black, the
following pale yellowish to cream color: elongate spot on mandibles; clypeus;
large supraclypeal spot; paraocular areas, extending up along inner eye margin
to slightly above level of upper margin of antennal sockets; thoracic maculae as
in female; outer surface of fore tibiae, basal spot on middle tibiae, basal one-
fourth to one-third of hind tibiae, hind basitarsi. Underside of scape and flagel-
lum, fore and middle tarsi entirely, hind medio- and distitarsi all dull ferrugi-
nous to ferruginous-yellow. Pubescence similar to that of female. Thoracic and
abdominal punctation similar to that of male of H. r. coquilletti.
Holotype female and allotype male from Tanbark Flat, Los Angeles Co.,
Calif., VII-7-1963 (R. R. Snelling), on Eriogonum fasciculatum, in the Los
Angeles County Museum of Natural History. Paratypes: same locality: 19,
1 8 , same data as Holotype (LACM); 1 8 , VI-22-1950 (P. D. Hurd); 1 8,
VI-23-1950 (R. Schuster), on Eriogonum; 1 8 , VI-24-1950 (T. R. Haig) ; 1 9,
VI-25-1950 (R. Schuster; all CIS); 1 8, VI-18-1956 (R. M. Bohart); 4 9 9,
VI-21-1956 (H. R. Moffitt); 2 8 8, 1 9 , VI-22-1950 (J. C. Hall) ; 1 8 , VI-22-
1952 (E. M. Evans); 1 9, VI-22-1952 (R. L. Anderson); 1 8, VI-22-1956
(B. M. Bartosh); 1 8, VI-24-1950 (A. T. McClay); 1 8, VI-25-1956 (R. M.
Bohart); 1 8, VI-25-1956 (H. R. Moffitt) ; 1 9 , VII-14-1956 (R. M. Bohart) ;
1 9 , VII-17-1952 (A. T. McClay; all UCD) ; 1 9 , Crystal L., Los Angeles Co.,
VI- 29-1950 (F. X. Williams; CAS); 2 8 8, Los Angeles Co., VI (Coquillett;
USNM) ; 2 8 8 , San Gabriel Mts., Los Angeles Co., VI-5-1910 (F. Grinnell);
1 8 , same locality and collector, VI- 18- 19 10; 1 9 , same locality and collector,
VII- 19-1 9 10 (all USNM); 1 8, Camp Baldy, VIII-18-1929, on Eriogonum
fasciculatum; 1 9 , same locality, VIII-2 1-1929, on Mentzelia laevicaulis; 1 9 ,
same locality, VIII-22-1928 (all P. H. Timberlake; all UCR), on Adenosticia
filifolia. Ventura Co.: 3 8 8, Hungry V., 5 mi. S. Gorman, V-6-1959 (J. Pow-
ell), on Haplopappus cooperi; 1 8, same locality and date (G. I. Stage; all
CIS), on Lupinus; 1 9, Sespe Canyon, VII-10-1959 (F. D. Parker, UCD).
San Luis Obispo Co.: 1 9, Creston, 5 mi. S., VI-20-1959 (P. M. Marsh); 2
8 8, 1 9, Mill Potrero, VII-6-1959 (R. W. Spore; all UCD). Santa Barbara
Co.: 1 8 , Santa Ynez Mts., VI-24- 1959 (R. M. Bohart; UCD) . San Bernardino
Co.: 1 8, Strawberry V., San Jacinto Mts., VII-2 1-1912 (J. C. Bridwell); 2
8 8, 1 9, San Jacinto Mts., VII-29-1912 (J. C. Bridwell); 1 9, same locality
and collector, VII-7-1912 (all USNM); 1 9, Sugar Loaf, IX-21-1954 (J. C.
Hall); 1 9, Forest Home, VII-22-1953 (J. C. Hall); 1 9 , Camp Baldy, VI-29-
1956 (H. R. Moffitt); 1 9, same locality, VI-30-1956 (B. M. Bartosh); 1 8,
Camp Baldy Rd., VI-26-1956 (B. M. Bartosh; all UCD); 1 8, same locality
and date (G. I. Stage; LACM) ; 1 9 , Dollar L. trail, VII-1 1-1956 (R. C. Bech-
tel; UCD); 2 8 8, same locality, VII-10-1956 (L. A. Stange; LACM); 1 8,
1966
Bees of the Genus Hylaeus
15
L. Arrowhead, VII-9-1956 (E. G. Linsley; CIS), on Achillea millefolium; 1
8,5 9 9, Big Bear Lake, VII-7-1934 (I. McCracken), on Geranium; 1 8,
same locality and collector, VII-9-1934 (all CAS); 1 $ , same locality, VII-16-
1934 (C. D. Michener); 1 8 , same locality and collector, VIII-21-1932 (both
UK); 5 8 8 , Bear V., VIII- 19 13 (F. C. Clark; CAS) ; 1 9, Mentone, 12 mi. E.,
VII-11, 1956 (G. I. Stage); 2 9 9, Miller Can., VII-1-1956 (G. I. Stage); 2
8 8 , same data except (L. A. Stange; all LACM); 2 8 8 , Seven Oaks, VIII-
30-1949, on Eriogonum gracile; 1 8 , Mt. Home Cr., VIII-6-1949, on E. fascic-
ulatum; 1 9 , Camp Angelus, 2.5 mi. above, VIII-30-1949; 2 9 9, near Barton
Flat, VIII-30-1949, on Solidago californica; 2 9 9, Santa Ana R., VIII-30-
1949 (all P. H. Timberlake; all NCSC), on Melilotus alba; 1 9, Mt. Home,
VII- 27-19? (T. D. A. Cockerell), on Rhamnus californicus; 1 9 , same locality,
collector and host, VI-17-19?; 1 9 , U. Santa Ana R., VII-21-1946, on Erio-
gonum fasciculatum var. poliafolium; 1 9, same locality, VIII-2-1946, on
Salvia carnosa; same locality and host, VIII-4-1946; 1 8, same locality,
VIII- 18- 1946, on Gilia gilioides; 1 8,2 9 9, same locality, VIII-26-1946,
on E. fasciculatum; 1 9, same locality, VIII-29-1946, on E. nudum; 1 8,
same locality, IX- 1-1 946, on E. fasciculatum; 2 8 8, same locality, IX- 10-
1948 (all G. H. & J. L. Sperry; all UK); 1 8, Santa Ana Can., IX-7-1931
(C. D. Michener; UK); 3 8 8, 12 9 9, Mill Creek, 5700-6000', V-18 to
IX- 19, various years on Rhamnus californica, Penstemon grinnellii, Eriogo-
num fasciculatum, Chrysopsis villisa, Cordylanthus sp., Eriogonum subscapo-
sum; 1 9, Valley of the Falls, VII-28-1935, on Monardella, 1 9, Vivian Cr.
trail, VI-28- 1935, on E. subscaposum; 1 8 , trail to Dobb’s Cabin, 7000', VIII-
2-1936, on E. fasciculatum; 1 8, Forest Home, VII-5-1936, on E. fascicula-
tum; 10 9 9, Mountain Home Cr., VIII-14-1934, 8 on Phacelia ramosissima,
2 on E. elongatum; 52 8 8 , 50 9 9 , Big Bear V., VII-4 to IX- 14, various years,
on Potentilla glandulosa, Penstemon palmeri, Monardella linoides, Apocynum
sp., Geranium richardsonii, Lupinus cytisoides, Solidago confinis, Sphenosci-
adium sp., Eriogonum sp., and E. subscaposum; 12 8 8, 10 9 9, Big Pines
Camp, VII- 13 to 17-1927, on G. richardsonii, Penstemon labrosus and Eriogo-
num stenophythus; 4 9 9, Pine knob, IX-1-1936, on E. davidsonii; 1 8 , near
Crestline, V-13-1934, on P. glandulosa; 1 9, Lytle Cr., VII-4-1928 (all P. H.
Timberlake, all UCR), on E. fasciculatum. Riverside Co.: 1 8, the Gavilan,
V-17-1951 (E. I. Schlinger, UCD); 1 8, Idyllwild, VI-21-1940 (E. C. Van
Dyke; CAS); 1 9, Dutch Flat, VIII-14-1934 (C. D. Michener; UK); 1 9,
Idyllwild, VII-1936 (E. S. Ross; CAS); 3 8 8 , Herkey Cr., VI-11-1939 (E. S.
Ross); 1 8 , same locality, VI-14-1940 (C. D. Michener), on Amorpha fructi-
cosa; 1 9 , Keen Camp, 8 mi. W., V-17-1939 (E. S. Ross; all CIS), on Eriodic-
tyon\ 1 8, same locality, VI-10-1939 (J. G. Shanafelt; LACM), on Penste-
mon; 1 9, Ribbonwood, V-20-1939 (E. S. Ross; CIS), on Lupinus; 1 8, trail
above Glen Ivy, V-13-1928 (P. H. Timberlake), on E. fasciculatum; 2 8 8,
Idyllwild, VII- 12, 14-1912 (P. H. Timberlake; all UCR). San Diego Co.: 19,
Warner Spr., 7 mi. NW., VI-10-1956 (R. I. Schlinger; UCD); 1 9 , La Mesa,
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Contributions in Science
No. 98
IV-24-1953 (F. X. Williams; CAS); 6 $ $ , San Diego, VIII-24-1927 (J. C.
von Bloeker; USNM).
Paratypes are in the collections of the American Museum of Natural His-
tory, California Academy of Sciences, California Insect Survey, North Carolina
State College, United States National Museum, University of California at Da-
vis and Riverside, University of Kansas, and the Los Angeles County Museum
of Natural History.
Hylaeus ( Prosopis ) transvittatus (Cockerell)
Prosopis transvittata Cockerell, 1917. Ann. Mag. Nat. His. (8), 20:437-
438. $ 8.
This species was originally described from a small series collected by C. H.
T. Townsend at “Meadow Valley’’ Mexico. Since the original description, no
additional material has been recorded. Consequently, the new records given
below are of interest in providing clues to the distribution of this seemingly rare
species which barely enters the United States.
New Records: MEXICO: Distrito Federal: 2 $ $ , San Jeronimo, VI-11-
1946 (J. & D. Pallister; AMNH). Veracruz'. 2 $ $ , 16 mi. S. Alvarado, VI-23-
1958 (J. C. Schaeffer; UN). Durango : 1 9, Palos Colorados, 8000', VIII-5-
1947 (D. Rockefeller Exp., M. Crazier; AMNH); 1 9 , Otinapa, 8200', VIII-
1 1-1947 (D. Rockefeller Exp., C. D. Michener; AMNH). UNITED STATES:
Arizona : 2 9 9, Fly’s Peak, 8500-9700', Chiricahua Mts., Cochise Co., VIII-
5-1927 (J. A. Kusche; CAS).
Hylaeus (Prosopis) insolitus, new species
Figures 3, C-F
This new species does not seem to be closely allied to any of the Nearctic
species of the subgenus Prosopis, and differs from the other western spe-
cies in several characters. In the male the face is broader above so that the
inner orbits are more strongly convergent below; the thoracic punctures are
much coarser and better defined; the post-scutellum is distinctly punctate; ter-
gites II- VI are impunctate, transversely lineolate; ventrites VIII and IX are as
illustrated. The female may be recognized by the broad face, the vertical me-
dian clypeal stripe, the presence of the supraclypeal mark, the facial foveae
ending at the midpoint between the eyes and ocelli and the distinctly, closely
punctate post-scutellum.
Male : Integument black, the following dull lemon-yellow: mandibles, except
piceous to ferruginous apical one-fifth; labrum; clypeus, except piceous apical
margin; paraocular areas; stripe extending broadly along lower three-fourths
of inner eye margin; large supraclypeal spot (fig. 3C); narrow stripe on pro-
notal collar, broadly interrupted medially; entire pronotal tubercules; large
spot on tegulae; small apical spot on fore femora; anterior tibiae, except narrow
1966
Bees of the Genus Hylaeus
17
dark stripe on inner surface; outer, basal half of middle and hind tibiae; all
basitarsi. The following clear ferruginous-yellow: narrow stripe on underside
of antennal scape; underside of all flagellar segments; extreme apex of all basi-
tarsi; all medio- and distitarsi; tarsal claws; tibial spurs. Pubescence of head and
thorax sparse, long, whitish; of abdomen sparse, shorter, tending to form apico-
lateral fasciae on tergites I and II. Wings hyaline, slightly brownish, veins and
stigma brownish-ferruginous.
Head. UFD 0.95 x FL; LFD 0.66 x UFD; OCD 0.78 x TFD; clypeus as
wide basally as distance between latero-basal clypeal angle and inner eye mar-
gin; distance between antennal sockets slightly greater than a socket diameter,
subequal to distance between sockets and eyes; clypeus dull, densely tessellate,
punctures fine, scattered, obscure; supraclypeal area dull, densely tessellate,
with scattered punctures a little coarser than those of clypeus; lateral, maculate,
areas of face tessellate, a little shinier than clypeus, with scattered punctures
equal to those of supraclypeal area; nonmaculate areas of face and vertex ap-
pearing rugose from large, dense punctures; genal punctures smaller, well-
separated; antennal scape 0.60 times as broad as long; minimum length of first
flagellar segment less than breadth; maximum length of first flagellar less than
minimum length of pedicel or second flagellar.
Thorax. Outer margin of fore coxa straight; mesopleural punctures coarse,
subcontiguous, interstices tessellate, slightly shining; mesoscutal punctures sub-
equal to those of mesopleurae, subcontiguous, becoming slightly more sepa-
rated posteriorly; mesoscutellar punctures a little finer, more separated than
those of mesoscutum; metanotum densely tessellate with distinct subcontiguous
punctures equal to those of mesoscutellum; basal area of propodeum broad,
inner striae longitudinal, outer transverse or largely so; lateral and latero-basal
areas distinctly punctate.
Abdomen. Tergite I shining, punctures fine, mostly separated by about a
puncture diameter; tergite II duller, punctures of disc much finer than those of
tergite I; tergites III- VII, dull, transversely lineolate, impunctate; ventrites
VIII and IX as illustrated (Fig. 3, E, F) .
Measurements. Body length (front of vertex to apex of tergite II), 5.9 to
6.7 mm.; forewing length 4.8 to 5.8 mm.
Female’. Integument black, following dull lemon-yellow: Broad longitudinal
median clypeal stripe; variable supraclypeal spot; paraocular areas and along
lower three-fourths of inner eye margin (Fig. 3D) ; narrow, medially interrupt-
ed stripe on pronotal collar; pronotal lobes largely; spot on tegulae and post-
tegulae; basal spot on fore and middle tibiae; basal one-fourth to one-third of
hind tibiae. Tibial spurs and tarsal claws ferruginous vellow. Tarsal segments
rufescent. Wings and pubescence as in male.
Head. UFD 0.8 1 x FL; LFD 0.85 x UFD; OCD 0.64 x TFD; basal clypeal
width about 1.5 times distance between laterobasal angle and eye margin; di-
ameter of antennal sockets slightly less than half distance between sockets,
about two-thirds distance between sockets and eye margins; clypeus and macu-
18
Contributions in Science
No. 98
late areas of face dull, densely tessellate, with scattered obscure punctures; non-
maculate areas with punctures coarse, contiguous; genal punctures fine, sepa-
rated by about a puncture diameter; first flagellar segment longer than second;
facial foveae widely removed from eye margin above, ending at or near mid-
point between eyes and lateral ocelli.
Thorax. As described for male except that metanotal punctures are much
finer than those of mesoscutellum.
Abdomen. Tergite I shining, finely punctate, the punctures mostly sepa-
rated by about a puncture diameter; tergite II transversely lineolate, with scat-
tered minute punctures; tergite III- VI transversely lineolate, impunctate.
Measurements. Body length (front of vertex to apex of tergite II), 6.2 to
7.2 mm.; forewing length, 4.8 to 5.9 mm.
Holotype male and allotype female from Sedona, Coconino Co., Ariz., VI-
28-1951 (R. S. Beal; CIS), in the California Academy of Sciences. Paratypes:
2 8 8 , 2 $ 9 , same data as holotype (CIS) ; 1 $ , Flagstaff, 8 mi. E., Coconino
Co., Ariz., VI-23-1959 (R. R. & M. D. Snelling; LACM), on Cleome serrulata;
1 9, Walnut Canyon, Coconino Co., Ariz., V 11-29- 1 950 (T. Cohn, P. Boone,
M. Crazier; AMNH); 1 8, Ramsey Canyon, Huachuca Mts., Ariz., VII- 13-
1955 (F. G. Werner & G. D. Butler; UA), on Agave; 1 $ , Carrizo Cr., Ariz.,
VI-16-1950 (J. G. Rozen), on “cane cactus”; 1 8 , Chiricahua Mts., Ariz.,
VI-27-1934 (Fowler); 1 $ , Oak Creek Cyn., Coconino Co., 6000', VII-?
(F. H. Snow); 2 8 8, 38 mi. NE. Globe, Gila Co., Ariz., VI-16-1950 (L. D.
Beamer; all UK) ; 1 8 , 3 $ $ , The Basin, Big Bend National Park, Brewster
Co., Tex., VI-14-1948 (M. A. Crazier; AMNH); 4 9$, Arroyo Mesteno, Si-
erra del Nido, Chihuahua, VII-15-1959 (W. C. Russel; CIS); 1 9, same data
except VII-18-1959 (CIS).
Paratypes are in the collection of the California Insect Survey, the Univer-
sity of Arizona, University of Kansas, the American Museum of Natural His-
tory, and the Los Angeles County Museum of Natural History.
Literature Cited
Cockerell, T. D. A.
1937. The bees of Alberta, III, Can. Ent, 69:126-127.
Cockerell, T. D. A., and Sumner, R.
1931. Rocky Mountain Bees. III. The genus Hylaeus ( Frosopis ). Amer. Mus.
Nov., 490:1-15.
Metz, C. W.
1911. A revision of the genus Prosopis in North America. Trans. Amer. Ent.
Soc., 37:85-156.
LOS
ANGELES
COUNTY
MUSEUM
CONTRIBUTIONS
TA* IN SCIENCE
UMBER 99
May 5, 1966
THE CALIFORNIA SPECIES OF PHILORUS :
TAXONOMY, EARLY STAGES AND DESCRIPTIONS
OF TWO NEW SPECIES (DIPTERA: BLEPHAROCERIDAE )
By Charles L. Hogue
Los Angeles County Museum of Natural History • Exposition Park
Los Angeles, California 90007
CONTRIBUTIONS IN SCIENCE is a series of miscellaneous technical papers
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by the Los Angeles County Museum of Natural History. Issues are numbered sepa-
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Editor
THE CALIFORNIA SPECIES OF PH1LORUS :
TAXONOMY, EARLY STAGES AND DESCRIPTIONS
OF TWO NEW SPECIES (DIPTERA: BLEPHAROCERIDAE )
By Charles L. Hogue1
abstract: The genus Philorus is known in North America
only from California. There are four species: yosemite Osten
Sacken, 1877, jacinto n. sp., vanduzeei Alexander, 1966 and cali-
fornicus n. sp. The early stages of the first and last are known and
described here for the first time with the adults of all the species.
An unidentified larva from Modoc County is also known and de-
scribed. It possibly represents a fifth species.
The name ancilla Osten Sacken, 1878 is synonymized with
yosemite and a lectotype is designated for the species.
The description below of two new species of Philorus raises the number in
this genus to four for California, as well as for North America and the Western
Hemisphere. With a complex of mountainous terrain in western North America
equally as diverse geologically and ecologically as that of Japan, the center of
abundance of Philorus in the Old World (10 species), the number should go
considerably higher. To discover yet unknown forms, I urge intensified collect-
ing especially in the Rocky Mountains and the Pacific Northwest at large.
I wish to thank C. P. Alexander, Amherst, Massachusetts, for his assistance
and advice and for providing material [ALEX]. For the loan of specimens, I
am also indebted to the following individuals and their respective institutions:
P. H. Arnaud, California Academy of Sciences [CAS], J. L. Bath, University
of California, Riverside [UCR], P. J. Darlington, Museum of Comparative Zo-
ology, Harvard University [MCZ], J. G. Edwards, California State College,
San Jose [SJS], L. L. Pechuman, Cornell University [CU], J. A. Powell, Cali-
fornia Insect Survey, University of California, Berkeley [CIS], A. Stone,
A.R.S., U.S. Department of Agriculture— U.S. National Museum [USNM] and
P. Wygodzinsky, American Museum of Natural History [AMNH]. [LACM]
indicates material in the Los Angeles County Museum of Natural History.
Paul Freeman, British Museum (Natural History), made possible the syn-
onymy of ancilla with yosemite by locating and examining the types of the
former.
Blepharocera yosemite Osten Sacken, 1877. Bull. U.S. Geol. Survey Terr.
3:194-196. Type: LECTOTYPE, ADULT 8, BY PRESENT DESIGNA-
TION: labelled as follows: “Type 12530/ Yosemite, Cal. June 6. O Sack/
Blepharocera yosemite O.S ./ 8 genitalia on slide number CLH 63 10 14-1/Lec-
1Curator of Entomology, Los Angeles County Museum of Natural History.
Philorus yosemite
Figures 1-2, 7-13
1
SMITHS,
1WTUTI0M
2
Contributions in Science
No. 99
totype Blepharocera yosemite O.S. C.L. Hogue 1966!’ [MCZ]. Selected as single
extant specimen of original cotype series of 3 males, all with same data.
Blepharocera ancilla Osten Sacken, 1878. Cat. Des. Dip. N. Amer., Smith.
Misc. Coll. 16:266-267. NEW SYNONYMY.
Liponeura yosemite, combination of Osten Sacken, 1878. Deutsche Ento-
mologische Zeitschrift 28:408-410.
Philorus yosemite, combination of Kellogg, 1903. Proc. Calif. Acad. Sci.,
Ser. 3 3:199.
Adult Female
Size. Medium; measurements (lengths in mm from typical specimen):
overall body 9; wing 9.9; fore femur 5.9, tibia 4.5, basitarsus 2.0; mid femur
5.7, tibia 4.5, basitarsus 1.3; hind femur 6.9, tibia 6.1, basitarsus 1.9.
Head. Eyes approximate above antennal bases, interocular distance less
than ocellus diameter; bisected, upper portions slightly more extensive than
lower and with larger ommatidia. Antenna short, shape and size of flagellar
segments about equal, as figured, except basal which is slightly more than 2.0
length of others, ultimate and penultimate segments subequal, former slightly
longer than the latter. Mouthparts with all normal elements, mandibles present;
maxillary palpus with segments 2 and 3 subequal, 4 and 5 subequal.
Thorax. Scutellum with patches of bristles restricted to lateral corners;
posterior pronotum (strongly convex sclerite immediately cephalodorsad of
mesothoracic spiracle) nude. Legs with tibial spurs 0-2-2; mid tibia approxi-
mately 3.5 basitarsus. Wing shape, venation and macrotrichia as figured, R5
branching off from R4 at near or greater than a 45° angle, M3 frequently with
small barb or irregular pigment blotch projecting cephalad in basal third, 1A
sometimes not reaching wing margin; membrane entirely hyaline.
Genitalia. As figured (Fig. 13).
Adult Male
Size. Medium, smaller than female; measurements (lengths in mm from
lectotype— apparently a small individual, most specimens are somewhat larger) :
overall body 8; wing 8.4; fore femur 4.3, tibia 3.9, basitarsus 1.95; mid femur
4.6, tibia 3.9, basitarsus 1.5; hind femur 5.9, tibia 5.5, basitarsus 1.7.
Head. Eyes well separated above antennal bases, interocular distance
about equal to (or slightly less than) width of ocellar tubercle; bisected, upper
portions smaller than lower (estimated one-third the total surface area), with
slightly larger ommatidia. Antenna as in female. Mouthparts with mandibles
absent; tip of labrum bare; tip of hypopharynx bare, evenly rounded, marginal
teeth present; maxillary palpus segments 2 and 3 subequal, 5 approximately
2.0 length of 4.
Thorax. Scutellum and posterior pronotum as in female. Legs with tibial
spurs 0-1-2, single spur of middle tibia very tiny; mid tibia approximately 2.7
basitarsus. Wing as figured, shape, etc. as in female, irregularities of M3 and
1A as described for female (except 1A not reaching wing margin in several
1966
California Diptera
3
specimens from San Gabriel Canyon material); membrane hyaline apically,
basal half (or slightly more) infuscated.
Genitalia. As figured (Fig. 12). Tip of paramere attenuo-uncinate, sub-
apical one-third of shaft with dense dorsolateral area of spinules distad, heavier
teeth proximad.
Pupa
Size. Medium; measurements (lengths in mm): 5. 5-8. 6; mean (n=25):
7.0.
Structure. As figured (Figs. 8-9).
Color (in alcohol) . General dorsal surface dark grey-brown or grey-black,
lighter on abdomen, intense to near black in vicinity of respiratory processes.
Lamellae of respiratory processes translucent, grey-brown.
Larva— Final lnstar
Size. Medium to large; measurements (lengths in mm) : range: 4.5-12.3;
mean (n=25) : 7.9.
Structure. As figured (Fig. 7) ; see also Table 1 .
Color (in alcohol). Dorsal integument dark grey-brown to black; area
surrounding anterior pair of dorsal tubercles on median division III creamy-
yellow in some specimens, especially in Kern Canyon material where this color
is extensive and extends cephalad onto the neighboring segment, and caudad to
encompass the posterior pair of dorsal tubercles; head black; other sclerotic
portions brown.
Material
Colusa Co. 3 pupae: Paradise Creek, el. 2400', July 24, 1953 (H. P.
Chandler) [CAS]
Mariposa Co. 1 $ : Upper Yosemite Falls, Yosemite Valley, June 6, 1876
(C. R. Osten Sacken) [MCZ-LECTOTYPE]
2 larvae: Middle Fork Chowchilla River, bridge east of Bootjack, April
13, 1960 (R. C. Jorgensen) [SJS]
Fresno Co. 1 $ : Toll House along Dry Creek, June 10, 1963 (C. P. Alex-
ander) [ALEX]
1 $ : small stream west General Grant Grove, Route 180, el. 4900', Se-
quoia Nat’l. Park, June 7, 1963 (C. P. Alexander) [ALEX]
1 5,2 $ : waterfall at 3500', Kings Canyon Road, May 31, 1963 (C. P.
Alexander) [ALEX, LACM]
5 5 : waterfall at 3500', Horseshoe Bend, Route 180, Sequoia Nat’l. Park,
June 1, 1963 (C. P. Alexander) [ALEX, LACM]
8 pupal skins: west cliff along Route 180, Kings Canyon Nat’l. Park, June
21, 1963 (C. P. Alexander) [ALEX]
5 5,1 9 : waterfall at 3500', Big Creek near Huntington Lake, June 9,
1963 (C. P. Alexander) [ALEX, LACM]
Kern Co. 21 larvae, 13 pupae: tributary to Kern River, east side Kern
Canyon, April 6, 1963 (C. L. Hogue, Coll. No. CLH 79) [LACM]
Table 1
4
Contributions in Science
No. 99
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1966
California Diptera
5
Los Angeles Co. 22 $, 16 9 , 48 larvae, 64 pupae and pupal skins: Big
Tujunga Canyon, San Gabriel Mountains, April 22-May 27, 1962 (D. Gibo)
[LACM]
2 larvae: Ladybug Canyon, el. 4000', San Gabriel Mountains, April 24,
1963 (C. L. Hogue) [LACM]
Remarks
The type material and consequent identity of ancilla have been obscure for
a long time although it has been suspected that the name applied merely to the
female of yosemite (Walley, 1927: 1 15; Alexander, 1963, personal communica-
tion). With the cooperation of J. E. Collin, Newmarket, England, P. Freeman,
British Museum (Natural History) located two female specimens of ancilla
labeled as types by Osten Sacken. At my request he compared them against the
characters of the other California species and found them to agree with yosem-
ite. I was able to verify this during a visit to the Museum in 1964.
Notes on the biology of this species in Los Angeles County are given by
Gibo (1964).
Philorus jacinto, new species
Figures 14-15
Adult Female
Size. Medium; measurements (lengths in mm from allotype) : overall body
10; wing 1 1.5; fore femur 6.2, tibia 4.9, basitarsus 2.2; mid femur 6.2, tibia 4.9,
basitarsus 1.3; hind femur 7.9, tibia 6.9. basitarsus 2.2.
Head. Eyes approximate above antennal bases, interocular distance less
than ocellus diameter; bisected, upper portions slightly more extensive than
lower and with larger ommatidia. Antenna short, shape and size of flagellar seg-
ments about equal, except basal which is slightly more than 2.0 length of others,
ultimate and penultimate segments subequal, former slightly longer than latter.
Mouthparts with all normal elements, mandibles present; maxillary palpus seg-
ments 3-4 subequal, 4-5 subequal, 5 very slightly shorter than 4.
Thorax. Scutellum with dense patches of bristles restricted to lateral cor-
ners; posterior pronotum nude. Legs with tibial spurs 0-2-2; mid tibia approxi-
mately 3.8 length of basitarsus. Wing shape and venation as figured for yo-
semite (Fig. 1), except for barb on M3, R5 branching off from R4 at near or
greater than a 45° angle; membrane entirely hyaline.
Genitalia. As figured (Fig. 15).
Adult Male
Size. Medium, smaller than female; measurements (lengths in mm from
holotype) : overall body 7.0; wing 9.0; fore femur 5.2, tibia 4.7, basitarsus 2.3;
mid femur 5.4, tibia 4.5, basitarsus 1 .7 ; hind femur 6.9, tibia 6.3, basitarsus 2.2.
Head. Eyes well separated above antennal bases, interocular distance
slightly less than width of ocellar tubercle; bisected, upper portions smaller than
lower (estimated one-fourth the total surface area), with slightly larger omma-
6
Contributions in Science
No. 99
tidia. Antenna as in female. Mouthparts with mandibles absent; tip of labrum
bare; tip of hypopharynx bare, evenly rounded, marginal teeth present; maxil-
lary palpus segments 2-5 subequal.
Thorax. Scutellum and posterior pronotum as in female. Legs with tibial
spurs 0-2-2; mid tibia approximately 2.6 basitarsus. Wing shape etc. as in fe-
male; membrane entirely hyaline.
Genitalia. As figured (Fig. 14). Tip of paramere straight-attenuate, sub-
apical one-fourth of shaft serrately toothed dorso-laterad.
Early Stages
Unknown.
Material
Holotype $ (genitalia on slide No. CLH 650823-2) : Riverside Co. Straw-
berry Creek, el. 3000', San Jacinto Mountains, June 29, 1965 (P. A. Rauch)
[UCR, in alcohol]
Allotype $ (genitalia on slide No. CLH 650823-3): same data as holo-
type.
Paratypes 3 $ , 3 $ : same data as holotype [UCR, LACM]
Philorus vanduzeei
Figures 5-6, 16-20
Philorus vanduzeei Alexander, 1966. Bull. Brook. Ent. Soc., 58: 134-135.
Adult Female
Size. Medium; measurements (lengths in mm from allotype, unless other-
wise noted) : overall body (not determinable); wing 8.9; fore femur 6.5, tibia
4.9, basitarsus 3.0; mid leg (not allotype), femur 6.2, tibia 5.0, basitarsus 1.8;
hind femur 7.7, tibia 6.8, basitarsus 3.4.
Head. Eyes approximate above antennal bases, interocular distance less
than ocellus diameter; bisected, upper portions slightly more extensive than
lower, with larger ommatidia. Antenna short, shape and size of flagellar seg-
ments about equal, as figured, except basal which is about 2.5 length of others,
ultimate and penultimate segments subequal. Mouthparts with all normal ele-
ments, mandibles present; maxillary palpus segment 3 slightly longer than 2,
segment 5 approximately 1.7 times length of 4.
Thorax. Scutellum with patches of bristles restricted to lateral corners;
posterior pronotum nude. Legs with tibial spurs 0-2-2; basitarsi all exceptionally
long, mid tibia approximately only 2.8 length of basitarsus. Wing shape, vena-
tion and macrotrichia as figured, R4 and R5 forming a symmetrical fork from
petiole (R4+5); membrane entirely hyaline.
Genitalia. As figured (Fig. 20).
Adult Male
Size. Smaller than female; measurements (lengths in mm from holotype
1966
California Diptera
7
unless otherwise noted) : overall body (not determinable) ; wing 9.0; fore femur
6.5, tibia 6.7, basitarsus 4.5; mid leg (not holotype), femur 6.2, tibia 5.8, basi-
tarsus 3.2; hind femur 8.1, tibia 8.0, basitarsus 4.2.
Head. Eyes well separated above antennal bases, interocular distance about
equal to width of ocellar tubercle; bisected, upper portions much smaller than
lower (estimated one-fifth the total surface area), with slightly larger omma-
tidia. Antenna as in female. Mouthparts with mandibles absent; extreme tip of
labrum bare, subapical lateral margins with spiculate denticles; tip of hypo-
pharynx bare, marginal teeth present but weak; maxillary palpus segments 2-5
subequal.
Thorax. Scutellum and posterior pronotum as in female. Legs with tibial
spurs 0-0-2; basitarsi very long as in female, mid tibia only about 1.8 length of
basitarsus. Wing shape etc. as in female; membrane entirely hyaline.
Genitalia. (Known only from incomplete holotype.) As figured for disti-
style (Fig. 18) and IX tergite lobes (Fig. 19); shape of basal process of disti-
style not determinable; tegmen oblong, apex rounded; tip of paramere broadly
uncinate, distal half of shaft with sparse, slender spines (similar to yosemite but
fewer in number).
Early Stages
Unknown.
Material
San Diego Co. 1 8,1 9 : Alpine, April 9, 1915 (M. C. Van Duzee)
[ALEX-holotype and allotype]
1 8,1 9 : same data as above, from same series but unknown at the time
of original description. [CU]
Remarks
Unfortunately vanduzeei is known only from four imperfect specimens.
Collecting in the Laguna Mountains in southern California should produce ad-
ditional material. The species is known only from the type locality presently but
probably will be found in the neighboring Santa Ana Mountains and possibly
in other ranges to the north and south in Baja California, Mexico.
Philorus californicus, new species
Figures 3-4, 21-27
(?) Philorus yosemite, identification of Kellogg, 1903. Psyche 10:186-187.
(?) Bibiocephala sp. identification of Peterson, 1951. Larvae of insects,
Pt. II, p. 274, Fig. D6-H-I.
Adult Female
Size. Medium, generally smaller and less robust than the other species;
measurements (lengths in mm from typical specimen): overall body 8; wing
8
Contributions in Science
No. 99
8.9; fore femur 5.3, tibia 4.9, basitarsus 2.6; mid femur 5.2, tibia 4.7, basitarsus
1.9; hind femur 7.2, tibia 6.7, basitarsus 2.5.
Head. Eyes approximate above antennal bases, interocular distance less than
ocellus diameter; bisected, upper portions slightly more extensive than lower,
with larger ommatidia. Antenna short, shape and size of flagellar segments ap-
proximately equal, as figured, except basal which is about 2.5 length of others,
ultimate segment 1.5 length of penultimate. Mouthparts with all normal ele-
ments, mandibles present; maxillary palpus segment 3 approximately 1.4 length
of 2, segment 5 approximately 2.7 length of 4.
Thorax. Lateral corners of scutellum with dense patches of bristles, patches
thinning mesad and connecting across posterior margin; posterior pronotum
with a few bristles. Legs with tibial spurs 0-2-2; mid tibia approximately 2.5
length of basitarsus. Wing shape, venation and macrotrichia as figured, R4 and
R5 forming a symmetrical fork from petiole (R4+5); membrane entirely
hyaline.
Genitalia. As figured (Fig. 27).
Adult Male
Size. Smaller than female; measurements (lengths in mm from typical
specimen) : overall body 4.5; wing 6.7; fore femur 4.1, tibia 4.2, basitarsus 2.2;
mid femur 4.2, tibia 3.8, basitarsus 1.8; hind femur 5.5, tibia 5.2, basitarsus 2.2.
Head. Eyes well separated above antennal bases, interocular distance near-
ly one-half width of ocellar tubercle; bisected, upper portions smaller than low-
er (estimated one-fourth the total surface area) , with slightly larger ommatidia.
Antenna as in female. Mouthparts with mandibles absent; tip of labrum densely
spiculate; tip of hypopharynx with elongate spicules, attenuate, marginal teeth
present; maxillary palpus segments 2 and 3 subequal, 5 approximately 3.0
length of 4 (sometimes less).
Thorax. Scutellum and posterior pronotum as in female. Legs with tibial
spurs 0-0-2, mid tibia approximately 2. 1 basitarsus. Wing as figured, as in fe-
male; membrane entirely hyaline.
Genitalia. As figured (Fig. 26). Tip of paramere ensiform, shaft without
spines; dorsal bristles of basistyle usually numerous (as figured), sometimes
reduced to 1 or 2.
Pupa
Size . Medium, smaller than all above species; measurements (lengths in
mm) : range: 5. 0-7.0; mean (n=25) : 5.9.
Structure. As figured (Figs. 22-23).
Color (in alcohol) . General dorsal surface evenly medium brown; lamel-
lae of respiratory processes translucent, pale grey.
Larva— Final Instar
Size. Medium to large; measurements (lengths in mm): range 3.5-10.1;
mean (n=25) : 6.9.
1966
California Diptera
9
Structure . As figured ( Fig. 2 1 ) ; see also Table 1 .
Color (in alcohol). Dorsal integument light brown; head and other scle-
rotic portions dark brown.
Material
Holotype 8 : Madera Co. Mugler Meadow, July 31, 1946 [CAS]
Allotype $ : Sierra Co. Sierra City, Aug. 10-14, 1963, disturbed from large
rock by stream— Yuba River, shady, netted in flight (R. Westcott) [LACM]
Paratypes : Siskiyou Co. 1 8 (partly dissected from pupal case): 10 mi.
SW Shasta City, S. Fork Sacramento River, July 21, 1948 (W. Wirth) [USNM]
2 $ : S. Fork Sacramento River, el. 4000', Aug. 4, 1953 (H. P. Chandler)
[CAS]
Sierra Co. 5 8 (one specimen totally dissected and mounted on slides, Nos.
CLH 63 1 1 14-3a-d) : Same data as allotype. [LACM]
Madera Co. 1 8 : Same data as holotype, genitalia on slide No. CLH
631204-5 [CAS]
Mariposa Co. 1 $ : Yosemite, July 2, 1947 (A. L. Melander) [USNM]
Other. Siskiyou Co. 2 8,3 $ , 66 larvae, 60 pupae (adults dissected from
pupal skins, partly or wholly, mounted on slides) : 10 mi. SW Shasta City, July
21-22, 1948 (W. Wirth) [USNM, CIS]
Humboldt Co. 11 larvae: 6 mi. W. Dinsmore, June 18, 1950 (L. W.
Quate) [CIS]
9 larvae, 1 pupa: Willow Creek, August 12, 1948 (W. Wirth) [USNM]
Sierra Co. 102 larvae, 83 pupae: Sierra City, June 16, 1940 (T. Aitken &
M. Crazier) [AMNH]
El Dorado Co. 3 larvae: Riverton, June 27, 1950 (Ting & Quate) [CIS]
Mono Co. 18 larvae: Sardine Creek, el. 8500', June 28, 1951 (Silver)
[LACM]
Inyo Co. 39 pupae: Whitney Portals, August 31, 1951; on rocks under
waterfall (Coleman) [USNM]
Fresno Co. 2 larvae, 3 pupae: Granite Creek, Kings River Canyon, July
13, 1930 (V. L. Kellogg) [USNM]
Tulare Co. 3 larvae: Giant Forest-Marble Fork, Sequoia Nat’l. Forest,
Kings River Trail, el. 6500-7100'; July 24, 1907 (J. C. Bradley; Johannsen Lot
2275) [CU]
Remarks
It appears that Kellogg’s “re-discovery” ( 1903) of yosemite in Kings Can-
yon (Granite Creek) was actually the discovery of californicus (Hogue, foot-
note p. 44 in Gibo, 1964) , however, it is not possible to say this with certainty,
from Kellogg’s description or material. The specimens he collected are not
available although I have examined a larva and pupa which are definitely cali-
fornicus collected by Kellogg in Granite Creek, July 13, 1930. References in
his paper to the contiguous respiratory flaps— although this also applies to yo-
semite (but in strict regard to the posterior flap only, not the whole gill struc-
10
Contributions in Science
No. 99
ture as in calif or nicus') —and similarity of the lateral processes of Agathon (as
Bibiocephala ) comstocki which is more like calif or nicus than yosemite, also
lead me to suspect that he found calif ornicus. The question must remain unan-
swered until, if ever, his material is found.
The larva figured and described by Peterson (1951:274, Fig. D6-H-I) is
certainly a Philorus very closely related to P. calif ornicus. It is even likely that
it is that species.
The number of gill filaments in the mature larva of calif ornicus (3) is less
than previously cited as minimum (5) for fourth instar Philorus larvae by Ki-
takami (1941:56), Stukenberg (1958:124) and Alexander (1963:41).
The structure of the pupal gills is similar to that of Philorus alpinus, a
Japanese species (figured by Kitakami, 1931 : Fig. 52) and pupa “Q” ( Philorus
sp.) , an Indian species (figured by Tonnoir, 1930: Fig. 39) .
UNIDENTIFIED LARVA FROM MODOC COUNTY2
Figures 28-29
From two localities in the Warner Mountains of Modoc County, I have
larvae representing an unidentifiable species, there being no way of correlating
them with adults. Their structure is basically similar to that of yosemite, the
most conspicuous difference being the much greater development of the dorsal
and dorsolateral tubercles. Other differences are summarized in the diagnosis
and in Table 1.
Larva— Final Instar
Size. Medium; measurements (lengths in mm) : range 4. 2-8. 2.
Structure. As figured (Fig. 28-29) ; also see Table 1.
Color (in alcohol). Dorsal integument light brown; head and other scle-
rotic portions dark brown.
Material
Modoc Co. Cedar Pass, May 15, 1948 (W. Wirth) [USNM]
Willow Creek, Fandango Pass Road, May 15, 1948 (W. Wirth) [USNM]
DIAGNOSIS AND KEY TO SPECIES
MALES
1. General: Wing vein R5 branching off from R4 at near or greater than a 45°
angle (Fig. 2). Genitalia : Apex of dorsal dististyle lobe broadly rounded
2
General’. Wing veins R4 and R5 forming a symmetrical fork from petiole
2Material which has come to hand since this manuscript was submitted indicates
strongly, though not conclusively, that this is the larva of Agathon elegantulus von
Roder, 1890.
1966
California Diptera
11
(R4 + 5) (Figs. 4, 6). Genitalia : Apex of dorsal dististyle bifurcate or at-
tenuate 3
2. General : Basal half of wing infuscated. Genitalia : Dorsal lobe of dististyle
small, not extending beyond posterior margin of ventral lobe .... yosemite
General : Wing membrane entirely hyaline. Genitalia : Dorsal lobe of outer
dististyle large, extending well beyond posterior margin of ventral lobe ....
jacinto
3. General : Larger species, wing length 9.0 mm. Scutellum with patches of
bristles restricted to lateral corners. Fore tibia 1.5 length of basitarsus. Gen-
italia: Dorsal lobe of outer dististyle undivided, attenuate, with numerous
normal bristles concentrated at the tip vanduzeei
General : Smaller species, wing length 6.7 mm. Scutellum with short bristles
all along posterior border, connecting the patches at the lateral corners. Fore
tibia 1.9 length of basitarsus. Genitalia : Dorsal lobe of outer dististyle bifur-
cate, posterior fork tipped with two large bristles, anterior fork with two
spiniforms californicus
FEMALES
1. General : Wing vein R5 branching off from R4 at near or greater than a 45°
angle (Fig. 1) 2
General: Wing veins R4 and R5 forming a symmetrical fork from petiole
(R4 +5) (Figs. 3, 5) 3
2. General: No characters presently known. Genitalia: Oviscapt tapering cau-
dad; only two setae arising from small sclerite immediately caudad of ovi-
scapt; small lobe laterad of oviscapt devoid of setae yosemite
General: No characters presently known. Genitalia: Oviscapt roughly quad-
rangular in outline, lateral borders parallel; four setae arising from small
sclerite immediately caudad of oviscapt; setae present on small lobe laterad
of oviscapt jacinto
3. General: Scutellum with patches of short bristles restricted to lateral cor-
ners. Genitalia: Spermatheca elongate ovoid vanduzeei
General: Scutellum with short bristles all along posterior border, connecting
the patches at the lateral corners. Genitalia: Spermatheca pear-shaped ....
californicus
MATURE LARVAE
( jacinto and vanduzeei unknown)
(See also Table 1)
1 . Dorsal sclerotized process in the form of tubercles. Lateral gill filaments 5
in number 2
Dorsal sclerotized processes in the form of transverse plates. Lateral gill fila-
ments 3 in number californicus
12
Contributions in Science
No. 99
2. Lateral sclerotized processes divided into dorsal and ventral subequal, api-
cally swollen lobes yosemite
Lateral sclerotized processes undivided
UNIDENTIFIED LARVA FROM MODOC CO.
PUPAE
( jacinto and vanduzeei unknown)
1. Respiratory lamellae large, convoluted, the two median ones entire. Dorsal
integumentary granules large and dense. Median leg case of female shorter
than adjoining ones yosemite
Respiratory lamellae small, flat, the two median ones deeply incised. Dorsal
integumentary granules small, sparse. All leg cases of equal length in both
sexes calif or nicus
Literature Cited
Gibo, D. L.
1964. Notes on the biology of Blepharocera micheneri and Philorus yosemite
(Diptera: Blepharoceridae) in southern California. Bull. So. Calif.
Acad. Sci., 63:44-53.
Kellogg, V. L.
1903. The re-discovery of Philorus ( Blepharocera ) yosemite Osten-Sacken.
Psyche, 10:186-7.
Kitakami, S.
1931. The Blepharoceridae of Japan. Memoirs Coll. Science, Kyoto Imperial
University, Ser. B., 6:53-108 + 7 pi.
Peterson, A.
1951. Larvae of insects Part II. Columbus, Ohio: pub. by author. 416 pp.
Stuckenberg, B. R.
1958. Taxonomic and morphological studies on the genus Paulianina Alex-
ander (Diptera: Blepharoceridae). Memoires de L’lnstitut Scientifique
de Madagascar, Ser. E., 10:97-198.
Tonnoir, A. L.
1930. Notes on Indian blepharocerid larvae and pupae with remarks on the
morphology of blepharocerid larvae and pupae in general. Rec. Indian
Mus., 32:161-214.
Walley, G. S.
1927. Review of the Canadian species of the dipterous family Blephariceridae.
Can. Ent., 59:112-116.
1966
California Diptera
13
14
Contributions in Science
No. 99
y o s e m i t e
1966
California Diptera
15
9
y o Semite
16
Contributions in Science
No. 99
c?
yosemite
1966
California Diptera
17
j a c i n t o
18
Contributions in Science
No. 99
vanduzeei
1966
California Diptera
19
21
calif ornicus
20
Contributions in Science
No. 99
22
23
cal if ornicus
1966
Cf
9
27
cali
California Diptera
21
22
Contributions in Science
No. 99
29
LOS
ANGELES
COUNTY
MUSEUM
Dumber 100
CONTRIBUTIONS
c\7l«d IN SCIENCE
May 5, 1966
A NEW GENUS OF FISSURELLIDAE AND A NEW NAME
FOR A MISUNDERSTOOD SPECIES OF
WEST AMERICAN DIODORA
By James H. McLean
Los Angeles County Museum of Natural History • Exposition Park
Los Angeles, California 90007
A NEW GENUS OF FISSURELLIDAE AND A NEW NAME
FOR A MISUNDERSTOOD SPECIES OF
WEST AMERICAN DIODORA
By James H. McLean1
Abstract: A new genus, Scelidotoma, type species Emar-
ginula bella Gabb, is proposed for three North Pacific species, and
a new name, Diodora arnoldi, is proposed, with adequate descrip-
tion, for “Diodora murina” of authors, not of Arnold, 1903.
During the course of a revision of the Fissurellidae of the temperate prov-
inces of western North America, I have found that one new genus is needed
and also a new name for a species. Preliminary to the use of these names in a
dissertation submitted to Stanford University I am here describing them.
ACKNOWLEDGMENTS
I am grateful to Dr. A. Myra Keen and Dr. Rolf Bolin for criticism and
reading of the manuscript. Dr. Robert Robertson provided comparative ma-
terial from the collection of the Philadelphia Academy of Natural Sciences
(ANSP). Photographs were made by Perfecto Mary and Armando Solis. This
investigation was supported (in part) by a fellowship, number 18613, from the
Division of General Medical Sciences, U.S. Public Health Services.
The generic assignment of the uncommon Californian species known as
Hemitoma bella (Gabb) has not been questioned by American malacologists,
although Japanese authors have for some time been using a different genus for
two related species that occur in Japanese waters. Consideration of this discrep-
ancy has convinced me that usage of Hemitoma Swainson is no longer appro-
priate; neither is the Japanese solution satisfactory. Thus, the erection of a new
genus to contain these species seems advisable.
Scelidotoma, new genus
Type species: Emarginula bella Gabb, 1865 (equals Subemarginula yatesii
Dali, 1901). Figures 1 and 2.
Diagnosis : Shell large, white, apex posterior to center. Radial sculpture of
numerous primary and secondary ribs, intersecting the concentric growth lines
as imbrications. Juvenile shell deeply notched by an Emarginula-like slit, ma-
ture shell with arched selenizone indenting the margin. Interior with channel
corresponding to selenizone; muscle scar horseshoe-shaped, with inturned
hooked process.
Additional assignable species : Subemarginula gigas von Martens, 1881
(Kira, 1962: 6, pi. 5, fig. 14; Habe and Ito, 1965: 6, pi. 3, fig. 4); Emarginula
1Curator of Invertebrate Zoology, Los Angeles County Museum of Natural History.
2
1966
New Genus of West American Diodora
3
vadosinuata Yokoyama, 1922 (Kira, 1962: 6, pi. 5, fig. 13;Habe and Ito, 1965:
6, pi. 3, fig. 3).
Remarks : The juvenile shell of Scelidotoma bella figured here is 4.9 mm.
in length; its slit measures 0.85 mm., 17% of the shell length. Judging from the
growth marks visible on mature shells, a similar deep incision is present in juve-
nile shells of S. gigas. Scelidotoma vadosinuata is morphologically very close to
S. bella; I can detect no distinguishing features in the illustrations. According
Figure 1. Scelidotoma bella (Gabb). Holotype, Subemarginula yatesii Dali, USNM
162062, Monterey, California. Long. 51, lat. 36, alt. 13 mm.
Figure 2. Scelidotoma bella (Gabb). Juvenile, LACM, Carmel, California. Long.
4.9, lat. 2.9, alt. 1.8 mm.
Figures 3 and 4. Diodora arnoldi new species. Holotype, LACM 1143, Carmel, Cali-
fornia. Long. 17.7, lat. 10.6, alt. 6.2 mm.
SMITHSONiAri
iwimmoN
4
Contributions in Science
No. 100
to Kira ( 1962), it has been taken at 150 to 200 fathoms off Iwate Prefecture,
Japan, considerably deeper than has been recorded for S. Bella. Scelidotoma
vadosinuata may qualify as a synonym of S. Bella.
Hemitoma Swainson, 1840 (type species Patella octoradiata Gmelin,
1791), and its subgenus Montfortia Recluz, 1843 (type species Emarginula
australis Quoy and Gaimard, 1834), differ from Scelidotoma in the following
ways: ( 1 ) Species of Hemitoma are usually higher in proportion but reach only
one-fourth the length of Scelidotoma, (2) the epipodium is brightly pigmented
and the interior of the shell is colored in Hemitoma, whereas these features lack
color in Scelidotoma, (3) radial sculpture in Hemitoma includes eight to ten
strong primary ribs; the number in Scelidotoma is over eighteen, (4) the seleni-
zone in Hemitoma coincides with one of the primary ribs, which is no larger
than the two adjacent primary ribs; the primary rib forming the selenizone of
Scelidotoma is larger than all other primary ribs, (5) juvenile shells of Hemi-
toma lack the deep anterior slit of Scelidotoma, (6) the fifth lateral tooth of the
radula in Hemitoma has two small cusps adjacent to the shorter of the two
large cusps, making a total of four cusps, whereas the radula of Scelidotoma
Bella is of the basic emarginulid type, with only two cusps on the fifth lateral2,
(7) Hemitoma is a tropical group; Scelidotoma is a cool-water group, ranging
north to southern Alaska.
The genus Clypidina Gray, 1847 (type species Patella notata Linnaeus,
1758), is similar to Hemitoma, but its radula departs markedly from the emar-
ginulid type; the central and all of the lateral teeth are cusped, as figured by
Thiele (1929: 34, fig. 19).
Japanese authors (Kuroda and Habe, 1952; Habe, 1953b; Kira, 1955) as-
signed Scelidotoma gigas and S. vadosinuata to the genus Tugali Gray, 1843
(type species Emarginula parmophoidea Quoy and Gaimard, 1834). Most
species of Tugali that I have studied have fine radial and concentric sculpture
with a barely perceptible selenizone. The anterior margin is rounded, as seen
from above, and early growth stages show no evidence of the Emarginula- like
slit. However, one species examined, Tugali decussata A. Adams, 1852 (see
Habe, 1953b: 45, pi. 2, figs. 25, 29), shows coarse radial and concentric sculp-
ture with a more prominent anterior rib, but the margin is not sinuous, nor is
there evidence of the slit in early stages ( ANSP 224670) .
Kira (1962) and Habe and Ito (1965) placed the two Japanese species in
the genus Tugalina Habe, 1953a (type species Tugalina radiata Habe, 1953a).
Kira used Tugalina as subgenus of Tugali, but Habe and Ito gave it full generic
status. The type species, Tugalina radiata (see Habe, 1953a: 183; 1953b: 47,
pi. 2, fig. 2), was described from Okinawa, Ryukyu Islands, and is said to have
a thin shell with broad irregular ribs and raised fasciole. There is no anterior
2The radula of Hemitoma octoradiata was illustrated by Thiele (1929: 33, fig. 18)
and that of Hemitoma emarginata (Blainville) by Turner (1959: 338, pi. 17). Al-
though Turner showed only two cusps on the fifth lateral in her illustration of H.
octoradiata, my own mounts of both of these species show the four cusps.
1966
New Genus of West American Diodora
5
slit in the juvenile shell implied in the description and figure, and the anterior
termination of the selenizone is shown to be slightly projecting. Another species
clearly congeneric with Habe’s type species and therefore also referable to Tu-
galina is T. plana (Schepman, 1908) (see Habe, 1964: 4, pi. 2, fig. 2). Tuga-
lina plana is probably equivalent to the Tugali scutellaris of Habe ( 1953b: 46,
pi. 2, fig. 1) not A. Adams, 1852 (as figured by Reeve, Conch. Icon., vol. 17,
Tugalia, fig. 1). Examination of shells I identify as Tugalina plana (ANSP
224894) shows that there is a strong anterior ridge in mature shells (16 mm.
in length), which markedly projects beyond the anterior shell margin. There is
no evidence of the slit in juvenile shells in this lot.
The genus Tugalina is evidently the generic unit closest to Scelidotoma,
but it is unsatisfactory for the allocation of the species concerned because of
the fundamental difference in the selenizone. In Tugalina the deep slit of the
juvenile shell is lacking and in the mature shell the selenizone comes to pro-
ject beyond the margin of the shell. Tugalina may further be distinguished from
Scelidotoma in size and distribution. The largest known specimens of Tugalina
species reach only one quarter of the length of large Scelidotoma. T ugalina is a
tropical group, whereas the species of Scelidotoma occur in cold provinces of
the North Pacific.
Although the three generic units, Tugali , Tugalina, and Scelidotoma could
be ranked as subgenera of Tugali, the structural distinction in the juvenile shell
of Scelidotoma readily distinguishes it from both Tugali and Tugalina. In this
feature, and in size and distribution, Scelidotoma forms a compact group. Tu-
galina is distinguished from Tugali by its raised, projecting selenizone. I there-
fore favor recognition of the three units as full genera.
The name Scelidotoma is derived from the Greek noun skelis, rib cage, and
tome (f.), cutting or separation, with reference to the arched selenizone.
Diodora arnoldi, new species
Figures 3 and 4
“ Fissurella ( Glyphis ) murina Carpenter”— Dali, 1885: 543 [nomen nudum].
“Fissuridea murina (Carpenter) Dali”— Dali, 1892: 197 [nomen nudum].
“Fissuridea murina Carpenter”— Dali, 1894: 200 [nomen nudum].
“ Fissuridea murina (Carpenter) Dali”— Keep, 1904: 260 [not Arnold, 1903].
“ Diodora murina (Arnold, 1903)”— Burch, 1946, no. 60, p. 26 [not Arnold,
1 903]— Palmer, 1 958, p. 1 20 [not Arnold, 1 903].
Not “Fissuridea murina (Carpenter) Dali”— Arnold, 1903: 399 [a description
of Diodora aspera (Rathke)].
Not “Fissuridea murina Carpenter”— Arnold, 1907: 545, pi. 50, fig. 3 [a speci-
men of Megathura crenulata (Sowerby)].
6
Contributions in Science
No. 100
Description : Shell of medium size for the genus; anterior slope straight,
posterior and lateral slopes convex, sides nearly parallel, anterior end narrowed.
Foramen oval, one third of the distance back from the anterior margin of the
shell. Sculpture of approximately 30 narrow primary ribs, with secondary ribs
appearing at later growth stages, crossed by fine concentric ribs, producing
beading and squarish pits at intersections. Color yellow-white, often with inter-
rupted gray radial rays. Margin finely crenulate, interior glossy white, muscle
scar dull surfaced, callus truncate posteriorly. Dimensions (in mm.): long.
17.7, fat. 10.6, alt. 6.2 (holotype).
Type Locality : Three miles south of Carmel, Monterey Co., California
(San Jose Creek Beach), depth 70 feet. James H. McLean, collector, February
6, 1964.
Type Material : Holotype, Los Angeles County Museum of Natural History,
cat. no. 1143; paratypes, cat. no. 1 144. Additional paratypes to be distributed
to U.S. National Museum (USNM), Stanford University, California Academy
of Sciences (CAS), Santa Barbara Museum of Natural History, and the San
Diego Museum of Natural History.
Distribution : Crescent City, Del Norte Co., California (USNM 104121)
to San Martin Island, Baja California (CAS 24041). Dali (1921) cited Mag-
dalena Bay as the southern limit for “Diodor a murina',’ but the specimen
(USNM 212710, Santa Margarita Island) is a specimen of Diodor a inaequalis
(Sowerby), having an elongate fissure.
Remarks'. The description of Diodor a arnoldi as new, to replace the famil-
iar “D. murina',’ is necessary to solve a long standing controversy in the litera-
ture. “ Diodora murina” has been variously cited as of Carpenter or Dali, but as
pointed out by Keen in Burch (1964) and Palmer (1958), the earliest descrip-
tion of a shell under that name was given by Arnold (1903). Three earlier
citations by Dali are but notes concerning equivalence. Dali (1885) : “This is
Glyphis densiclathrata of Californian conchologists, and for a time of Car-
penter, but not of Reeve!’ Dali (1892): “This is Glyphis densiclathrata of
Californian conchologists but not of Reeve; G. saturnalis of Pilsbry not of Car-
penter, and G. densiclathrata var. murina of Carpenter!’ Dali’s remarks of
1894 are identical to those of 1892, but the reference is to specimens from
the Gulf of California and must apply to some other species.
No description accompanied the name “murina” until that of Arnold
(1903: 399), quoted in full:
Shell resembling F. aspera in general outline; apical hole
nearly central, circular; radiating ridges numerous and smooth,
except for fine incremental lines; inner margin of shell quite
evenly crenulated; color white. Dimensions: long. 46, lat. 30, alt.
16 mm. Distinguishable from F. aspera by lack of coloration,
finer and more numerous ribs, more central and more nearly
round apical hole.
1966
New Genus of West American Diodora
7
This description clearly pertains to a white-shelled specimen of Diodora
aspera (Rathke); it can not include “D. murina’' of authors because the fora-
men of the latter is never centrally located and the maximum size observed for
“£>. murina” is 22 mm., less than half the size cited by Arnold.
A shell figured by Arnold ( 1907) as “Fissuridea murina Carpenter” is evi-
dently a juvenile specimen of Megathura crenulata (Sowerby), judging from
its central fissure and low elevation. Despite the misconceptions of Arnold,
some authors of the period were correctly referring to the species in question.
Keep (1904) presented an acceptable description of “ Diodora murina
Fissuridea murina (Cpr.) Dali, the white key -hole-limpet,
( Glyphis densiclathrata) . This species has a much smaller and
more delicate shell than the last. It is oblong in shape, with curved
ends. The roundish oval hole is one-third of the shell’s length from
one extremity, and there are numerous fine ribs, checked by con-
centric ridges. The color is pure white, at least in dead specimens,
and the length is about 15 mm., a little less than the diameter of
a silver dime.
Had Arnold’s description not been published a year earlier, “ Diodora murina’
could have been cited as of Keep and the name retained.
Literature Cited
Arnold, Ralph
1903. The paleontology and stratigraphy of the marine Pliocene and Pleisto-
cene of San Pedro, California. Mem. Calif. Acad. Sci., 3:1-419 pp.,
37 pis.
1907. New and characteristic species of fossil mollusks from the oil-bearing
Tertiary formations of southern California. Proc. U.S. Natl. Mus., 32:
525-546, pis. 38-51.
Burch, J. Q. (ed.)
1944-46. Distributional list of the West American marine Mollusca from San
Diego, California, to the Polar Sea. [Extracts from the] Minutes of the
Conchological Club of So. Calif., nos. 33-63.
Dali, W. H.
1885. In C. R. Orcutt. Notes on the mollusks of the vicinity of San Diego,
California, and Todos Santos Bay, Lower California, with comments by
W. H. Dali. Proc. U.S. Natl. Mus., 8: 534-552, pi. 24.
1892. In M. B. Williamson. An annotated list of the shells of San Pedro Bay
and vicinity, with a description of two new shells by W. H. Dali. Proc.
U.S. Natl. Mus., 15:179-220, pis. 19-23.
1894. In R. E. C. Stearns. The shells of the Tres Marias and other localities
along the shores of Lower California and the Gulf of California. Proc.
U.S. Natl. Mus., 17: 139-204.
1921. Summary of the marine shell-bearing mollusks of the northwest coast
of America, from San Diego, California, to the Polar Sea, mostly con-
8
Contributions in Science
No. 100
tained in the collection of the United States National Museum, with
illustrations of hitherto unfigured species. Bull. U.S. Natl. Mus., 112:
1-127, 22 pis.
Habe, Tadashige
1953a. Tugalina radiata n. gen et n. sp. Illustrated catalogue of Japanese shells,
edited by Dr. Tokubei Kuroda, 1: 183.
1953b. Fissurellidae in Japan (2). Publ. Seto Marine Biol. Lab., 3: 33-50, pi. 2.
1964. Shells of the Western Pacific in color. Vol. II. Osaka, Japan: Hoikusha,
233 pp., 66 pis.
Habe, Tadashige, and Kiyoshi Ito
1965. Shells of the world in colour. Vol. I, The Northern Pacific. Osaka,
Japan: Hoikusha, x -f 176 pp., 56 pis. [in Japanese].
Keep, Josiah
1904. West American shells. San Francisco: Whitaker and Ray, 360 pp.
Kira, Tetsuaki
1955. Coloured illustrations of the shells of Japan. Osaka, Japan: Hoikusha,
viii + 204 pp., 67 pis. [in Japanese].
1962. Shells of the Western Pacific in color. Osaka, Japan: Hoikusha, vii +
224 pp., 72 pis. [English ed. of Kira, 1955].
Kuroda, Tokubei, and Tadashige Habe
1952. Checklist and bibliography of the Recent marine Mollusca of Japan.
Tokyo, Japan: L. W. Stach, 210 pp.
Palmer, K, V. W.
1958. Type specimens of marine Mollusca described by P. P. Carpenter from
the west coast (San Diego to British Columbia), vi + 376 pp., 35 pis.
Thiele, Johannes
1929. Handbuch der systematischen Weichtierkunde. Erster Teil, Loricata.
Gastropoda. I: Prosobranchia (Vorderkiemer). Jena: Fischer, 376 pp.,
470 figs.
Turner, R. D.
1959. The genera Hemitoma and Diodora in the Western Atlantic. Johnsonia,
3 (39): 335-344, pis. 176-179.
angeles CONTRIBUTIONS
uZll IN SCIENCE
fUMBER 101 May 5, 1966
A POSSIBLE ANCESTOR OF THE LUCAS AUK (FAMILY
MANCALLIDAE ) FROM THE TERTIARY OF
ORANGE COUNTY, CALIFORNIA
By Hildegarde Howard
J
Los Angeles County Museum of Natural History • Exposition Park
Los Angeles, California 90007
A POSSIBLE ANCESTOR OF THE LUCAS AUK (FAMILY
MANCALLIDAE ) FROM THE TERTIARY OF
ORANGE COUNTY, CALIFORNIA
By Hildegarde Howard1
Abstract: A recently discovered marine deposit, of presum-
ably late Miocene age, at Laguna Hills, California, has yielded an
avifauna of eight or more species. One of these is described as a
new genus and species of flightless “auk” family Mancallidae. The
wing bones of the new form are less modified for swimming than
are those of typical Mancalla and suggest an earlier evolutionary
stage of development.
The development of the senior citizens’ town of “Leisure World” in La-
guna Hills, near El Toro, Orange County, California, has brought to light sev-
eral fossil-bearing deposits of possibly two epochs of Tertiary age. Two sites
have yielded a number of bones of the flightless auk, Mancalla, previously re-
corded only from the Pliocene. At a nearby site, however, Mancalla, is absent
and is replaced by a related form of the same family (Mancallidae) in which
the wings were less markedly specialized as paddles. This site (Los Angeles
County Museum of Natural History locality no. 1945) has yielded a large ver-
tebrate fauna including quantities of shark teeth, together with bones of ma-
rine mammals and over a hundred identifiable bones of birds. The presence of
desmostylians among the marine mammals suggests Miocene age for this lo-
cality; Mitchell and Repenning (1963) show that all North American recov-
eries of this group of sea mammals have been from Miocene strata. Initial judg-
ment of the shark fauna, by Dr. Shelton P. Applegate of the Los Angeles Coun-
ty Museum of Natural History suggests a late Miocene age (oral communi-
cation).
The bird bones at locality 1945 are fragmentary, with no evidence of asso-
ciated elements. They are well petrified and of an even texture; the color varies,
some bones being very dark brown, others lighter gray-brown. Some are
smooth of contour as if slightly water worn. Preservation is in contrast to the
rough-textured, discolored bones of Mancalla found at the other Laguna Hills
sites. The matrix is sand and conglomerate, which, for the most part, is readily
removable from the bones. On a few specimens, a very hard, adhering matrix
suggests an original concretionary deposition.
Five wing and shoulder bone fragments and a portion of lower mandible rep-
resent the mancallid. The rest of the avifauna comprises seven or more species
allocated to five families. Detailed identifications are yet to be made, but none
of the bones is assignable to species living today. On the whole, the avifauna is
oceanic, made up of the kinds of birds usually found in open ocean or on rocky
Research Associate in Vertebrate Paleontology, Los Angeles County Museum of
Natural History.
2
1966
California Fossil Birds
3
shores, including shearwaters (family Procellariidae), boobies (family Su-
lidae), and auklets (family Alcidae). One fragment of lower mandible, bear-
ing two bony toothlike projections, is assigned to the extinct family of marine
birds, the Pseudodontornithidae, and is presumably of the genus Osteodontor-
nis, described (Howard, 1957) from the Miocene of Santa Barbara County,
California. Two bones suggest a large goose— the only species that cannot be
classed as typically oceanic.
The present study is directed to a careful analysis of the mancallid bones,
and has resulted in the description of a new genus and species. Comparisons of
the new form have been made with Los Angeles County Museum of Natural
History (LACM) specimens of Mancalla from the Pliocene of Corona del Mar
(Orange County) and San Diego, California. Specimens from the latter local-
ity were also kindly loaned by the University of California, Los Angeles
(UCLA) and the University’s Museum of Paleontology on the Berkeley cam-
pus (UCMP) . Unfortunately the type humerus of Mancalla calif or niensis from
Los Angeles (and the only specimen from that locality) is lacking the distal
condyles which characterize the specimen from Laguna Hills. Except for the
area of the ectepicondylar process of the humerus, this element, and the lower
mandible, can be compared only with specimens from San Diego. The other
elements (carpometacarpus, coracoid and scapula) can be compared with both
San Diego and Corona del Mar specimens. Several elements of the flightless
Great Auk, Pinguinis impennis, in the collection of the Los Angeles County
Museum of Natural History have also been available for comparison. The hu-
merus from Laguna Hills is selected as the type of the new form.
Description
The new genus, described below, is assignable to the family Mancallidae on
the basis of the following characters : humerus with ectepicondylar process ex-
tending high above distal condyles in straight line with shaft and closely ap-
pressed thereto, shaft straight (longitudinally) on anconal surface above distal
end, distal end diagonally compressed in lateral direction so that external con-
dyle tends to overlap internal condyle and internal and median tricipital crests
are more developed than external; external condyle blunt and knoblike at its
proximal termination, with greatest palmar projection at this point; carpometa-
carpus with long, straight process of metacarpal 1 , facet for alar digit 1 relative-
ly small, trochlear area flat proximoanteriorly, narrowing posterodistally, and
of short extent proximally beyond the process of metacarpal 1 .
Although these characters are associated with a specialization of the wing
for swimming rather than for flight, they are not found in Pinguinis , the flight-
less auk of the family Alcidae. In the latter, both humerus and carpometacarpus
closely resemble these elements in other alcids. The distal end of the humerus is
typically alcid with the greatest palmar projection of the external condyle be-
low its proximal tip, the tip itself attenuated, the shaft and distal condyles
bending palmad to an even greater degree than in Uria. The process of meta-
4
Contributions in Science
No. 101
carpal 1 of the carpometacarpus of Pinguinis is slightly lengthened, but has the
convex anterior contour of Uria, and the trochlea is broad with centrally de-
pressed surface as in all alcids.
Praemancalla, new genus
Type : Praemancalla lagunensis, new species.
Diagnosis’. See species diagnosis.
Praemancalla lagunensis, new species
Figure 1 (A, C, D, E, G)
Type : Distal end of right humerus, LACM 15288; collected by M. K.
Hammer, December, 1964.
Locality and Age : LACM locality no. 1945, Laguna Hills, l3A miles
southwest of town of El Toro, in the SW/4 of NE/4 of SW/4, Sec. 34, T 6 S,
R 8 W, Orange County, California; San Juan Capistrano quadrangle. Late
Miocene?
Paratype : Proximal half of right carpometacarpus, LACM 15287; col-
lected by W. Earl Calhoun, January, 1965, at type locality.
Diagnosis’. Humerus: ectepicondylar process separated at its base from
external condyle; compression of distal end less marked than in Mancalla, tri-
cipital grooves less deeply incised and tricipital crests less prominent, more
rounded, with internal crest less distally extended and bending slightly mediad;
on palmar surface impression of brachialis anticus muscle faintly impressed,
running diagonally from ectepicondylar process to a point slightly proximal to
attachment of anterior articular ligament; area between impression and exter-
nal condyle smooth, lacking the papilla at tip of external condyle found in
Mancalla', shaft rounded, less deep than in Mancalla, breadth 66 per cent of
depth.
Carpometacarpus: length of process of metacarpal 1, 117% of depth of
proximal end of element (measured through internal crest of trochlea and
process of metacarpal 1), with less than half of process distal to level of
proximal symphysis; internal surface of carpal area with small, blunt pisiform
process; trochlear area with narrow, deep grove between internal and external
crests posteriorly; metacarpal 2 relatively broad, with more rounded anterior
contour and more angular internal contour than in Mancalla.
Referred Material’. Fragment of upper end of carpometacarpus, LACM
15290; incomplete upper end of right coracoid, LACM 15289; articular end of
scapula, LACM 15294; articular portion of lower mandible, LACM 15428. All
are from type locality.
Description of referred material'. In carpometacarpus 15290 the small
portion of metacarpal 3 that is preserved adjacent to the proximal symphysis
curves away from metacarpal 2 to a greater extent than in Mancalla, with the
internal surface of the symphyseal area distinctly depressed. This portion of the
1966
California Fossil Birds
5
Figure 1. A, C, D, E and G, Praemancalla lagunensis : A, paratype carpometacarpus,
LACM 15287, internal view; C, D, coracoid, LACM 15289, internal and dorsal views;
E, G, type humerus, LACM 15288, palmar and external views. B, F and H, Mancalla
calif or niensis? from San Diego: B, carpometacarpus, LACM 2541, internal view;
F, H, humerus LACM 2296, palmar and external views, x IV2
element is broken away in the paratype carpometacarpus of Praemancalla’,
other portions of the referred specimen are similar to the paratype insofar as
they are preserved. The coracoid resembles that of Mancalla in having a
glenoid facet that is convex and a coracohumeral surface that is slightly twisted
anteriorly at its upper end, but both areas are heavier; the scapular facet is
larger and more cupshaped; the triosseal canal faces internally, contrasting with
the posterointernal direction of the canal in Mancalla’, the furcular facet is
6
Contributions in Science
No. 101
broken, but is seemingly flatter proximally and more extended internally than
in Mancalla. The scapula is poorly preserved, but is basically similar in contour
to that of Mancalla, with relatively wide glenoid facet, and small, papillalike
coracoidal facet continuous with it; the blade is thick, with the ventral surface
tending to be concave, but less markedly so than in Mancalla. The articular
portion of the lower mandible is massive, as in Mancalla, with a broad, straight-
sided internal articular process projecting more posteriorly than in the Alcidae,
but slightly more laterally than in Mancalla', ventrally, the ramus is less sharply
angled th&n in Mancalla, and posteriorly the surface is less deeply depressed.
Measurements'. See tables 1 and 2 for humerus and carpometacarpus com-
pared with these elements of Mancalla and Pinguinis. Coracoid: distance from
head to below scapular facet, 18.3 mm.; least breadth of coracohumeral sur-
face, 4.7 mm.; breadth across triosseal canal, 5.9 mm. Lower mandible: pos-
terior breadth from external articular facet to internal articular process, 10.5
mm.; distance from anteriormost edge of internal lip of articular area to poste-
riormost tip of internal articular process, 11.5 mm.; greatest depth of ramus
externally, 8.4 mm. Scapula: approximate breadth across proximal end (tip of
acromion broken), 13.0 mm.; dimensions of blade, 2.6 x 6.2 mm.
Remarks'. In general size the available elements of Praemancalla lagunen-
sis are comparable to specimens of Mancalla from the Pliocene of Corona del
Mar and to the largest of those from San Diego, which, in turn, seemingly rep-
resent birds of the approximate dimensions of Mancalla calif or niensis and
have been, at least tentatively, referred to that species. A reevaluation study of
Pliocene Mancalla, now under way, based on material available since the 1949
report of Miller and Howard, and including collections from new localities as
TABLE 1
Measurements and Proportions of Humerus
Praemancalla lagunensis compared with Mancalla californiensis
and Pinguinis impennis
(Measurements in millimeters, proportions in per cent)
Praemancalla Mancalla 1 Pinguinis
Breadth of distal end
Depth of distal end, internally
Distance from top of ectepicondylar
process to distal extremity, externally
Breadth of shaft immediately above
ectepicondylar process
Depth of shaft immediately above
ectepicondylar process
Ratio of breadth to depth of distal end
Ratio or breadth to depth of shaft
Type
(9 specimens)
(2 specimens)
7.8
6.5- 8.0
10.3 - 10.8
10.5
9.6- 11.2
12.1-12.4
15.3
15.5-19.8
16.2-16.3
5.1
4.1 - 5.2
5.8- 6.1
7.7
8.2- 10.5
10.4- 11.0
74.3
62.5 - 72.6
83.1 - 88.5
66.0
47.7-51.1
52.7 -58.5
^Ten largest specimens from San Diego: LACM 2304, 2480, 2481 (2 bones), 2670,
2695, 6180; UCMP 45856 (2 bones); specific assignment tentative.
TABLE 2
Measurements and Proportions of Carpometacarpus
Praemancalla lagunensis compared with Mancalla californiensis
and Pinguinis impennis
(Measurements in millimeters, proportions in per cent)
1966
California Fossil Birds
7
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8
Contributions in Science
No. 101
well as from sites previously recorded, suggests that at least three species of Pli-
ocene Mancalla should now be recognized. The diagnostic features of the La-
guna Hills Praemancalla lagunensis markedly separate it from all of the Plio-
cene populations, and justify its assignment to a distinct genus. Although the
wing bones of Praemancalla show distinct specialization toward a paddle-like
wing, they are less advanced in this specialization than those of Mancalla. The
carpometacarpus in the latter genus lacks the pisiform process, has an even
longer process of metacarpal 1 than in Praemancalla, with 50% or more ex-
tending distad to the proximal symphysis, and the entire element is more com-
pressed, with the posterior division between the external and internal crests of
the trochlea less distinctly grooved. The humerus of Mancalla is also more com-
pressed, the shaft deep and bladelike, the tricipital crests and grooves more
sharply and deeply developed (respectively). The degree of development of
the wing of Praemancalla suggests a stage of evolution that might well be
looked for in the ancestors of the Pliocene Mancalla. It seems not unreasonable
to look upon Praemancalla as exactly what its name implies, a form anticipat-
ing Mancalla. These evolutionary implications of the new genus, therefore,
afford contributing evidence for a late Miocene age for Laguna Hills locality
LACMno. 1945.
Acknowledgments
The material from Laguna Hills has been acquired largely through the in-
terest and generosity of persons in that area who have kept the museum in-
formed as excavations opened up, and have themselves made collections for the
museum. I should like, in particular, to acknowledge the aid of Mr. W. Earl
Calhoun, of Fullerton, California.
Specimens illustrated were photographed by George Brauer, and expertly
retouched by Pearl Hanback to bring out detail which in these darkened speci-
mens is very difficult to show photographically. My continuing thanks go to
the John Simon Guggenheim Memorial Foundation. This study is an outgrowth
of a project sponsored by the Foundation under a research grant of 1962-63 for
investigation of avian fossils of western United States. The illustrations were
financed from this grant.
Literature Cited
Howard, Hildegarde
1957. A gigantic “toothed” marine bird from the Miocene of California. Santa
Barbara Mus. Nat. Hist. Bull. Dept. Geol., 1 : 1-23.
Miller, Loye, and Hildegarde Howard
1949. The flightless Pliocene bird, Mancalla. Carnegie Institution Washington
Publ. 584:210-228.
Mitchell, Edw. D., Jr., and Charles A. Repenning
1963. The chronologic and geographic range of desmostylians. Los Angeles
County Mus., Cont. in Sci., 78:1-20.
LOS
ANGELES
COUNTY
MUSEUM
CONTRIBUTIONS
Skh'r IN SCIENCE
UMBER 102
May 5, 1966
A NEW SYRRHOPHUS FROM MEXICO
(AMPHIBIA: LEPTODACT YLIDAE )
By James R. Dixon and Robert G. Webb
Los Angeles County Museum of Natural History
Los Angeles, California 90007
Exposition Park
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A NEW SYRRHOPHUS FROM MEXICO
(AMPHIBIA: LEPTOD ACTYLID AE )
By James R. Dixon1 and Robert G. Webb2
Abstract: The leptodactylid frog, Syrrhophus nivicolimae,
is described and is apparently confined to the lower tropical mon-
tane forest of the Nevado de Colima, Jalisco, Mexico. Remarks on
eggs, habitat, call, and meristic variation are included.
The discovery of an undescribed Syrrhophus from the Nevado de Colima,
Jalisco, Mexico, provided the impetus for a study of the herpetofauna of this
area. Specimens were obtained and observations made, in the three visits to
Nevado de Colima (Webb and Michigan State University field party on July
24, 1963; Webb and Dixon on June 1 1, 1964; and R. W. Ax tell, M. P. McKel-
vey and Webb on July 20, 1964).
The Nevado de Colima is a conspicuous topographic feature with a maxi-
mal elevation of approximately 14,200 feet. Four major habitats are recogniz-
able: (1) tropical deciduous and thorn scrub forests at the lowest elevations;
(2) lower tropical montane forest between elevations of 5,800 and 7,800 feet;
(3) tropical montane forests to approximately 13,800 feet; (4) Alpine-like as-
pect at the highest elevation. Gadow (1908: 507-516) and Goldman (1951:
180-181) published general observations about the Nevado de Colima.
Syrrhophus nivicolimae, new species
Figure 1
Holotype : LACM 3200, adult male from Nevado de Colima, six miles (air-
line) west of Atenquique, Jalisco, 7,800 feet; collected by Robert G. Webb on
July 20, 1964.
Paratypes : LACM 3201-09, taken at the type locality on the same date as
the type; LACM 3210-14, five miles west of Atenquique, 6,500 feet, collected
by Robert G. Webb on July 24, 1963.
Diagnosis : Syrrhophus nivicolimae is a member of the western Mexico
species complex that has an outer palmar tubercle smaller (usually much small-
er) than the first supernumery tubercle of the fourth finger, just distal to the
outer palmar. It may be distinguished from all western species of Mexican Syr-
rhophus by the mid-dorsal brown band extending from the rear of head to
anus; a short, blunt, rounded snout; combination of an interorbital light bar,
small snout-vent size (M= 20.4 mm.) and large tympanum (M= 52.5% of
diameter of eye).
‘Curator of Herpetology, Los Angeles County Museum of Natural History (LACM ) .
-Department of Biological Sciences, Texas Western College, El Paso.
SMfTHSQIttM Mm i <3
i
2
Contributions in Science
No. 102
Figure 7. Topotype of Syrrhophus nivicolimae, new species.
Description of Holotype : Adult male, snout short, somewhat rounded;
loreal region slightly concave and almost vertical in outline; head relatively
wide with small eyes; tibiotarsal articulation reaching to anterior edge of eye
when leg is brought forward along side of body; inner metatarsal tubercle three
to five times larger than outer one; skin of dorsum and limbs somewhat warty,
warts are relatively small but visible to the naked eye; small tubercle on outer
edge of wrist; tongue ovoid, as broad as long; choanae oval, hidden by maxillary
when viewed from ventral aspect; tibia length 48.4% of snout-vent length; foot
length 46.9% of snout-vent length; head length 36.1% of snout-vent length;
head width 38.1% of snout-vent length; eyelid width 48.4% of interorbital dis-
tance; tympanum length 17.1% of head length; tympanum length 52.5% of
1966
New leptodactylid frog
3
diameter of eye; length of digits, shortest to longest respectively, hand 1-2-4-3;
foot 1-2-5-3-4; tips of outer two fingers expanded, truncate, twice the width of
the narrowest part of digit.
Measurements in millimeters : Snout-vent length 19.4; foot length 9.1;
tibia length 9.4; total leg length 29.0; forearm length 4.5; head length 7.0; head
width 7.2; eye to nostril distance 2.0; greatest diameter of eye 2.3, of tympanum
1.2; internarial distance 2.0; interorbital distance 2.9.
Coloration in life : Ground color yellow-orange with a broad dark brown
mid-dorsal band from rear of head to anus; lateral margins of mid-dorsal band
well defined, in contrast to ground color; a dirty white interorbital bar bor-
dered posteriorly by a dark brown crossband; loreal region with a dark brown
(almost black) line from nostril through eye to arm insertion; tibiotarsal por-
tion of leg with dark brown crossbands, one and one half times as narrow as the
ground color interspaces; dorsal surface of the femur area without banding or
spotting; a small black spot covering anus; forearm with dark brown cross-
bands; upper arm orange-red, without dark markings on dorsal surface; poster-
olateral surface of forearm with one large dark brown spot; belly translucent,
dirty white; ventrolateral parts of belly lightly mottled with dark brown; pos-
teroventral surfaces of foot dark brown, almost black, from tibiotarsal articu-
lation to tip of longest toe.
Variation : The ground color varies from a gray through buff, pale yellow,
orange-red to brown. In most cases the neck and arms are darker than the body
and legs. The barring found on the tibia is often lacking on the femur. The in-
terorbital light bar is present in all but one specimen, which has the dark cross-
band behind the light bar, and the light bar broken up into a series of small
white spots. One specimen has a thin whitish line from snout to anus and from
heel to heel, across the anus. The intensity of mottling along the ventrolat-
eral margin of the belly varies from sparse to dense.
The average snout- vent length for males is 20.4 mm. ( R= 19.0 to 21.5).
The only known female has a snout-vent length of 23.5 mm. The variation in
ratios of the following measurements are expressed in per cent: tibia length/
snout length, M =44.9 (R=40.5 to 48.7); foot length/ snout-vent length, M=
42.6 (R= 40.0 to 47.1); head length/ snout- vent length, M= 34.6 (R= 32.6 to
36.8); tympanum length/head length, M= 16.9 (R= 15.4 to 18.8); eye-lid
width/ interorbital distance, M= 53.5 (R= 51.7 to 57.1); tympanum length/
diameter of eye, M— 52.5 (Rz= 50.0 to 59.0) .
Habitat: Syrrhophus nivicolimae is apparently confined to the lower tropi-
cal montane forest on the Nevado de Colima. Principal trees of this forest are
oaks and pines, including the characteristic drooping-needle pine, locally called
“pino triste!’ Poison ivy, bracken fern (Pteridium), pokeweed (Phytolacca), the
herb Eupatorium and the shrub Baccharis are prominent species in the under-
story. The understory is often a near-impenetrable thicket.
The tropical lower montane forest is continuous from the Nevado de
Colima to its sister peak to the south, the Volcan de Colima on the Jalisco-
4
Contributions in Science
No. 102
Colima state line. Suitable habitat on the two volcanic mountains seems to be
isolated, except perhaps to the south in Colima, by lowlands of thorn scrub.
Individuals of S. nivicolimae were not observed during the dry season
(June 11), but were found calling on the ground and in shrubs to a height of
five feet during the rainy season (July 20, 24). Their call is a single, rather
high peep. Some individuals were found calling from fully exposed positions,
but most made some attempt at concealment. One clutch of 12 eggs was laid
by a captive female. The smallest and largest eggs of the clutch, respectively,
were 5.3 and 5.7 mm. in diameter.
A species of Tomodactylus closely related to T. angustidigitorum (cur-
rently under study by us) is also confined to the tropical lower montane forest.
Both Syrrhophus and Tomodactylus are sympatric at about 7,800 feet. At
higher elevations only individuals of Tomodactylus are found, whereas at
lower elevations Syrrhophus occurs to the exclusion of Tomodactylus. Other
amphibians and reptiles obtained in our visits to the tropical lower montane
forest of Nevado de Colima are Eleutherodactylus occidentalis, A noils nebulo-
sus, Sceloporus grammicus, Sceloporus sp. (currently being studied by Webb)
Eumeces dugesi, and Rhadinaea taeniata.
Relationships: Syrrhophus nivicolimae does not appear to be closely re-
lated to any of the western or eastern Mexican Syrrhophus. It is closest to
S. modestus in snout-vent length and eyelid/ interorbital distance ratio; to S.
leprus and S. rubrimaculatus in foot/ snout- vent length ratio; to S. inter orbit alls
in the presence of an interorbital light bar and head/ snout-vent length ratio; to
S. pipilans in tympanum/ head length ratio and tympanum/ eye diameter ratio.
Duellman ( 1958) gives an account of the relationships of the above species and
provides a table of meristic values for the species.
Acknowledgments
William E. Duellman, Museum of Natural History, University of Kansas;
Kenneth S. Norris, University of California, Los Angeles; Jay M. Savage, Uni-
versity.of Southern California; Hobart M. Smith, University of Illinois Museum
of Natural History; and Donald W. Tinkle, University of Michigan Museum of
Zoology graciously permitted us to examine comparative material in their care.
The photograph of a topotype (Figure 1 ) was taken and donated to us by
Ralph W. Axtell. We are grateful to Rollin H. Baker for permitting Webb to
participate as a member of the field party from Michigan State University and
to the Michigan State Development fund for financial assistance in 1963, and
to the American Philosophical Society (Penrose Fund) for providing financial
assistance to Webb in 1964.
1966
New leptodactylid frog
5
Literature Cited
Duellman, W. E.
1958. A review of the frogs of the genus Syrrhophus in western Mexico. Occ.
Paps. Mus. Zool. Univ. Michigan (594): 1-15.
Gadow, H.
1908. Through southern Mexico, being an account of the travels of a natural-
ist. London: Witherby and Co., xvi+527 p.
Goldman, E. A.
1951. Biological investigations in Mexico. Smithsonian Misc. Coll., 115 (4017):
1-476.
LOS
ANGELES
COUNTY
MUSEUM
CONTRIBUTIONS
fllld IN SCIENCE
Ijmber 103 May 5, 1966
||” " ‘
A NEW SPECIES OF BOETICA
FROM THE PLIOCENE OF CALIFORNIA
By George P. Kanakoff
!
Los Angeles County Museum of Natural History • Exposition Park
Los Angeles, California 90007
A NEW SPECIES OF BOETICA
FROM THE PLIOCENE OF CALIFORNIA
By George P. Kanakoff1
Abstract: Boetica hertleini, a new species of mollusk ap-
parently belonging to the family Lacunidae, is described from
Pliocene deposits of Los Angeles County, California
On January 9, 1954, Mr. Stanford Lane brought several fossil shells im-
bedded in Miocene shale to the Los Angeles County Museum of Natural
History for determination. This material was collected in the vicinity of Sand
Canyon Road in Los Angeles County.
On January 30, 1954, the author, with a group of Invertebrate Paleontol-
ogy students, visited the Miocene localities reported by Mr. Lane and while
investigating the outcrops discovered a small triangular fill of Pliocene age.
The excavations in this solidly packed black silt (Pico Formation) yielded
an interesting marine fauna with several new species (Kanakoff, 1954, 1956).
A small lot of five specimens represented a new species of the genus Boetica
Dali. As this genus was described in 1918 and was based on a unique specimen
not since mentioned in the literature, Dali’s (1918: 137) original description
is repeated here :
‘ Boetica new genus, is proposed for B. vaginata Dali, a small shell
resembling conchologically a very solid Lacuna but with a sulcus at the
posterior commissure of the aperture and one like that of Trichotropis
anteriorly; the surface smooth, but the operculum unknown. Habitat:
San Diego, California, in 199 fathoms!’
Dali later (1919: 349) gave a more complete description of Boetica
vaginata :
“Shell small, solid, conical, white, smooth except for faint incre-
mental lines, of about five rapidly enlarging whorls including a minute
subglobular smooth nucleus; suture distinct, not deep; base rounded, ap-
erture subovate, a distinct sharp groove in the subsutural callus, the outer
lip simple, thick; the body with a thick coat of enamel curving into the
concavely arcuate pillar lip; umbilicus perforate, the area bounded by a
thickened spirally striated ridge parallel with the pillar lip, with the area
between them excavated; at the anterior end of the pillar is a shallow,
narrow sulcus, somewhat as in Trichotropis; height of shell, 4 mm.; of
last whorl, 3 mm.; diameter, 2.5 mm. U.S. Nat. Mus. Cat. no. 209891a.
“Type locality.— U.S. Fish Commission Station 4322, off La Jolla,
San Diego County, California, in 199 fathoms, shelly mud.
Curator of Invertebrate Paleontology, Los Angeles County Museum of Natural
History.
2
1966
New fossil mollusk
3
“The exact position of this genus awaits the discovery of the oper-
culum and soft parts!’
One additional dead specimen of B. vaginata is represented in the United
States National Museum according to Mr. James H. McLean (USNM 211308;
USFC station 2902, 53 fathoms off Santa Rosa Island, California).
And, finally, Mr. McLean collected one dead specimen of this species off
Monterey, California in 45 fathoms. It is through the courtesy of Mr. McLean
that Dali’s holotype (Fig. 3) and the specimen from Monterey (Fig. 2) are
figured here.
The extreme rarity of this genus, the fact that the known specimens are
dead, and finally the finding of a small lot of them in the Pico formation, opens
the way for speculation that the genus Boetica may now be extinct.
As Dali pointed out, the exact position of the genus Boetica is not certain,
as long as the soft parts remain unknown. Following Dali, the genus is tenta-
tively retained in the family Lacunidae.
Specimens of Boetica from the Pliocene of Pico formation closely resem-
ble specimens of B. vaginata, but differ considerably in proportion, and in the
shape of the wide umbilical plate. The species is therefore being put on record
as:
Boetica hertleini, new species
Figure 1
Diagnosis : Shell minute, porcellaneous, globose-conic, of five whorls;
sutures distinct, base rounded, outer lip simple, flaring, thickened; inner lip
adnate, arcuate, with a turn at the commissure, forming a tooth-like protru-
sion; umbilicus deeply perforate, bound by the thick spiral ridge, forming a
Figure 1. Boetica hertleini, new species, holotype (LACM).
Figure 2. Boetica vaginata, dead specimen collected in 45 fathoms off Monterey,
California (LACM).
Figure 3. Boetica vaginata Dali, holotype (USNM).
4
Contributions in Science
No. 103
platform narrow at the umbilicus and rapidly widening until it reaches the
inner lip; nucleus smooth, subglobular; the semilunar wide flat area formed
by the inner lip and the fasciole, under stronger magnification (X 60) appears
uniformly tuberculated, and the body whorl shows the fine spiral incremental
lines below the suture and at the base.
Dimensions’. The holotype measures 5.1 mm. in altitude, 3.5 mm. in
width; the body whorl includes 3A of the shell height.
Type Material: Holotype LACMIP No. 1145, 4 paratypes, LACMIP
No. 1146.
Type Locality : LACMIP No. 291 (Los Angeles County Museum, Inver-
tebrate Paleontology locality) : An exposed stratum of black silt, weathering
into gray, in a gully in the center of the south half of Sec. 27, T. 4 N, R. 15 W,
Mt. San Bernardino B. and M. (which is probably the same as Kew’s (1924)
locality No. 3390.5) ; it is exactly one half mile south of the Humphreys Rail-
road Station, Los Angeles County, California.
Age and Formation: Upper Pliocene, Pico formation.
This shell is named for Dr. Leo George Hertlein, Curator of Paleontology
of the California Academy of Sciences, a prominent scholar, on whose knowl-
edge, time and kindness the author has imposed for many years.
Literature Cited
Dali, W. H.
1918. Changes in and additions to molluscan nomenclature. Proc. Biol. Soc.
Wash., 31: 137.
1919. Description of new species of Mollusca from the north Pacific Ocean
in the collection of the United States National Museum. Proc. U.S.
Natl. Mus., 56: 293-371.
Kanakoff, G. P.
1954. A new Kelletia from the Pliocene of California. Bull. So. Calif. Acad.
Sci., 53 (2): 114-117.
1956. Two new species of Nassarius from the Pliocene of Los Angeles County,
California. Bull. So. Calif. Acad. Sci., 55 (2) : 110-114.
Kew, W. S. W.
1924. Geology and oil resources of a part of Los Angeles and Ventura Coun-
ties, California. U.S. Geol. Surv. Bull., 753: 77-81.
LOS
ANGELES
COUNTY
MUSEUM
Dumber 104 July 22, 1966
CONTRIBUTIONS
Sliv IN SCIENCE
OBSERVATIONS ON THE DISTRIBUTION, COLORATION,
BEHAVIOR AND AUDIBLE SOUND PRODUCTION OF THE
SPOTTED DOLPHIN, ST EN ELLA PL AGIO DON (COPE)
By
David K. Caldwell and Melba C. Caldwell
Los Angeles County Museum of Natural History • Exposition Park
Los Angeles, California 90007
CONTRIBUTIONS IN SCIENCE is a series of miscellaneous technical papers
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Editor
OBSERVATIONS ON THE DISTRIBUTION, COLORATION,
BEHAVIOR AND AUDIBLE SOUND PRODUCTION OF THE
SPOTTED DOLPHIN, ST EN ELLA PL AGIO DON (COPE)
By David K. Caldwell1 2 and Melba C. Caldwell-
Abstract: The spotted dolphin ranges from off Cape May,
New Jersey, to Panama. Present data indicate that it is gen-
erally restricted to a spatial zone from about five to 200 miles
offshore of mainland North and Central American shores, al-
though in the northeastern Gulf of Mexico it comes into shallow
waters much closer inshore in late spring. The young are un-
spotted. Several forms of breathing behavior in captivity were
observed which seem related to specific life needs in the wild.
Audible sounds of several types were recorded from captive
animals. A possible eastern Atlantic record is noted.
Introduction
Although apparently it is a common species offshore, relatively little has
been published on the biology of the spotted dolphin, Stenella plagiodon
(Cope). Consequently, it seems appropriate to add additional information on
the natural history of this species, particularly as observed in the northeastern
Gulf of Mexico, and to describe certain captive behavior as it seems to corre-
late with known behavior by this species in the wild. In addition, we have had
the opportunity to study the coloration of a number of individuals of different
sizes, and to make tape recordings of two captive animals.
Acknowledgments
J. B. Siebenaler of Florida’s Gulfarium at Fort Walton Beach generously
allowed us free access to two captive spotted dolphins in his charge and was
also most helpful in obtaining specimens and data on wild animals from that
region. Leonard P. Hutchinson of the Gulfarium also helped gather informa-
tion on wild spotted dolphins and provided several of the photographs used
in this paper. B. C. (Cliff) Townsend of Marineland of Florida at St. Augustine
was especially helpful in providing data on wild and captive animals and
photographs of captives. Others who helped in various ways by providing
data, photographs and/or other valued help and suggestions were: F. C.
Fraser of the British Museum (Natural History); Winfield Brady of the
1Curator of Ichthyology, Los Angeles County Museum of Natural History; also
Research Associate, Florida State Museum, and Collaborator in Ichthyology, Insti-
tute of Jamaica.
2Research Associate, Los Angeles County Museum of Natural History; Staff Re-
search Associate, Allan Hancock Foundation, University of Southern California.
1
2
Contributions in Science
No. 104
Aquarium of Niagara Falls, New York; Richard W. Edgerton of Marineland
of Florida; John H. Prescott of Marineland of the Pacific, Los Angeles; Ray-
mond M. Gilmore of California Western University, San Diego; Dewey
Destin, Marjorie Siebenaler, Gregory Siebenaler, Lowell Longaker, Ronald
Ward and Chuck Ray of Destin (Florida), Fort Walton Beach and the Gulf-
arium; and Robert L. Brownell, Jr., and Armando Solis of the Los Angeles
County Museum of Natural History. Partial support for certain phases of the
study was received from the Los Angeles County Museum Associates and
through grants from the National Institute of Mental Health (MH-07509-01 )
and the National Science Foundation (GB-1 189).
Geographical Distribution
Stenella plagiodon was originally described by Cope (1866: 296) as Del-
phinus plagiodon from a skull of unknown geographical origin in the collec-
tions of the United States National Museum (No. 3884). True (1885: 323)
compared the skull with one from a specimen of the typical Gulf of Mexico
spotted dolphin (from off Pensacola, Florida) and found that they did not
differ in any significant way. Later (1889: 66), and with the same results, he
also compared the type of plagiodon with the skull from an animal, in external
appearance apparently of the same spotted species as the Gulf specimen, col-
lected off Cape Hatteras, North Carolina. In his 1885 paper he believed that
plagiodon was a junior synonym of Prodelphinus {—Stenella) doris (Gray),
but he changed this opinion in his later paper and concluded with some cer-
tainty that doris and plagiodon are distinct and that the specific name for the
common spotted dolphin of the Gulf of Mexico and south Atlantic coast of
the United States (the species under discussion and illustrated in the present
report) should stand as plagiodon. Despite the lack of a definite type locality
for this species, present evidence suggests that it usually occurs in offshore
waters (but within somewhat restricted spatial limits) along the mainland
Atlantic coasts of North and Central America from Cape May, New Jersey,
to Panama. Consequently, the type locality for plagiodon almost surely lies
within that spatial and geographical range. However, Dr. F. C. Fraser is
studying the entire genus Stenella at this writing and any further restriction of
a possible type locality for S. plagiodon should await his judgment. There ap-
parently is no firm basis for published statements that the type locality is the
eastern coast of the United States ( e.g ., Elliot, 1901: 31, 1904: 58).
The range of this species as presently understood apparently is restricted
to the western North Atlantic and extends from 60 miles off Cape May, New
Jersey (Schevill and Watkins, 1962: 9), southward to Golfo de San Bias,
Panama (Miller and Kellogg, 1955: 657). We find no published records or
specimens in collections to form the basis for the note of Hall and Kelson
(1959: 818) that this species occurs in South America, although the record
from nearby Panama in Central America may account for their statement.
1966
Spotted Dolphin Biology
3
General statements on geographical range for this species which are less
extensive (usually indicating occurrence from Cape Hatteras, North Carolina,
to the Gulf of Mexico) but which are included in the New Jersey to Panama
limits, have been given by many writers. Included among these are: True
(1889: 66, 165), Elliot (1901. 31; 1904: 58), Lyon and Osgood (1909: 9),
Miller (1924: 508), Fraser (1937: 332), Gunter (1941; 1954: 548), Poole
and Schantz (1942: 113), Lowery (1943: 256), Burt and Grossenheider
(1952: 170), Palmer (1954: 332), Hall and Kelson (1959: 818) and Cald-
well and Golley ( 1965 : 25 ) .
It is interesting to note that none of the records that we have seen make
reference to the occurrence of this species among the islands of the West
Indies, although it is primarily an inhabitant of offshore waters and would
seem to be expected there. Elliott (1904) did not include it from the West
Indies, and in our studies of West Indian marine mammals made during the
past several years we have also failed to find specimens or other reports of
S. plagiodon in that region.
The possible occurrence of S. plagiodon in the eastern Atlantic has been
suggested by Cadenat (1959) and Cadenat and Lassarat (1959). In late
February, 1958, a rather robust male spotted dolphin was harpooned ten
miles off the entrance to the port of Abidjan, Ivory Coast, Africa. This speci-
men, 188 cm. (about 74 inches) in snout to caudal-notch length, was photo-
graphically illustrated in several views by Cadenat (1959: figs. 34-37) and
Cadenat and Lassarat (1959: figs. 1-4) and was questionably identified by
those writers as S. plagiodon. Even considering possible postmortem changes
which usually tend to darken delphinids that have been dead for more than
a few hours, the pigmentation of this specimen appeared darker in the photo-
graphs than any specimen of S. plagiodon from the western Atlantic that we
have seen alive or illustrated. On the other hand, the general conformation of
the body of the African specimen, the general patterning of its light spots
and the tooth counts are similar enough to western Atlantic S. plagiodon to
warrant positive consideration for the possible occurrence of this species in
the eastern Atlantic. However, until a direct comparison of specimens from
both sides of the Atlantic is made, both on a living or freshly-dead and a
skeletal basis, we prefer only to draw attention to the African specimen and
for the present question the occurrence of S. plagiodon outside the western
Atlantic.
Ecological Distribution
Caldwell (1960) summarized data which indicated that the spotted dol-
phin is primarily an offshore species ( 12 miles or more) in the Gulf of Mexico
and elsewhere in its range. However, data were also presented which recorded
the occurrence of this species much closer inshore than is usually expected ( 3
miles from the nearest land in mid-April, and 7 miles in mid-July). Sport fish-
4
Contributions in Science
No. 104
ing captains working out of Destin, Florida, corroborated this general offshore
distribution. It is the belief of these fishermen that spotted dolphins generally
occur five miles or more offshore in waters of about five fathoms or greater,
and that the dolphins are replaced inshore of these general points by the com-
mon inshore porpoise of the region, the Atlantic bottlenosed dolphin, Tursiops
truncatus (Montagu). Raymond M. Gilmore told us in early 1966 that he had
found a similar spatial distribution for S. plagiodon and T. truncatus off Rock-
port, Texas, in late March, 1951, and Cliff Townsend told us in early 1966 that
these two species usually distribute themselves in a similar spatial manner in
the vicinity of St. Augustine, Florida. However, Mr. Townsend added that in
that region S. plagiodon are rarely seen closer inshore than about nine miles off
the outer beaches, and then only in summer. He further stated that while typi-
cal T. truncatus of some eight to nine feet in maximum length are found in the
back rivers and estuaries and within five or six miles offshore of the outer
beaches, he has seen several very large Tursiops, some 12 feet in length, only
at a point about 40 miles offshore of St. Augustine.
It is thus of considerable interest in this regard to record the seasonal oc-
currence of spotted dolphins close inshore in the vicinity of Destin and Fort
Walton Beach, Florida.
During the period from mid-April to mid-May, spotted dolphins appear
regularly each year so close inshore that they are collected for display by the
same means that are used for capturing Tursiops ; namely, by seines worked in
very shallow water (usually wading depth) immediately adjacent to or within
less than a mile of the beach (see Caldwell, Caldwell and Siebenaler, 1965: 4,
for a description of technique).
Two animals were captured near the beach in the vicinity of Destin in late
spring in 1964 and were sent to the Aquatarium at St. Petersburg Beach,
Florida, where we observed them in late June of the same year. Several addi-
tional spotted dolphins were taken at the same time but these failed to survive
for any length of time.
On May 10, 1965, a portion of a school of spotted dolphins was captured
close to the beach near Crystal Beach pier, about five miles east of Destin. Two
of these, both females, 51 and 89 inches in snout to caudal-notch length (Figs.
1 through 5), believed not to be mother and daughter by the collectors, were
kept at the Gulfarium where we studied their phonations and general captive
behavior in June of the same year. Several months after this, the adult of this
twosome died of unknown causes and her skull and most of the postcranial
skeleton are now in the collections of the Los Angeles County Museum of
Natural History (No. 27057). Another from this school, a female of adult
size but undetermined exact length (Fig. 6) died after about one day at the
Gulfarium. Two more of this school, an adult female about 90 inches in length
and a juvenile male about 63 inches in length, believed by the collectors to be
mother and son because the young animal sometimes apparently attempted to
nurse, were sent to Cape Coral (near Ft. Myers), Florida. Two additional
1966
Spotted Dolphin Biology
5
Figure 1 Stenella plagiodon. Live adult female (89 inches in snout to caudal-notch
length) and juvenile female (51 inches in snout to caudal-notch length) collected
near the beach on May 10, 1965, about five miles east of Destin, Florida, in the
northeastern Gulf of Mexico. Shown here stranded on the bottom of their holding
tank when it was drained at Florida’s Gulfarium in late June, 1965. Dark flecks on
the sides of the small animal are dirt and not pigment. Note the slightly darker
trailing edges of dorsal and pectoral fins of juvenile. (Photograph by Leonard P.
Hutchinson.)
animals (Fig. 7) from this school, both females, about 48 and 90 inches long,
were sent to the Aquarium of Niagara Falls, New York. We were told that the
school also included adult males, but the number was not noted and those cap-
tured were released as probably unsuitable for the display purposes intended
for the others.
The Destin sport fishing captains were of the opinion that spotted dolphins
are found all year round in that region offshore, but that the numbers markedly
increase during the spring and summer months. This information, along with
the especially noteworthy occurrence of the species close inshore in late spring,
suggests corroboration for the suggestion by Moore (1953: 132) that there is
an annual migration by this species into the waters of northern Florida during
the warmer months, or even in the winter months when the water is unusually
warm. On the other hand, data given us by Cliff Townsend indicate that the
migration more likely may be a matter of inshore-offshore movement in the
same latitudes rather than an alongshore migration between different latitudes.
The comments of the Destin fishermen suggested this possibility, but Mr.
Townsend’s notes provide even more positive evidence for such a phenomenon.
During the course of his collecting activities for Marineland of Florida, Mr.
Townsend has noted that in January and February, off St. Augustine, most S.
plagiodon are found some 90 miles offshore near the western edge of the Gulf
Stream. By April and early May he found that they were present some 40 to 50
miles offshore, and that by late May or early June they were 10 to 20 miles off-
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Contributions in Science
No. 104
Figure 2. Stenella plagiodon. Same two animals as shown in Figure 1. Note im-
maculate ventral surfaces of juvenile.
shore. During the rest of the summer they occurred closest to the beach (about
nine miles) and then as the year progressed the movement was reversed until
by winter they were again most common some 90 miles offshore.
The Destin fishermen indicated that spotted dolphins are frequently seen
up to 50 miles offshore. While in this case 50 miles is the usual offshore limit
for the fishing boats, Caldwell (1955: 468; 1960: 135) failed to find spotted
dolphins in the Gulf of Mexico at distances greater than 166 miles from the
nearest land, and in most cases the distances were 70 miles or less. The cruises
on which those data were gathered included numerous opportunities to look
for cetaceans at much greater distances offshore and a cetacean watch was
maintained from the bridge of the vessel during all daylight hours and when
the ship was running at night. The first of these cruises included a complete
transect across the Gulf of Mexico during which spotted dolphins were seen
within the above limits of distribution as the vessel moved offshore and several
days later when it again approached the land on the other side, but none were
seen in the middle of the Gulf (Caldwell, 1955: fig. 1). Consequently, while
S. plagiodon normally appears to be an inhabitant of offshore waters at dis-
tances greater than about five miles from the nearest land, present evidence
suggests that there may also be a general maximum limit to the distribution of
this species of about 150 to 200 miles, with a usual maximum of some 75 to
100 miles. Furthermore, the lack of West Indian records suggests that this
species occurs within the aforementioned spatial limits only off mainland
shores. The one possible record of this species from the eastern Atlantic fits
this same distributional pattern.
Further evidence for a primarily offshore occurrence of S. plagiodon
1966
Spotted Dolphin Biology
7
Figure 3. Stenella plagiodon. Same two animals as shown in Figure 1. Note light
caudal keel and dark underside of caudal flukes of juvenile; the trailing edge is
darker than the rest of the fluke.
comes from the fact that there is only one positive report of a stranding, even
though there has been a special effort in recent years to record such data.
Caldwell and Golley (1965: 25) included an earlier stranding record by Brim-
ley of an individual stranded at Cape Hatteras, North Carolina. The stranding
of an offshore-living animal at Cape Hatteras is of reduced significance when
one considers that dead or dying animals from more offshore waters are likely
to be deposited on the shores of the Cape by the Gulf Stream as it flows north
and washes the Cape. Moore (1953) and Layne (1965) failed to report
strandings of the spotted dolphin in Florida, as did Gunter (1954) from any-
where in the Gulf of Mexico. There are skulls of S. plagiodon in collections
from unstated circumstances of collection that possibly may represent strand-
ings. However, with the Cape Hatteras exception, museum specimens which
have good data on how they were obtained came from animals collected alive
offshore (e.g., True, 1885, 1889; Moore, 1953; Schevill and Watkins, 1962;
Caldwell and Golley, 1965), or, as in the case of the Destin records, from
animals occurring close inshore as a regular part of their seasonal life history.
It should be emphasized, however, that the seasonal distribution of S. plagi-
odon close inshore at Destin apparently is very unusual and perhaps unique.
While young animals are included in the schools of some 25 to 30 animals
that move close inshore near Destin in the late spring, no young were observed
in the wild that were still nursing. However, we have been told that rarely they
apparently do try to nurse in captivity, although they readily take dead fish as
food. The attempts by these animals to nurse apparently are so infrequent as
to be of little significance other than an artifact, perhaps for emotional security,
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Contributions in Science
No. 104
Figure 4. Stenella plagiodon. Same two animals as shown in Figure 1, showing pig-
ment pattern of dorsal surfaces.
from younger days (see Caldwell and Caldwell, 1966: 764, for a summary of
data concerning long nursing periods, after weaning, in captive T. truncatus) .
Homosexual behavior, which can easily be misinterpreted as nursing behavior,
may also be involved under these circumstances as well (see Brown, Caldwell
and Caldwell, 1966: 14). These young spotted dolphins, while readily taking
food fish instead of nursing (if that is what they are actually doing), still lack
the spotting of the typical adult coloration.
The adult female captured on May 10, 1965, and kept at the Gulfarium
(see above) gave birth to an unspotted female calf (Fig. 13) there on about
August 10, 1965. J. B. Siebenaler reported that the calf made only feeble
movements after birth and then immediately died. No details of the birth were
1966
Spotted Dolphin Biology
9
Figure 5. Stenellci plagiodon. Same adult animal as shown in Figure 1. Photograph
is distorted due to surface ripple on the water, but shows the degree of spotting on
the ventral surface and immaculate area in genital region.
noted, and we do not even know if the small calf was full term. However, it
was certainly nearly full term, and by August the school of spotted dolphins
would have long been back in its expected offshore habitat and the calf would
have been born at sea at that time or at still later date. The calf was much too
well developed to have been conceived inshore in May of that same year.
We suggest, therefore, that the young are born offshore and that they are
not brought close inshore by their parents until they are weaned. Consequently
the seasonal move inshore in the Destin area is clearly not for the purpose of
calving.
Over the years it has been the observation of Dewey Destin, a master
commercial seine fisherman who captures both on a commercial basis in the
Destin area, that the arrival of the spotted dolphins along the beach in May
seems to coincide with the late spring arrival of large numbers of “hardtails^
carangid fishes of the genus Caranx— usually C. crysos (Mitchill). However,
while these fish usually remain in the area all summer, according to Mr. Des-
tin, the dophins soon move back offshore. Consequently, while feeding should
be considered as a motive for the movement inshore, this possible source of
food may be coincidence rather than the reason for the movement, en masse ,
by the dolphins into the shallow waters. The spotted dolphins that have been
held in captivity at Destin and Fort Walton Beach do readily feed on dead
“hardtails’’ but the one stomach of a wild-caught spotted dolphin that was
examined, from an individual that died shortly after capture, contained large
10
Contributions in Science
No. 104
Figure 6. Stenella plagiodon. Autopsy of adult female collected with the animals
shown in Figure 1. Especially note pigmentation on ventral surfaces of head. (Photo-
graph by Leonard P. Hutchinson.)
numbers of small cephalopod beaks. Unfortunately, none of these beaks were
saved, and a photograph of a mass of them lining the stomach of the dolphin
was insufficient for Gilbert L. Voss of the University of Miami Institute of
Marine Science to provide an identification. On the other hand, fish must form
at least a portion of the natural diet of S. plagiodon as evidenced by observa-
tions made at sea by Mr. Townsend. He reported to us in 1966 that he had
observed spotted dolphins off St. Augustine feeding on small fishes of the
families Clupeidae (herrings) and Engraulidae (anchovies), and somewhat
further offshore on carangid fishes of the genera Decapterus or Selar. Captive
spotted dolphins at the Gulfarium readily take dead fishes of the latter two
genera.
1966
Spotted Dolphin Biology
11
Figures 7. Stenella plagiodon. Live adult female (about 90 inches in length) and
juvenile female (about 48 inches in length) in captivity at the Aquarium of Niagara
Falls, New York. These animals were collected at Destin, Florida, with those shown
in Figures 1 through 6. (Photograph courtesy ol the Aquarium of Niagara Falls.)
When held in captivity, it has been observed and reported to us that pairs
(either adults or adult and young, of any combination of sexes) or more
spotted dolphins seem to survive better than single animals. Reportedly this
holds true whether the spotted dolphins are segregated or in a community tank
with other kinds of dolphins. However, that this is not always a prerequisite
for survival in captivity is evidenced by the long-time survival of a single fe-
male spotted dolphin at Marineland of Florida (Figs. 1 1 and 12). This animal
was collected on August 31, 1956, and at this writing (April, 1966) is still
living and healthy although there have been many, often extended, periods
when no other spotted dolphins were in the tank with her. When we visited
Marineland in April, 1966, and observed this and one other spotted dolphin
then with her, Mr. Townsend told us that the long-captive female furthermore
is unusual because she has lived so well in a community tank with a number of
T. truncatus (see Fig. 11). Mr. Townsend noted that most spotted dolphins
are completely dominated by the bottlenosed dolphins in such a tank and that
the spotted dolphins are usually unable to compete under these circumstances.
He further noted that spotted dolphins as jumpers generally are erratic in their
aim and hence are not satisfactory as performers under conditions where the
attraction uses guest porpoise feeders. However, these dolphins will sometimes
satisfactorily perform in this manner (Fig. 12).
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Contributions in Science
No. 104
Figure 8. Stenella plagiodon. Adult of undertermined sex and size, probably col-
lected at sea off St. Augustine, Florida. Photographed by D. K. Caldwell at Marine-
land of Florida about 1955. Especially note the heavy lateral and ventral pigmen-
tation in this animal as compared to the lighter pigmentation of the other illustrated
animals.
Coloration
Despite a large number of published illustrations, the color pattern of S.
plagiodon has, until now, remained imperfectly known, particularly with re-
gards its changes in relation to the size (and presumably age) of the animal.
The illustrations of adult S. plagiodon that we have seen range from some-
what inadequate and generalized drawings (e.g., Elliot, 1904: fig. 21; Kellogg,
1940: 75; Burt and Grossenheider, 1952: 173; Hoffmeister, 1963: 47)
through adequate drawings made from specimens or photographs (e.g., True,
1885: pis. 1 and 2, 1889: pi. 18, retouched photograph; Elliot, 1901: fig. 9,
based on True, 1889; Nishiwaki and Yabuuchi, 1965: 180f.) to actual photo-
graphs, from various angles, of living or freshly-dead animals (e.g., McBride,
1940: 24, 26; Moore, 1953: 130; Schevill and Watkins, 1962: 9, and record
cover; Schevill, 1962: 2f.; Belkovitch, Kleinenberg and Yablokov, 1965: be-
tween pages 120 and 121, same photograph as the one appearing herein as Fig.
8). Schevill told us in 1965 that the specimen illustrated in the booklet of
Schevill and Watkins (1962: 9) is an animal taken from the group observed
and recorded off Cape May, New Jersey, and that the skull is now in the col-
lections of the Museum of Comparative Zoology at Harvard (MCZ 51074).
Schevill further told us that the animals illustrated in Schevill (1962: 2f.) and
Schevill and Watkins, 1962: record cover) were photographed at sea off Pen-
sacola, Florida, in the northeastern Gulf of Mexico.
1966 Spotted Dolphin Biology 13
Figure 9. Stenella plagiodon. Adult female (83 inches in length) collected at sea off
Rockport, Texas, on March 22, 1951. Note variations in pigmentation between this
specimen and the animals shown in other figures herein. (Photograph courtesy of
Raymond M. Gilmore.)
In addition to these various illustrations, the color pattern of adult S.
plagiodon was described in considerable detail by True (1885: 318) and in
much less complete, but useful, detail by Hall and Kelson (1959: 818), Kel-
logg (1940: 83), Gunter (1954: 548), True (1889: 164). The adult descrip-
tion by Palmer (1954: 332) is incorrect.
As shown by the photograph in Schevill (1962: 2f.) and Schevill and
Watkins (1962: record cover) and by our own experience (see Figs. 1 through
12), the degree of spotting (i.e., number of spots) is somewhat variable in
adults although the general pattern of spotting remains the same. There is also
some variation in the coloration of the snout and lower jaw (ranging from
light to dark) as pointed out by Caldwell (1960: 136) and suggested by the
photograph in Schevill, (1962: 2f.) and Schevill and Watkins (1962: record
cover).
Fraser (1950: 63), using the descriptions of True (1885, 1889), com-
pared S. plagiodon with a spotted dolphin from off the Atlantic coast of Africa
which he referred to the species S. frontalis (Cuvier). True’s descriptions and
figures, while very useful, are lacking in certain specific and minute details and
Fraser understandably erred in stating the plagiodon lacks white on the snout
tip or chin (see our comments on this above) and that plagiodon lacks a dark
band from the angle of the mouth to the flipper (while it is diffuse, such a band
does exist in plagiodon as shown herein in Figs. 5, 9, 10, and 11). Fraser was
correct in differentiating his specimen of frontalis from plagiodon on the basis
of the gray belly of the former as compared to the white belly of the latter.
However, for the sake of clarity it should be noted that while the base color of
the belly of plagiodon is pure white, it is for the most part covered with large
dark blotches and spots in the adult. In comparing the photographs of Fraser’s
frontalis with those of plagiodon included here (especially our Figs. 6 through
12), it becomes apparent that the spotting on the lower posterior surfaces of
the mandible of Fraser’s frontalis consists of light spots on a dark ground.
14 Contributions in Science No. 104
Figure 10. Stenella plagiodon. Captive of adult size of undertermined sex and exact
size at Marineland of Florida; collected at sea off St. Augustine, Florida. Note un-
usually sparse spotting. Apparently this same individual, or one equally sparsely
spotted, was figured less clearly by Lauber (1963: 25) and Kay (1964: 24). (Photo-
graph courtesy of Marineland of Florida.)
while in plagiodon the spots are apparently always dark on a light ground. In
addition, the mandibular spots on the specimen of frontalis appear to be rela-
tively smaller than those of plagiodon.
Knowledge of the coloration of the young of S. plagiodon is much less
complete, and as noted previously (Caldwell, 1955: 470), it is so different that
it may well be the basis for some reports of mixed schools of dolphins offshore
( e.g ., Nichols, 1920). Kellogg (1940: 85) first noted that the small calf is
gray and so illustrated it in general terms (p. 75). Palmer (1954: 332) stated
that the calf is gray, but neither he nor Kellogg gave further details. Lowery
(1943: 256) noted that S. plagiodon in Louisiana waters “is sometimes
spotted’’ which suggests a knowledge of the ontogenetic differences in colora-
tion in this species. Gunter (1954: 548) incorrectly indicated that the calves
are uniformly gray, although in an earlier report (Gunter, 1941) he had in-
cluded the more correct note that a 5-foot specimen from Texas had been
described to him as having been solid grayish-black on top and lighter under-
neath. Actually, the color of the young shades from dark purplish-gray on the
dorsal side to white on the ventral, and this can best be described by reference
to the figures we include herein (Figs. 1 through 4, and 13). The trailing edges
of the flukes of the young are notably darker than the rest of the fluke. As
noted elsewhere, the darker ground colors of living animals contain consider-
able purple (see Kellogg, 1940: 75; Gunter, 1941 ) and this is the case for both
adults and young (see Kellogg, 1940: 75; Caldwell, 1955: 470, 1960: 136).
We were able to reconfirm this purplish ground color in our close-hand obser-
vations on captive animals.
Although their text indicated that it was spotted, and Gilmore told us in
1965 that Mahnken had told him such was the case, the photograph of a wild
1966
Spotted Dolphin Biology
15
Figure 11. Stenella plagiodon. Captive female of adult size at Marineland of Florida;
collected at sea off St. Augustine, Florida. This is the same animal that appears in
Figure 12. (Photograph courtesy of Marineland of Florida.)
S. plagiodon included by Mahnken and Gilmore (1960) seems instead to be
an unspotted young animal. It may be that the spotting was sq faint or incom-
plete that it was just developing and so showed up more clearly at sea or on
the original photograph than in the published reproduction. The size of the
sucker-fish ( Remilegia australis Bennett) attached to the dolphin would also
indicate that the latter was a small animal even if the sucker-fish was a large
example of its species (known to reach a maximum total length of some two
feet, according to Radford and Klawe, 1965). The development of the spotted
coloration takes place gradually, and while to our knowledge it has never been
described in detail and to completion, Mr. Gregory Siebenaler told us in 1965
that a small captive animal that had been completely unspotted at capture in
May was beginning to develop spots on the dorsum, posterior to the dorsal fin,
about one month later. This animal, a male, was still partially unweaned and
was approximately 63 inches in snout to caudal-notch length. Another young,
a female captured at the same time (in May) as the small male just discussed,
was 51 inches in snout to caudal-notch length when we studied here in mid-
June. She had no spots at that time, but Mr. Siebenaler told us in mid-Novem-
ber of the' same year (1965) that this animal was somewhat larger and was
16
Contributions in Science
No. 104
Figure 12. Stenella plagiodon. Captive female of adult size jumping for food at
Marineland of Florida; collected at sea off St. Augustine, Florida. This is the same
animal that appears in Figure 11. (Photograph courtesy of Marineland of Florida.)
beginning to develop the darker and somewhat spotted tongue pigmentation of
the adult, and that sometime later she began to develop spots on the side of the
head. Unfortunately, she died during the winter of 1965-66 so that this de-
velopment could not be traced further. However, in early 1966 Mr. Townsend
wrote, and later told us in conversation, that it was his impression that in cap-
tive subadult-size S. plagiodon at Marineland of Florida the spots first appear
low on the sides of the animal, the full length of the body, and then gradually
Figure 13. Stenella plagiodon. Dorsal, ventral and both lateral views of preserved
unspotted female young (about 30.5 inches in snout to caudal-notch length) which
died immediately after birth in captivity at Florida’s Gulfarium. The spotted mother
is the animal shown in Figures 1 through 5. Note the similarity of the pigment pattern
1966
Spotted Dolphin Biology
17
of this specimen to that of the juvenile S. plagiodon shown in Figures 1 through 4,
and also note the text for important details of comparative pigmentations between
the two unspotted animals. Also note the single row of hairs, typical of newborn
delphinids, on the dorso-posterior part of the snout near its junction with the melon.
(Photographs by Armando Solis, Los Angeles County Museum of Natural History.)
18
Contributions in Science
No. 104
spread upward with increase in size and age so that an animal has almost its
full complement of spots about the time it reaches adult size. It was Mr. Town-
send’s impression that the process from an unspotted to a spotted animal takes
some six to eight months. He also noted that most of the spots are acquired on
the ventral surface during this period, but that some additional ones may ap-
pear there after the lateral and dorsal spotting has fully developed. Special
note should be made that according to Mr. Townsend the development of
spotting does not take place until the individual has reached nearly an adult
size, and that once the spotting has developed the particular pattern developed
by an individual essentially remains the same throughout the remainder of its
life. This is certainly true of captive animals and there seems to be no reason
to assume that it would be different in wild individuals. The late development
of the spotting could easily explain the report by Nichols (1920) of mixed
schools of spotted and unspotted dolphins which he assumed consisted of more
than one species. The size differential between the spotted and unspotted in-
dividuals possibly was not great enough to suspect that spotted adults and
unspotted young were involved.
Unquestionable proof that the young of S. plagiodon are not spotted is
shown by an event which took place at the Gulfarium. The spotted adult fe-
male (Figs. 1 through 5), that we studied there, later on gave birth to a totally
unspotted calf (Fig. 13) that was clearly conceived in the wild since the
mother’s only companion in captivity was a juvenile female spotted dolphin
captured with her. The calf died immediately after birth and is preserved in-
tact in the collections of the Los Angeles County Museum of Natural History
(No. 27058). We did not see the calf before it was preserved but except for
the slightly darker overall pigmentation, especially on the under surfaces
(which may be an artifact of preservation), the color pattern of the newborn
calf is essentially the same as that of the live juvenile we studied (compare
Figs. 1 through 4, of the juvenile, with Fig. 13, of the preserved calf). The
newborn calf clearly lacks spots, has the light line running posterio-ventrally
from the eye to the pectoral flipper base, the separate dark line on the flank
below the dorsal fin and is beginning to show the somewhat streaked pattern-
ing on the flank anterior to the dorsal fin. The pectoral flippers are dark in the
calf as in the juvenile, and the low keel formed where the caudal peduncle
joins the ventral surface of the flukes is lightly pigmented in both animals.
There is no doubt that the pigmentation pattern of the unspotted newborn
calf out of a spotted mother was the same as that of the live unspotted juvenile
we studied, or, consequently, that the live juvenile from a school of spotted
animals in turn was S. plagiodon.
The adult animal that we studied alive at Destin (Figs. 1 through 5) and
the Texas adult (Fig. 9) captured by Gilmore had light spotting on the leading
edges of the caudal flukes. Otherwise the flukes appeared to retain the homo-
geneous dark coloration (except for the darker trailing edge) shown by the
1966
Spotted Dolphin Biology
19
juveniles. True (1885: 318), on the other hand, indicated that the rest of the
fluke of a dead specimen that he studied bore spotting like that of the other fins.
Except for the ventral surface of the body around the genital region which
remains white, the rest of the body of the adult (and including the dorsal and
pectoral fins and the tongue) bore some degree of spotting from light on dark
on the dorsal surfaces to dark on light on the ventral (Figs. 1 through 9).
The marked variation in color pattern between young and adult apparent-
ly is somewhat unusual in Cetacea. Similar variation, in which the young is
essentially plain and the adult spotted, has been reported for the narwhal,
Monodon monoceros Linnaeus ( e.g ., Fraser, 1937: 284; Kellogg, 1940: 80)
and for Stenella graffmani (Lonnberg) in the eastern North Pacific (John H.
Prescott, pers. conversation, 1966, told us that an unspotted calf of this species
had been taken from the uterus of a spotted mother). The two varieties of
Prodelphinus (=Stenella) froenatus (F. Cuvier) listed by Liitken (see True,
1889: pi. 19) may be another example of a spotted adult and an unspotted
young (as suggested to us by F. C. Fraser, pers. comm., 1965). Less marked
differences, in which the young has a white belly and anterior part of the head
while the adult is a solid color, have been reported in Risso’s dolphin, Grampus
griseus (G. Cuvier) (e.g., Flower, 1874: pi. 1; Millais, 1906: pi. following
p. 310, after Flower; Richard, 1936: pi. 7). Species of cetaceans with adults
and young of different colors or shades of the same color, but with the same
color pattern, are not as unusual.
If in the development of coloration ontogeny follows phylogeny to some
degree, then it appears that the plain gray coloration is the more primitive,
followed by shades of gray and finally by complicated patterns of some kind
such as the one shown herein for S. plagiodon. Another aspect of delphinid
coloration that is found in several species and thus may be a primitive charac-
ter is a “smear” of the lighter color usually found on the sides of the animals
which extends dorso-ventrally into the darker dorsal coloration, in the region
generally beneath the dorsal fin (see Figs. 1 through 4 and 7). Although not
obvious in the figures included herein, the young plagiodon also showed this
“smear” of lighter color into the darker. We have also seen this same kind of
coloration in Atlantic and Pacific forms of Tursiops and in other species of
Stenella from the Pacific, all of which have a basic coloration of a dark dorsal
shading into a lighter ventral region.
Yablokov ( 1963: 41 ) noted that cetaceans with spots of the kind described
herein for S. plagiodon are usually species with gregarious habits and sug-
gested that the spots serve as recognition marks. Stenella plagiodon certainly
is a gregarious species, but while there may be some basis for Yablokov’s
second suggestion, we believe that it is more likely that the spots serve as camo-
flage to help protect the animals from possible predators such as killer whales,
Orcinus orca (Linnaeus), in the sparkling and often white-capped waves which
are common in their normal open-sea environment.
20
Contributions in Science
No. 104
Breathing
When observed unseen from the higher roof of an adjacent building, the
two captive spotted dolphins were seen to engage in breathing in three ways.
First, when swimming around the tank in a leisurely and undisturbed
manner with no humans in sight, the animals surfaced on an average of about
once every 22 to 23 seconds. Although difficult to measure, the actual time of
the blow while the animal was on the surface lasted about half a second, and
no more than one second. The juvenile apparently blew slightly faster than the
adult, but our equipment did not permit a precise measurement and this ob-
servation is subjective. Although the range of values for the interval between
blows was essentially the same for the juvenile and adult animals, and only
slightly more than one second difference was calculated in mean values for
a number of blows, our data show that the young animal actually tended to
blow more often than the adult. McBride and Kritzler (1951: 257) made
similar, but more marked, observations on differences in average times be-
tween blows in Tursiops truncatus of different ages, and Layne and Caldwell
(1964: 91) did the same in Inia geoffrensis (Blainville) . While swimming in
this leisurely manner, the two spotted dolphins gently rolled between breaths
and showed progressively only their snout, blowhole, predorsal surface of the
back, and dorsal fin. They did not show their flukes. Similar behavior by wild
spotted dolphins was noted by Caldwell ( 1960: 135).
In the second type of breathing behavior, the two animals frequently en-
gaged in a pattern which we called “standing on the head!’ At such times the
animals would take from two to five short breaths at intervals of two to 12
seconds, and then arch over to stand vertically in the water, head down, for
from 28 to 50 (usually the upper end of this range) seconds before returning
to the surface for another series of short blows. When “standing on their heads’’
completely submerged, the two animals maintained their vertical position by
gently waving their caudal flukes. In this position the tip of the rostrum did not
touch the bottom of the deep pool, but instead the animals were actually sus-
pended in the water. When they returned to the surface from this position, the
animals stopped waving their flukes and their bouyancy brought them to the
surface tail first— which they gradually arched into the horizontal plane parallel
to the surface so that the entire body reached the surface while still submerged.
At that time the head broke the surface for the series of short breaths. When
the animal again dove, it showed its flukes momentarily above the surface as
it completed the dive by either arching straight over and down or arching over
and down in a corkscrew fashion so that once submerged its body axis was
some 90° to its position on the surface before the dive.
We suspect that the first two forms of breathing are related in the first
instance to normal swimming to get from one point to another, and in the
second instance to deeper diving probably associated with feeding. The second
form of breathing, in which the animal appears to hyperventilate in prepara-
tion for a deep dive, is reminiscent of the behavior of the sperm whale.
1966
Spotted Dolphin Biology
21
Physeter catodon Linnaeus, under such circumstances (see Caldwell, Cald-
well and Rice, 1966).
In the third form of breathing, the captive animals at the Gulfarium some-
times rested in a horizontal fashion just below the surface, rising to blow only
infrequently, but we never observed them to rise to blow as infrequently as the
once every 120 seconds that Caldwell (1955: 468) observed in wild individ-
uals.
Frequently, just before taking a food fish held in the hand of the feeder, the
captive animals released large amounts of air in a single large bubble under-
water. At this time the animal was usually positioned motionless underwater
before the feeder and appeared to be uncertain as to whether or not to accept
the fish. During the period of our study the animals had been captive for only
about a month and appeared to still remain very wary of humans. The captive
Inia geoffrensis studied by Layne and Caldwell (1964: 91) on occasion re-
leased similar large bubbles of air underwater.
Sometimes, when swimming at the surface, tne spotted dolphins blew
very explosively, and sometimes even caused a slight “spout” when water pos-
sibly present in the vestibule was blown out. Lawrence and Schevill (1956:
135) made similar observations in T. truncatus, as did Layne and Caldwell
(1964: 91) for Inia geoffrensis. At the time, we suspected that this behavior,
like the release of the large air bubble noted above, was related to the wari-
ness of the spotted dolphins and that it served to convey the disturbed attitude
of the animals.
Play
At Marineland of Florida the younger T. truncatus have devised a game
of catch with human observers who are often positioned around the open top
of the dolphin community tank in order to watch the feeding show. In this
game the dolphins grasp a small rubber ball in their mouth and then toss it out
of the tank in the direction of the human participant. The dolphins exercise
remarkable aim, and the human is then apparently expected to catch the ball
and return it to the waiting dolphin. Mr. Townsend told us in 1966 that the
older adult T. truncatus do not participate in this particular activity (although
they play with objects underwater) , but that the fully adult S. plagiodon do. An
adult female spotted dolphin (“Dottie”), who has been in captivity at Marine'
land of Florida for ten years, still takes part in this play. We observed this first-
hand in early April, 1966, and also noted that a young male spotted dolphin,
in captivity only about three weeks at that time, had also learned this activity
and was performing it accurately.
Sound Production
The vocalizations of two female S. plagiodon, 51 and 89 inches in snout
to caudal-notch length, were recorded at the Gulfarium. The animals were 28
days captive when the recording sessions began and they were recorded inter-
mittently during the two weeks following.
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Contributions in Science
No. 104
Wood (1953: 122) noted that this species is less vocal than the oft-studied
T. truncatus. Our studies reinforce this statement, particularly in the “whistle”
component. However, on the rare occasions when whistles were emitted, there
were several in succession and very loud. There were two contours that made
up most, but not all, of the contour repertoire, and this leads us to believe that
individual spotted dolphins are characterized by a signature whistle as has
been suggested for T. truncatus by Caldwell and Caldwell (1965). Schevill
and Watkins (1962: phonograph record) recorded “squeals” (^“whistles”)
from a number of wild S. plagiodon and a sonagram of these phonations has
been published (Schevill and Watkins, 1962: 9; Schevill, 1962: 3).
During the times when the two animals that we studied at the Gulfarium
were clearly in acoustical isolation, a “chirp” was also recorded. This is a brief
pure tone that rises in frequency. We do not classify this as a true whistle con-
tour because of its simplicity and universality. We have recorded the same
chirp from captive T. truncatus.
Low intensity pulsed sounds (click trains) that were directly correlated
with feeding were recorded. However, there is a possibility that acoustical
isolation may have been broken at that time between the tank with the S.
plagiodon and an adjoining tank containing T. truncatus. In view of this, addi-.
tional confirmation is needed for the click train in this species.
Additional confirmation is also needed for a number of other audible
sounds that we recorded under circumstances that we felt may have been
acoustically contaminated by the nearby T. truncatus. However, we are rea-
sonably confident that the two spotted dolphins emitted several types of pulsed
phonations that we could separate audibly on playback, and these we have
listed as “squawks” “squeaky-squawks’’ “barks’’ “growls’’ and “cracks!’ Under
even less acoustically— isolated conditions, Wood (1952; 1953) listed “whis-
tles” and “barks” which he attributed to S. plagiodon. Caldwell, Caldwell and
Evans (1966) have discussed some of the problems in the use of subjective
terms to describe cetacean phonations.
All of the sounds listed for the two S. plagiodon at the Gulfarium were re-
corded at a tape speed of 7.5 inches per second using a Uher model 4000-S
Report recorder coupled with an Atlantic Research Corporation Model LC-57
hydrophone and a special preamplifier constructed for the system by William
E. Sutherland of the Lockheed-California Company, Los Angeles. This system
had a flat capability of 40 to 20,000 cycles per second with a good signal to
noise ratio.
On August 17. 1963, a female spotted dolphin became entangled in a sport
fisherman’s line off Destin, Florida. This animal was brought to the Gulf-
arium, but only lived a few days before succumbing to injuries received at cap-
ture. However, before the animal died, Mr. Lowell Longaker of Ft. Walton
Beach was able to record whistles and click trains from it. Mr. Longaker
kindly allowed us to listen to his recordings, but we heard no additional phona-
tions which were not duplicated in our own studies of the two other captive
animals in 1965.
1966
Spotted Dolphin Biology
23
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Radford, Keith W., and Witold L. Klawe
1965. Biological observations on the whalesucker, Remilegia australis ;
Echeneiformes : Echeneidae. Trans. San Diego Soc. Nat. Hist., 14(6):
65-72.
Richard, M. Jules
1936. Documents sur les Cetaces et Pinnipedes provenant des Campagnes du
Prince Albert ler de Monaco. Resultats des Campagnes Scientifiques
Accomplies sur son Yacht par Albert ler Prince Souverain de Monaco,
94: 1-72, pis. 1-8.
26
Contributions in Science
No. 104
Schevill, William E.
1962. Whale music. Oceanus, 9(2) : 2-7.
Schevill, William E., and William A. Watkins
1962. Whale and porpoise voices. A phonograph record. Woods Hole Massa-
chusetts: Woods Hole Oceanogr. Inst., 24-p. booklet and phonograph
record.
True, Frederick W.
1885. On a spotted dolphin apparently identical with the Prodelphinus doris
of Gray. Smithsonian Ann. Rept., for 1884, pt. 2 pp. 317-324, 6 pis.
1889. Contributions to the natural history of the cetaceans, a review of the
family Delphinidae. Bull. U.S. Natl. Mus., 36: 1-191, 47 pis.
Wood, F. G., Jr.
1952. Porpoise sounds. A phonograph record of underwater sounds made by
Tursiops truncatus and Stenella plagiodon. Produced by the Marine-
land, Florida, Research Laboratory.
1953. Underwater sound production and concurrent behavior of captive por-
poises, Tursiops truncatus and Stenella plagiodon. Bull. Mar. Sci. Gulf
and Caribbean, 3(2) : 120-133.
Yablokov, A. V.
1963. On the types of color of Cetacea. Bull. Moscow Soc. Experimentation
in Nature, Biol. Sec., 68(6) : 27-41. (In Russian, with English summary)
Addendum
After this paper was in galley proof, a publication by Philip Hershkovitz
appeared (1966, Catalog of living whales. Bull. U. S. Natl. Mus., 246 : viii +
259) wherein, on page 40, he assigned the name Stenella pernettyi (Blain-
ville) to the form we have discussed herein as S. plagiodon.
True (1885: 322) and Fraser ( 1950: 64) each briefly discussed pernettyi,
which was described (1817, Nouveau Dictionnaire d’Histoire Naturelle, 9:
154) from a specimen collected in Atlantic South American waters at 16°44' S,
35° 10' W. Another animal, believed at the time to have been the same species,
was said to have been sighted earlier at 6°43' N, 25° 17' W, somewhat closer to
Africa than to South America. Neither True nor Fraser made a definite assign-
ment for pernettyi to a species represented by specimens, but each writer sug-
gested that it may have affinities with plagiodon.
None of the original material of pernettyi was saved, and so no direct
comparisons with plagiodon can now be made, and the description of pernet-
tyi is too incomplete to make non-specimen comparisons. We have examined
collector Pernetty’s figure (reproduced in Philippi, 1893, Los delfines de la
punta austral de la America del sur. Annales del Museo Nacional Chile, ( 1)
Zool., 6: 14, pi. 5, fig. 3) and find it too lacking in detail and unlike our present
understanding of plagiodon to make a case for considering the two forms
synonyms. Actually, in our estimation the figure is crude to the degree that it
1966
Spotted Dolphin Biology
27
is doubtful if it can be legitimately used to represent any known species of
delphinid.
Consequently, as True ( 1885: 322) suggested, we do not believe that it is
advisable at this time, if ever, to try to revive the name pernettyi, but instead
suggest that it be added to Hershkovitz’ list of cetacean incertae sedis. How-
ever, it must be pointed out that the capture and observational localities for the
nominal form pernettyi are in the general geographical region of capture for
the “ plagiodon ?” of Cadenat (1959) and Cadenat and Lassarat (1959) dis-
cussed above, and consequently the same species could at least in part be in-
volved—whatever it may be.
Even if the form named pernettyi could be shown to be the same as the
form named plagiodon, it would be of significance only in providing a basis for
listing plagiodon from South America. Under Article 23b of the International
Code of Zoological Nomenclature for 1964, the name pernettyi should be con-
sidered a nomen oblitum inasmuch as it has been over 50 years (73 to be
exact) since even the listed use of the name as a senior synonym was applied
by Philippi in 1893. The only other use of the name that we can find is that of
True (1885) and Fraser (1950) when they discussed it as a questionably-
identifiable nominal species in their consideration of the possible identification
of other specimens. To our knowledge, pernettyi has never been directly ap-
plied to the western North Atlantic form named plagiodon, until Hershkovitz
did so in 1966. To try to apply the name of such a poorly-described form as
pernettyi to a well-known species such as plagiodon serves no purpose.
Hershkovitz (1966: 36) is in error in placing the Prodelphinus doris of
True ( 1885) in the synonymy of Stenella frontalis G. Cuvier. There is reason
to consider the doris of Gray a synonym of frontalis (see Fraser, 1950: 68),
but this is no basis for the inclusion of the doris of True (1885) in that syn-
onymy. True (1889) clearly noted that his earlier application of the name
doris was incorrect and his material from his 1885 paper should be referred
only to the nominal form plagiodon. Hershkovitz (1966: 41) apparently was
aware of this later correction by True.
Although we did not include his remark in our discussion of juvenile
pigmentation in plagiodon above, because the species involved was uncertain,
True (1885: 322) did mention in his comments on pernettyi that the lack of
spotting on the back of the animal in Pernetty’s figure was not a significant
difference between that species and doris (= plagiodon in True’s paper).
True made this statement because he had been informed by a naturalist on
board the research vessel Albatross that the young in schools of spotted
dolphins (believed by True to be plagiodon) which had been seen off Cape
Hatteras did not have the back spotted.
Also after our manuscript was in galley proof, Dale W. Rice pointed out
a paper by Glover M. Allen (1931, Ocean dolphins. Bull. Boston Soc. Nat.
Hist., 61: 3-7) in which he illustrated (p. 4) as Prodelphinus froenatus, a small
dolphin harpooned at sea “south of Bermuda!' The pigmentation of the small
28
Contributions in Science
No. 104
animal is very like that of the young Stenella plagiodon illustrated herein
(especially the lateral view shown in our Figure 1). We suspect this may
represent a record of plagiodon from near Bermuda, but withhold final judg-
ment until more is learned of the relationships of that species and froenatus
(= frontalis)', and if they are different, as we consider them to be, until the
juvenile of the latter species is described.
Rice also reminded us of a book by F. H. Van Den Brink (1957, Die
Saugetiere Europas, westlich des 30. Langengrades. Hamburg: Paul Parey,
225 p., 32 pis.) wherein (p. 159) Stenella plagiodon is questionably placed in
the synonymy of Stenella dubia G. Cuvier. We strongly question such a place-
ment of plagiodon with some of the other species also included therein with
dubia. The drawing of the latter species (pi. 22) would be a poor likeness of
plagiodon under any circumstance.
J. B. Siebenaler told us in mid June, 1966, that the expected late spring
arrival of the spotted dolphins inshore near Destin and Ft. Walton Beach
failed to take place this year. The winter and early spring of 1966 were un-
seasonably cold in that part of Florida; so much so that even the estuarine
Tursiops truncatus for the most part moved much further offshore than usual
for that time of year— apparently to escape the cold shallow bay waters during
the sub-freezing weather. Moreover, the region just to the east suffered the
direct effects of a major hurricane in early June and the fringe of this adversely
affected the weather in the Destin-Ft. Walton Beach area. Any of these
meteorological conditions, and especially the very low water temperatures
inshore, probably were enough to deter the regular inshore movement of the
spotted dolphins.
LOS
ANGELES
COUNTY
MUSEUM
Number 105
July 22, 1966
COMPARISON OF THE EARLY PERMIAN VERTEBRATE
FAUNAS OF THE FOUR CORNERS REGION AND
NORTH-CENTRAL TEXAS
CONTRIBUTIONS
IN SCIENCE
By Peter Paul Vaughn
Los Angeles County Museum of Natural History • Exposition Park
Los Angeles, California 90007
CONTRIBUTIONS IN SCIENCE is a series of miscellaneous technical papers
in the fields of Biology, Geology and Anthropology, published at irregular intervals
by the Los Angeles County Museum of Natural History. Issues are numbered sepa-
rately, and numbers run consecutively regardless of subject matter. Number 1 was
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COMPARISON OF THE EARLY PERMIAN VERTEBRATE
FAUNAS OF THE FOUR CORNERS REGION AND
NORTH-CENTRAL TEXAS1
By Peter Paul Vaughn2
Abstract: It has been thought for a long time that the
differences between the Early Permian vertebrate faunas of
north-central Texas and north-central New Mexico were due to
the presence of a water barrier between these areas. New finds of
vertebrates in southeastern Utah, in particular the discovery in
this area of Ectosteorhachis, Diplocaulus, Seymouria, and most
recently, the long-spined pelycosaur Dimetrodon, cast doubt on
the long-term effectiveness of any such barrier. The presence of
these animals in southeastern Utah indicates special resemblance
of this fauna to that of north-central Texas. It is suggested that
this resemblance is due to environmental similarity; both north-
central Texas and southeastern Utah were deltaic regions in
Early Permian time, near the borders of persistent seaways. The
absence of these forms in the intervening areas of north-central
New Mexico and southwestern Colorado may be accounted for
by the apparently more “upland” conditions of these areas in
Early Permian time. The special ways in which all the Four
Corners faunas (southeastern Utah, southwestern Colorado, and
north-central New Mexico) resemble one another, for examples,
in the common possession of Platyhystrix and, in Utah and New
Mexico, Sphenacodon, are thought to be due to geographic prox-
imity.
Introduction
The best known Early Permian vertebrate fauna is that of the famous
redbeds of north-central Texas, summarized by Romer (1958). Also well
known is the contemporaneous fauna from north-central New Mexico, dis-
cussed by Langston (1953). The most recent general comparison of these
faunas is by Romer (1960). Lately, Lower Permian strata farther west have
begun to yield a varied assemblage of vertebrate fossils, in southeastern Utah
(Vaughn, 1962, 1964a, 1965) and southwestern Colorado (Lewis and Vaughn,
1965). These later finds have not only shown the existence of genera and
species previously unknown, but, in their demonstration of the basic similarity
of Early Permian vertebrate faunas from widely separated areas, they allow
new insight into particular paleozoogeographic phenomena. The general simi-
larity of the continental vertebrate faunas of this horizon in Europe and North
America has been discussed by Lewis and Vaughn (1965). This paper will
treat of similarity— and difference— on a more local scale: between the region
xThis study was supported by National Science Foundation grant NSF GB-1014.
2Research Associate, Los Angeles County Museum of Natural History; and De-
partment of Zoology, University of California, Los Angeles.
1
2
Contributions in Science
No. 105
of north-central Texas and the region of the Four Corners (where Utah, Colo-
rado, New Mexico, and Arizona meet).
It has been recognized for a long time that there are significant differences
between the Early Permian vertebrate faunas of north-central Texas and
north-central New Mexico. As Romer (1960:52-53) has summarized it:
“Texas . . . and New Mexico faunas differ in part in the rarity or absence in
the latter of many water-dwelling types; each has a few characteristic forms
absent in the other area; both share a number of common and familiar genera,
although it is probable that the species are different; differences in the case
of a number of rare types may be due to accidents of collecting. The general
picture is one of two essentially contemporaneous faunas, evolving along
similar lines but differing in a fashion to be expected of continental assem-
blages separated from one another by such a broad water barrier as is known
to have been present in eastern New Mexico and western Texas in early
Permian times!' Among the forms commonly found in Texas but never in
New Mexico are: the crossopterygian fish Ectosteorhachis, the nectridian
amphibian Diplocaulus, the advanced labyrinthodont amphibian Seymouria,
and the distinctively long-spined pelycosaurian reptile Dimetrodon. Dimetro-
don, as Romer (1960:52) has remarked, is the most dramatic example:
“ Dimetrodon is the most abundant form in every Texas horizon. . . !’ The
New Mexico pelycosaur phylogenetically closest to Dimetrodon is Sphenaco-
don, “definitely distinct generically but indistinguishable except for spine
development”; the neural spines in Sphenacodon are much lower, nearer to
the proportions of what was apparently normal for pelycosaurs.
Somewhat farther west, however, some of the animals common in Texas
but absent from New Mexico have been found in the Cutler sediments of
southeastern Utah. Already reported are Ectosteorhachis (Vaughn, 1962),
Diplocaulus (Vaughn, 1965), and Seymouria (Vaughn, 1966). In addition,
some of the elements rarer in Texas but totally unknown from New Mexico
have now also been reported from farther west: the pelycosaur Ctenospondy-
lus from southeastern Utah (Vaughn, 1964a) and the pelycosaur Myctero-
saurus from southwestern Colorado (Lewis and Vaughn, 1965). These finds,
and others to be noted below, tend to cast doubt on the long-term effectiveness
of any water barrier that may have existed between the Midcontinent and the
Four Corners. This doubt is intensified by the fact that Dimetrodon is now
known from southeastern Utah.
Some new explanation of Early Permian vertebrate distributions in the
western United States seems to be required. The answer would still seem to
lie in paleogeography, but with the additional consideration of newly un-
covered facts, and with attention to environmental dissimilarities. I think it is
probable that the faunas from southeastern Utah and north-central Texas
resemble one another because they lived in similar, deltaic environments, and
that they differ from the intervening New Mexico fauna because the latter
lived in a non-deltaic area; that is, that the differences between the Texas and
1966
Early Permian Vertebrates
3
the New Mexico faunas are not due to the presence of any broad barrier to
faunal distribution.
Throughout this discussion, reference will be made to Figure 1. In this
map, simplified outlines of major Wolfcampian (early Early Permian) and
early Leonardian (later Early Permian) paleogeographic features have been
superimposed onto outlines of parts of the states in the region. The positions
and extents of the positive areas, or “highlands’’ have been compiled from a
number of sources, cited in the caption. The outlines of the “seaways” will
Figure 1. Simplified map of Wolfcampian and early Leonardian vertebrate faunas,
highlands, and seaways, in the region from the Four Corners to the Midcontinent.
Faunas indicated by circles: CJ, Coyote-Jemez, in north-central New Mexico; LV,
Lisbon Valley, and MV, Monument Valley, in southeastern Utah; NT, North-central
Texas; and PL, Placerville, in southwestern Colorado. Highlands indicated by stipple:
A, Apishapa; AWC, Amarillo-Wichita-Criner Hills; B, Bravo; D, Defiance; FR,
Front Range; N, Nacimiento; P, Pedernal; R, Roosevelt; S, Sacramento; SG, Sierra
Grande; USL, Uncompahgre-San Luis; Z, Zuni. Seaways indicated by horizontal
ruling: LS, “Leonardian” seaway with approximate northern boundary — subject to
much fluctuation; EC, seaway represented by Elephant Canyon Formation. Pale-
ogeographic features compiled from maps and data published by: Baars, 1962; Hills,
1963; King, 1959; and Rascoe, 1962.
4
Contributions in Science
No. 105
receive special comment further on. The Wolfcampian and early Leonardian
faunas are indicated by encircled letters: “LV” and “MV” stand, respectively,
for the faunas known from the undifferentiated Cutler Formation of Lisbon
Valley and from the Cutler Group of Monument Valley, both of these in San
Juan County, southeastern Utah; “PL” stands for the fauna known from the
Cutler Formation near Placerville, San Miguel County, southwestern Colo-
rado; “CJ” stands for the faunas known from the Cutler and Abo Formations
in and about Coyote, Rio Arriba County, and Jemez Springs, Sandoval County,
north-central New Mexico; and “NT” stands for the fauna known from the
Wichita Group of north-central Texas. Only the Wolfcampian and early
Leonardian vertebrates from Texas are emphasized; the fauna from the higher
Clear Fork Group of Texas is of later Leonardian horizon, younger than the
Four Corners faunas, and will not be considered except incidentally— although
there are no animals known from the Clear Fork Group that would contradict
the explanations developed here. For stratigraphic correlations of these faunas,
see Dunbar, et al. (1960), Langston (1953), Lewis and Vaughn (1965),
Romer ( 1960), and Vaughn ( 1962, 1964a, 1965).
Dimetrodon in Southeastern Utah
In 1960, when I first undertook collection of vertebrate fossils from the
Cutler sediments of southeastern Utah, I expected that no trace of Dimetrodon
would be found, because of my conviction at the time that there was an effec-
tive Early Permian water barrier between the Four Corners and Texas. But
when, in addition to elements commonly found in both New Mexico and
Texas, good signs of Ectosteorhachis, Diplocaulus, and Seymouria were recov-
ered, I began to see the basic similarity of the Utah fauna to that of Texas; and
a detailed re-examination, including further preparation, of all fragmentary
materials collected during past field seasons was made in a search for Dime-
trodon. This search was rewarded by the finds of parts of characteristic neural
spines referable to Dimetrodon from six different localities low in the Organ
Rock Shale, a formation equivalent to some horizon in the upper part of the
Wichita Group of Texas, earliest Leonardian in age (see Vaughn, 1964a).
The first find was embarrassing. In my report on the vertebrates of the
Organ Rock Shale (Vaughn, 1964a), I described materials of Ctenospondylus
aff. C. casei, a pelycosaur with fairly long, laterally flattened neural spines
known theretofore only from a single specimen from the upper Wichita Group
of Texas. The Organ Rock Shale had yielded a good number of specimens, in-
cluding a skull and associated postcranial parts from one quarry. Among mate-
rials from other quarries that I thought were referable to Ctenospondylus,
I noted NTM VP 1018, a badly weathered string of three dorsal vertebrae with
partial neural spines, found near the base of the northern slope of Hoskinnini
Mesa in Monument Valley, in NW!4 sec. 8, T. 43 S., R. 14 E., San Juan
1966
Early Permian Vertebrates
5
County, Utah.3 it is now obvious that this identification was incorrect. Large
parts of the neural spines of two of the vertebrae are preserved, but about
70 mm. above the bases of the spines, the preservation is in impression only,
with the impression of only one of the spines completely bordered fore and
aft. I had thought that this rounded impression represented the posterior edge
of a distorted Ctenospondylus spine. Now, after re-examination with the aid
of a rubber mould of the impression, it is clear that what is really represented
is somewhat more than a lateral half of a subcircular spine complete with the
deep, longitudinal fore and aft grooves so characteristic of most species of
Dimetrodon (see Romer and Price, 1940). These grooves can be followed
onto the proximal part of the spine where, on the anterior surface, a thin,
longitudinal ridge appears within the groove— again as in Dimetrodon. The
lateral surfaces of the proximal portion of the spine are flattened, and the
distal, rounded surfaces are marked by longitudinal striae; these features are
also in complete accord with the pattern of Dimetrodon. The centra, badly
weathered, are about 37 mm. long and about 29 mm. high at their ends. These
dimensions are very close to those of a posterior dorsal vertebra of Dimetro-
don limbatus from the upper part of the Texas Wichita Group (Romer and
Price, 1940: table 3), but in the absence of better materials and in view of the
distance between Utah and Texas, it seems wise not to attempt specific identi-
fication at this time. The impressions of the two spines are incomplete distally;
the total length of each of the spines as preserved, including the basal bony
part and the distal impression, is about 180 mm. The better of the two impres-
sions shows that the subcircular part of the spine, about 140 mm. above the
base, had a transverse diameter of about 12 mm. The natural cast of the
neural canal shows that the floor of the canal was well ossified, as in sphena-
codontid pelycosaurs generally. The spines have been compared with vertebrae
of species of Dimetrodon on hand from both the Wichita and Clear Fork
Groups of Texas, and I am satisfied that NTM VP 1018 represents that genus.
Subsequently, more parts of neural spines referable to Dimetrodon were
turned up in collections from the Organ Rock Shale. These are catalogued as:
UCLA VP 1675, collected by J. R. Dyer in 1950 about 2 miles east of Mitchell
Butte in the Arizona part of Monument Valley; NTM VP 1039 from NE14
sec. 11, T. 43 S., R. 15 E., NTM VP 1040 and 1042 from SW14 sec. 12, T.
43 S., R. 16 E., NTM VP 1041 from about 2 miles southeast of Monument
Pass in approximately the southern part of sec. 17, T. 43 S., R. 17 E., and
NTM VP 1043 from NW14 sec. 18, T. 43 S., R. 17 E., San Juan County,
Utah. Most of these catalogue numbers designate more than only one frag-
ment of spine. UCLA VP 1675 includes two fragments that show the charac-
teristic transition from a compressed, laterally flattened proximal portion to
3The abbreviations “NTM” and “UCLA” stand for the collections of, respectively,
the Navajo Tribal Museum, Window Rock, Arizona, and the University of Cali-
fornia, Los Angeles.
6
Contributions in Science
No. 105
a swollen, transversely widened distal portion. All lots include fragments that
show the dumb-bell or figure-8-shaped cross-section of the distal part, com-
pressed from front to back, wide from side to side, and with deep fore and aft
grooves. The transverse diameters of the distal fragments range from about
8 to about 13 mm.
It would, of course, be desirable to be able to identify positively parts of
Dimetrodon other than neural spines from the Organ Rock Shale. Even
partially articulated skeletons are, however, rarely found in this formation, and
the difficulty in distinction of isolated skeletal elements, other than spines, of
the three sphenacodontine pelycosaurs Sphenacodon, Ctenospondylus, and
Dimetrodon has been commented on by several authors (see Vaughn, 1964a).
For the present, it is not possible to cite more than the given six occurrences
of Dimetrodon in the Organ Rock Shale, but it may be pointed out that this
record is greater than the one occurrence apiece that has established the
presence of the pelycosaurian genera Eothyris and Ctenospondylus in the
Wichita sediments of Texas (Romer and Price, 1940).
Similarity of the Early Permian Environments and Faunas of
Southeastern Utah and North-Central Texas
The Cutler Group in Monument Valley in southeastern Utah is divided
into, in ascending order: Halgaito Shale, Cedar Mesa Sandstone, Organ Rock
Shale, and De Chelly Sandstone (see Baars, 1962). The Halgaito Shale and
Organ Rock Shale are redbeds, lithologically continuous to the East with
arkosic sediments of the undifferentiated Cutler Formation derived through
erosion of the Early Permian Uncompahgre-San Luis highland of southwestern
Colorado (see Fig. 1). The intervening Cedar Mesa Sandstone is apparently
principally of shallow-water marine origin; and the uppermost unit, the De
Chelly Sandstone, seems to be of aeolian origin in this area although its south-
eastward extension, the Meseta Blanca Sandstone, may represent marine con-
ditions (Baars, 1962). Vertebrate faunas indicate a Wolfcampian age for the
Halgaito Shale, equivalent to the lower part of the Texas Wichita Group, and
a very early Leonardian age for the Organ Rock Shale, equivalent to the
upper part of the Wichita Group (Vaughn, 1962, 1964a). The De Chelly
Sandstone seems to be of early Leonardian age (Baars, 1962). The Halgaito
Shale interfingers toward the Northwest with Wolfcampian marine carbo-
nates of the Elephant Canyon Formation (Baars, 1962). This transition has
recently been examined by the author in the region of Comb Wash, and the
picture is one of genuinely deltaic conditions, with fossils in the Elephant
Canyon facies that would seem to represent animals washed northwestward
from the Halgaito delta. This corresponds with my earlier observation
(Vaughn, 1962) that Halgaito drainage was, in general, toward the North.
As Baars (1962:169) interprets it, “The depositional site of the Halgaito was
probably a broad marginal marine mud flat, alternatively receiving fluvial
1966
Early Permian Vertebrates
7
sedimentation and periodic marine invasions!’ The Elephant Canyon beds
have also produced many marine invertebrates and remains of an actinoptery
gian fish remarkably advanced for its time; these, and other details, will be
published later. The Organ Rock Shale thins northwestward to a pinch-out
between the Cedar Mesa Sandstone and the White Rim Sandstone— a De
Chelly equivalent. The Organ Rock, too, would seem to represent deltaic
conditions; as Baars (1962:187) says, “the wide distribution and uniformity
of the unit along with its proximity to the probably marine Cedar Mesa Sand-
stone suggest that deposition was on a broad coastal plain. It is possible that
fluvial deposits were partly reworked by marine waters to provide the uniform
distribution of the beds!’ These considerations provide the basis for the out-
line of the seaway northwest of Monument Valley presented in Figure 1. The
outline is drawn to correspond with the approximate zone of facies change
between, on the one hand, the Halgaito Shale of Monument Valley and the
undifferentiated Cutler sediments of the general region of Lisbon Valley, and,
on the other hand, the Elephant Canyon Formation— as this transition is
understood from Baars’ map (1962: fig. 9) and data and from my investiga-
tions currently underway. The same outline will serve for Organ Rock time,
as may be seen in an examination of the fence diagram presented by Baars
(1962: fig. 13). In summary, the region of Monument Valley would seem to
have been deltaic in Halgaito time, covered by shallow marine water in Cedar
Mesa time, and deltaic again in Organ Rock time. Later, in De Chelly time,
conditions changed more drastically, with deposition of sand dunes, on the
slopes of which are recorded only the trackways of tetrapods (see Vaughn,
1963b).
Romer (1958:165) has described the Early Permian paleogeography of
north-central Texas, and we may draw from his observations: “in the later
Carboniferous much of Texas was occupied by shallow seas. ... To the south,
marine conditions persisted through Wichita time, but in the collecting area
the deposits of the Cisco, highest of definitely Carboniferous groups, indi-
cate a trend toward continental conditions ... [In the lowest part of the Wichita
Group] limestones and marine shales disappear and continental conditions
dominate, to continue with little interruption throughout the Permian ex-
posures in northern Texas; during most of Wichita time the present line of the
Salt Fork of the Brazos River roughly marks the boundary between marine
beds to the south and continental sediments which extend northward across
Oklahoma. The Wichita continental . . . sediments . . . are presumed to come
from an emergent land mass to the east and a mountain chain to the north in
(modern) Oklahoma. . . . The Texas collecting area appears to have been a
broad deltaic region!’ The highland north of the delta is drawn in much sim-
plified form in Figure 1 , without attention to its subdivisions. The seaway to
the South, sometimes referred to as the “Leonardian seaway!’ was of variable
extent and occasionally transgressed far northward, as during the deposition
of the Lueders Formation— a formation variously assigned to the top of the
8
Contributions in Science
No. 105
Wichita Group (Dunbar, et al., 1960) or to the base of the Clear Fork Group
(Romer, 1958). The simplified boundary of this seaway presented in Figure 1
is intended only to outline approximately the more persistently marine part of
its basin, to illustrate the essentially deltaic nature of the north-central Texas
area during Wichita time.
Lists are available of the vertebrates known from the Halgaito Shale
(Vaughn, 1962), Organ Rock Shale (Vaughn, 1964a), and the Wichita
Group (Romer, 1958, 1960). These lists will not be repeated here; suffice it
to say that the known faunas, including such familiar genera as Xenacanthus,
Eryops, Diadectes, and Ophiacodon, demonstrate general similarity and per-
mit approximate stratigraphic correlation. 1 shall confine myself for the most
part to the significant ways in which the faunas of southeastern Utah and
north-central Texas resemble one another, and to the ways in which these
faunas differ from those of north-central New Mexico and southwestern
Colorado. I shall also limit myself to discussion at the generic level, because
of the frequent difficulty of comparison of species between widely separated
areas at this horizon.
The crossopterygian fish Ectosteorhachis is known from the Halgaito Shale
(Vaughn, 1962), and also from equivalent strata low in the undifferentiated
Cutler sediments of Lisbon Valley about twelve miles south-southeast of the
town of La Sal, San Juan County, Utah (Vaughn, 1965). In Figure 1, it may
be seen that the Lisbon Valley area is also to be included as part of the deltaic
region bordering the “Elephant Canyon seaway!’ The correlation between the
Halgaito Shale and the lower Cutler strata in Lisbon Valley is now strength-
ened by recent discoveries of the rhachitomous amphibian Platyhystrix rugosus
(UCLA VP 1677) in Lisbon Valley— previously known from the Halgaito—
and the pelycosaur Edaphosaurus novomexicanus (UCLA VP 1676) in the
Halgaito— previously known from Lisbon Valley. Further, small toothplates of
a lungfish similar to Gnathorhiza are now known from both places (Halgaito:
UCLA VP 1680; Lisbon Valley: UCLA VP 1678), as are also vertebrae of
aistopod amphibians (Halgaito: UCLA VP 1681; Lisbon Valley: UCLA VP
1679). Although this is the first published report of actual parts of Early
Permian lungfishes west of Texas, their presence in northern New Mexico has
been suspected on the basis of fossilized burrows in the Sangre de Cristo
Formation (Vaughn, 1964b), similar to burrows of Gnathorhiza in Texas
(Romer and Olson, 1954). Although aistopods are unknown from New Mexico
and Colorado, the lack of record may well be due to sampling error; the
record of these very small animals in the Midcontinent is extremely scanty
(see Baird, 1964). Probably not to be accounted for by sampling error is the
absence of Ectosteorhachis in Colorado and New Mexico. Specimens of this
fish are common in the Halgaito Shale, lower Lisbon Valley Cutler, and the
Wichita Group, and this would seem to constitute a special resemblance
between the faunas of southeastern Utah and north-central Texas. It may be
noted here that in Utah Ectosteorhachis is not found above the Halgaito,
1966
Early Permian Vertebrates
9
whereas in Texas this genus is known from higher levels in the Wichita. This
may be part of a general picture of somewhat earlier onset of drier conditions in
Utah than in Texas, a phenomenon indicated by both fauna and flora (Vaughn,
1 964a) , and corroborated by the earlier appearance of lungfish of Gnathorhiza
type in Utah; in Texas, the lungfish of the Wichita Group is the apparently non-
aestivating Sagenodus, replaced in the Clear Fork by Gnathorhiza (Romer,
1958).
The nectridian amphibian Diplocaulus is known from both the Wichita
and Clear Fork Groups of Texas although it is rare in the former (Romer,
1958), and it is also known from the Halgaito equivalent in Lisbon Valley
(Vaughn, 1965) where recent field work has uncovered a fairly large number of
specimens. Diplocaulus is not known from southwestern Colorado, but there
has been only one published report of Early Permian vertebrates from there
(Lewis and Vaughn, 1965). Its absence from north-central New Mexico
would seem to be genuine, in view of the long history of collection in that area
(see Romer, 1960).
The well known genus Seymouria, an animal seemingly near the phylo-
genetic borderline between the labyrinthodont amphibians and the reptiles, is
known from both the upper part of the Wichita Group and the lower part of
the Clear Fork Group of Texas, and it has recently been reported from the
Organ Rock Shale of the Cutler Group in Monument Valley (Vaughn, 1966).
Like Ectosteorhachis and Diplocaulus, this genus is totally unknown from the
well searched Lower Permian beds of north-central New Mexico. There is a
seymouriid, not Seymouria itself, known from a single vertebra found in the
undifferentiated Cutler Formation of southwestern Colorado (Lewis and
Vaughn, 1965), and perhaps this indicates some degree of special similarity
of Early Permian conditions in southwestern Colorado to the deltaic condi-
tions in southeastern Utah— Figure 1 shows that the Colorado area lay con-
siderably closer to the boundary of the “Elephant Canyon seaway” than did
the New Mexico area. It may also be noted that the Colorado seymouriid,
from a Wolfcampian horizon, seems to be more primitive than Seymouria.
Seymouria is not known in Utah below the Organ Rock Shale, of earliest
Leonardian age. Perhaps this indicates a movement of some faunal elements
from the somewhat more upland area of southwestern Colorado into the
region of southeastern Utah. This would resemble the apparent replenishment
of faunal elements in north-central Texas from Oklahoma (see Olson, 1962).
The presence of Dimetrodon in the Organ Rock Shale as well as in the
Texas Wichita Group, and its absence from southwestern Colorado and north-
central New Mexico have already been noted, but it may be re-emphasized
that the discovery of this pelycosaur in Utah removes the most dramatic of
cited examples of Early Permian faunal differences between the Four Corners
and the Midcontinent. Also, we now know that the three closely similar sphena-
codontine pelycosaurs Dimetrodon, Ctenospondylus, and Sphenacodon all
lived together in at least one region. Ctenospondylus is known from good
10
Contributions in Science
No. 105
specimens from the Organ Rock Shale, and although the definitely determina-
ble Organ Rock materials of Sphenacodon are scanty (Vaughn, 1964a), this
genus is also known from strata approximately equivalent to the Organ Rock
in Lisbon Valley. The discovery of cranial and appendicular elements of a large
sphenacodontid at this level in Lisbon Valley has already been published
(Vaughn, 1965), and additional materials more recently collected from the
same locality— including excellently preserved vertebrae in the UCLA collec-
tions—help show that this animal is closely comparable to Sphenacodon
ferocior Romer, although the neural spines are proportionately slightly longer
in the Utah form. Lest an old theory be revived that the difference between
Dimetrodon and Sphenacodon merely represents sexual dimorphism, let it
be remembered that Sphenacodon remains unknown from Texas, and in this
connection, it may be pointed out that the difference of Ctenospondylus from
both of these genera lies in more features than only the structure of the neural
spines (Vaughn, 1964a).
The total known fauna of the Cutler sediments in southeastern Utah,
including the recently discovered lungfish, helps demonstrate the general simi-
larity of all Early Permian vertebrate faunas— at least those that lived in basins
of continental deposition. But, the presence of Ectosteorhachis, Diplocaulus,
Seymouria, and Dimetrodon in the Lower Permian of southeastern Utah, and
the absence of these forms in north-central New Mexico and probably south-
western Colorado too, would seem to constitute special resemblance of the
Utah fauna to that of north-central Texas, and perhaps the presence of Ctenos-
pondylus in both these areas may be cited as additional evidence of similarity.
There remain, however, differences between the Utah and Texas faunas. In
part, these differences reflect uniqueness; as illustration, the seymouriamorph-
diadectomorph intermediate Tseajaia ( Vaughn, 1964a) is known from the
Organ Rock Shale and from nowhere else. For the rest, the differences lie in
special resemblances of the Utah fauna to that of north-central New Mexico,
as illustrated by the record of Platyhystrix in the Halgaito Shale and its Lisbon
Valley equivalent, a genus of limnoscelid cotylosaurs in the Halgaito Shale
(Vaughn, 1962), and Sphenacodon in the Organ Rock Shale and its Lisbon
Valley equivalent. Platyhystrix and a limnoscelid are also known from the
Lower Permian of southwestern Colorado (Lewis and Vaughn, 1965).
The Early Permian vertebrate faunas of north-central New Mexico and
southwestern Colorado are alike in most respects, including the absence of
Ectosteorhachis, Diplocaulus, Seymouria, and Dimetrodon, but each also has
its own characteristics. For examples, the rhachitomous amphibian Cheno-
prosopus is known only from New Mexico where it has been found at two
separate localities (Vaughn, 1963a), and the haptodontine pelycosaur Cut-
leria is known only from Colorado (Lewis and Vaughn, 1965). The presence
of the nitosaurid pelycosaur Mycterosaurus in southwestern Colorado is in-
teresting inasmuch as this genus is known otherwise only from Texas. Perhaps
this, like the Colorado seymouriid already noted, is a consequence of the
1966
Early Permian Vertebrates
11
relative proximity of the Colorado area to the delta of southeastern Utah, but
the picture is still far from clear.
Hypothesis
That faunas from widely separated areas should each show special charac-
teristics is to be expected, and it is also to be expected that the degree of
propinquity should at least roughly affect faunal resemblance. In this way,
the Early Permian vertebrate faunas from the several Four Corners areas
(southeastern Utah, southwestern Colorado, and north-central New Mexico)
not only each contain unique elements but also show special resemblance to
one another— in the common possession of Platyhystrix and, in Utah and New
Mexico, Sphenacodon. That these two elements are not known from Texas
may be due to geographic distance and, possibly, some other obstacle to distri-
bution. What this barrier (or perhaps better, “filter”) may have been is not
clear, but it does now seem clear that any water barrier that may have existed
could not have been very effective throughout the time involved.
The similarity of the Utah, Colorado, and New Mexico faunas is thus
interpreted as possibly due to geographic proximity, but the special resem-
blance of the Utah fauna to that of Texas is probably due to a different factor
—similar environment. The remarkable fact is that, as one proceeds north-
westward from the New Mexico collecting area, he encounters vertebrates in
southeastern Utah that are also present in north-central Texas but not in the
intervening region, specifically, Ectosteorhachis, Diplocaulus, Seymouria, and
Dimetrodon. This demands explanation, and I think the answer lies in the
fact that southeastern Utah and north-central Texas were alike in Early
Permian time in that they were both deltaic regions. The north-central New
Mexico and southwestern Colorado faunas lived near the southwestern flank
of the Uncompahgre-San Luis highland, relatively far removed from the per-
sistent seaways of the time. Because all of these faunas lived in basins of
sedimentation, it is difficult to draw a contrast between “lowlands” and “up-
lands” forms, but perhaps it may be said that the north-central New Mexico
and southwestern Colorado faunas do represent somewhat more upland con-
ditions.
Further collection and study will undoubtedly modify these conclusions.
The suggestion that the southwestern Colorado fauna is to some degree inter-
mediate between those of southeastern Utah and north-central New Mexico
seems particularly worthy of further attention. The north-central Texas area
has been searched for such a long time that it seems unlikely that, for example,
Sphenacodon will be found there, but this is far from certain; such a find
would modify the detailed analysis but would only support the main thesis
developed here. Contrariwise, discovery in north-central New Mexico of any
of the genera Ectosteorhachis, Diplocaulus, Seymouria, or Dimetrodon would
12
Contributions in Science
No. 105
seriously weaken the thesis. In the meantime, the explanations offered may
be regarded as a working hypothesis.
Literature Cited
Baars, D. L.
1962. Permian system of Colorado Plateau. Bull. Amer. Assoc. Petrol. Geol.,
46:149-218.
Baird, D.
1964. The aistopod amphibians surveyed. Breviora, Mus. Comp. Zook,
206:1-17.
Dunbar, C. O., et. al.
1960. Correlation of the Permian formations of North America. Bull. Geol.
Soc. Amer., 71:1763-1806.
Hills, J. M.
1963. Late Paleozoic tectonics and mountain ranges, western Texas to south-
ern Colorado. Bull. Amer. Assoc. Petrol. Geol., 47:1709-1725.
King, P. B.
1959. The Evolution of North America. Princeton: Princeton Univ. Press,
189 pp.
Langston, W., Jr.
1953. Permian amphibians from New Mexico. Univ. Calif. Publ. Geol. Sci.,
29:349-416.
Lewis, G. E., and P. P. Vaughn
1965. Early Permian vertebrates from the Cutler Formation of the Placerville
area, Colorado, with a section on Footprints from the Cutler Forma-
tion, by Donald Baird. U.S. Geol. Survey Prof. Papers, 503-C:l-50.
Olson, E. C.
1962. Late Permian terrestrial vertebrates, U.S. A. and U.S.S.R. Trans. Amer.
Philos. Soc., 52, pt. 2:1-224.
Rascoe, B., Jr.
1962. Regional stratigraphic analysis of Pennsylvanian and Permian rocks in
western midcontinent, Colorado, Kansas, Oklahoma, Texas. Bull. Amer.
Assoc. Petrol. Geol., 46:1 345-1370.
Romer, A. S.
1958. The Texas Permian redbeds and their vertebrate fauna. In Studies on
Fossil Vertebrates presented to David Meredith Seares Watson, Univ.
London, Athlone Press, pp. 157-179.
1960. The vertebrate fauna of the New Mexico Permian. New Mexico Geol.
Soc. Guidebook of Rio Chama Country, 11th Field Conf., pp. 48-54.
Romer, A. S., and E. C. Olson
1954. Aestivation in a Permian lungfish. Breviora, Mus. Comp. Zook, 30:1-8.
Romer, A. S., and L. I. Price
1940. Review of the Pelycosauria. Geol. Soc. Amer. Spec. Papers, 28:1-538.
1966
Early Permian Vertebrates
13
Vaughn, P. P.
1962. Vertebrates from the Halgaito tongue of the Cutler Formation, Permian
of San Juan County, Utah. J. Paleont., 36:529-539.
1963a. The age and locality of the late Paleozoic vertebrates from El Cobre
Canyon, Rio Arriba County, New Mexico. J. Paleont., 37:283-286.
1963b. A downslope trackway in the De Chelly Sandstone, Permian of Monu-
ment Valley. Plateau, 36:25-28.
1964a. Vertebrates from the Organ Rock Shale of the Cutler Group, Permian
of Monument Valley and vicinity, Utah and Arizona. J. Paleont., 38:
567-583.
1964b. Evidence of aestivating lungfish from the Sangre de Cristo Formation,
Lower Permian of northern New Mexico. Los Angeles County Mus.,
Cont. in Sci., 80: 1-8.
1965. Frog-like vertebrae from the Lower Permian of southeastern Utah. Los
Angeles County Mus., Cont. in Sci., 87:1-18.
1966. Seymouria from the Lower Permian of southeastern Utah, and possible
sexual dimorphism in that genus. J. Paleont., 40:603-612.
2
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»
LOS
ANGELES
COUNTY
MUSEUM
II
CONTRIBUTIONS
IN SCIENCE
Number 106
July 22, 1966
NEW DISTRIBUTION DATA FOR MARTAREGA, BUENO A AND
A BED US , INCLUDING THE FIRST RECORD OF THE GENUS
MARTAREGA IN THE UNITED STATES (HEMIPTERA:
NOTONECTIDAE, BELOSTOMATIDAE)
By Arnold S. Menke and Fred S. Truxal
Los Angeles County Museum of Natural History • Exposition Park
Los Angeles. California 90007
CONTRIBUTIONS IN SCIENCE is a series of miscellaneous technical papers
in the fields of Biology, Geology and Anthropology, published at irregular intervals
by the Los Angeles County Museum of Natural History. Issues are numbered sepa-
rately, and numbers run consecutively regardless of subject matter. Number 1 was
issued January 23, 1957. The series is available to scientific institutions on an ex-
change basis. Copies may also be purchased at a nominal price.
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or to papers dealing largely with specimens in the Museum’s collections. Manuscripts
must conform to CONTRIBUTIONS style and will be examined for suitability by
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David K. Caldwell
Editor
NEW DISTRIBUTION DATA FOR MARTAREGA, BVENOA AND
ABEDUS, INCLUDING THE FIRST RECORD OF THE GENUS
MARTAREGA IN THE UNITED STATES (HEMIPTERA:
NOTONECTIDAE, BELOSTOMATIDAE)
By Arnold S. Menke1 and Fred S. Truxal2
Abstract: Recent collections of aquatic Hemiptera reveal
significant range extensions for several poorly known species of
Notonectidae and Belostomatidae, including the first record for
the genus Martarega White in the United States. The first record
of flight in Abedus Stal is noted, as well as fragmentary notes on
the habits of Martarega. The taxonomy of Abedus signoreti is
discussed with a suggested change in the status of A. signoreti
vicinus.
Collections of aquatic Hemiptera made in Arizona, Mexico and Venezuela
during the last few years have helped to clarify the geographic ranges of several
poorly known species of Notonectidae and Belostomatidae. Most of the ma-
terial on which this report is based is located in the Los Angeles County Mu-
seum of Natural History (LACM), or in the A. S. Menke Collection (ASM)
which was recently acquired by the Los Angeles museum. Some specimens
from the collections of C. V. Reichart, Providence College, Rhode Island
(CVR); Snow Entomological Museum, University of Kansas (KU); and the
University of Michigan Museum of Zoology (UMMZ) also have been ex-
amined.
Family Notonectidae
Martarega White
Except for a brief mention of two new records from Brazil (Truxal, 1957 ) ,
nothing has been published since Truxal’s (1949) revision on this interesting
genus. Until recently, Martarega has been considered to be strictly a Neotropi-
cal group. The northernmost records for Martarega given by Truxal were from
central Mexico (M. mexicana Truxal). However, in 1958, A. S. Menke and
L. A. Stange collected a large series of M. mexicana in central Arizona, thus
establishing the fact that Martarega occurs much farther north. Since then,
F. S. Truxal, C. V. Reichart, and J. T. Polhemus3 have made additional collec-
tions of M. mexicana at various Arizona localities. All of these collections have
department of Entomology, University of California, Davis.
2Chief Curator of Life Sciences, Los Angeles County Museum of Natural History.
3After this paper was in press, a report by Polhemus was published concerning his
Arizona Martarega records. [Some Hemiptera New to the United States (Notonec-
tidae, Saldidae). Proc. Ent. Soc. Wash. 68 ( 1 ) : 57. March 1966].
1
2
Contributions in Science
No. 106
been made in either the Verde River or Salt River drainages in the Mogollon
Rim area of central Arizona. In view of the long history of collecting in Ari-
zona and other southwestern states with no previous evidence of Martarega, it
seems reasonable to assume that the genus has only recently invaded Arizona.
As is generally true for this genus, the great majority of specimens col-
lected in Arizona by Menke, Stange, Truxal and Reichart are brachypterous.
In fact, of the more than three hundred specimens collected, only three (by
Reichart) are macropterous.
Since nothing has been published on the habits of Martarega, we should
like to offer the following fragmentary notes. Several collectors have observed
Martarega jumping free of the water when their habitat is invaded by the col-
lector’s net, or when said habitat is otherwise endangered, as by an approach-
ing boat. A. S. Menke noted this behavior in M. chinai Hynes in Venezuela,
and C. V. Reichart observed the same behavior in M. mexicana in Arizona.
R. L. Usinger and F. S. Truxal have also noted this phenomenon for Marta-
rega in Peru and Brazil (respectively). The most obvious explanation for this
behavior is that it is a means of escaping enemies. Presumably Martarega serve
as food for fish and when endangered by such predators, one might assume
that they jump free of the water in an attempt to escape.
Martarega are stream inhabitants, and tend to be gregarious forming large
Figure 1. The East Verde River, seven miles north of Payson, Arizona (site of first
record for Martarega in the United States) typifies the stream habitat of the back-
swimmer genus Martarega.
1966
Records of Aquatic Hemiptera
3
schools in deep sheltered eddies. Like Buenoa, they appear to be in equilibrium
with the water and do not require active swimming movements or a clinging
to underwater vegetation to remain submerged. It has been proposed by Miller
(1964) that Buenoa and its Old World counterpart, Anisops, are able to main-
tain this neutral buoyancy in water because of the presence of large haemo-
globin-filled tracheal cells in their abdomens. Haemoglobin has not been re-
corded in Martarega or in any notonectids other than Buenoa and Anisops.
Martarega mexicana Truxal
UNITED STATES, ARIZONA, Gila Co.: East Verde River, 7 mi. N.
Payson, IX- 1 1-1958, 95 5, 68$ (A. S. Menke and L. A. Stange, LACM);
X-26-1959, 61 $ , 55$ (F. S. Truxal and L. Martin, LACM). Navajo Co.:
Carrizo Creek at Highway 60 (about 25 mi. S.W. Show Low), VII-3 1-1965,
46$ , 26 $ (C. V. Reichart, CVR, LACM).
MEXICO, Nayarit: Compostela, XII-30-1958, 24$, 19$ (A. S. Menke
and L. A. Stange, LACM). Vera Cruz: Tamazunchale, IV- 11- 1949, 5 5,4$
(F. S. Truxal, LACM).
Truxal (1949) records this species from the state of Morelos in Mexico,
and from Guatemala.
Martarega chinai Hynes
VENEZUELA: Cano Mariusa, Orinoco Delta (Orinoco River approxi-
mately 140 kms. N.E. Barrancas), VII-8-1958, 4 5,3$ (A. S. Menke, LACM).
42 kms. S.E. Maturin, Monagas, VII-3-1958, 1 $ (A. S. Menke, LACM).
Truxal (1949) records this species from Brazil and Bolivia.
Buenoa Kirkaldy
Buenoa hunger j or di Truxal
MEXICO, Jalisco : Santa Cruz Astillero, XII-30-1958 (A. S. Menke and
L. A. Stange, LACM). Puebla: Petlalcingo, VII-22-1959 (A. S. Menke and
L. A. Stange, LACM). Sonora: Cibuta, V- 18- 1954 (F. S. Truxal, LACM).
Truxal ( 1953) described this species from two widely separated areas : to
the south in the Mexican state of Chiapas, and to the north in Arizona and the
Mexican state of Sonora. The above records help to fill in this gap in the
known distribution of hungerfordi. Intensive collecting in Arizona has yielded
specimens of hungerfordi only from Sabino Canyon in Pima County, one of
the type localities.
Family Belostomatidae
Abedus Stal
Since Menke’s ( 1960) revision of this genus, the following new data are
worth noting. Especially interesting are the records of Abedus signoreti vicinus
4
Contributions in Science
No. 106
Mayr and A. signoreti sonorensis Menke taken at black light at night. These
are the first authentic records of flight in Abedus. Abedus signoreti is the most
Belostoma- like species in the genus Abedus, and is therefore, the species in
which one would expect to find the common Belostoma habit of flying to light.
Abedus signoreti Mayr
Menke (1960) recognized three subspecies of A. signoreti. Some of the
following records are interesting because they indicate a greater overlap in the
ranges of the subspecies than was indicated by Menke. In the zone of overlap
between A. signoreti vicinus Mayr and A. signoreti sonorensis Menke (the
state of Sinaloa), the characters which separate the two forms tend to break
down. However, character breakdown is not as common within the zone of
overlap of A. signoreti vicinus Mayr and A. signoreti s.s. suggesting that per-
haps A. vicinus should be elevated to specific status (with sonorensis as a sub-
species).
Abedus signoreti s.s.
MEXICO, Chiapas : Cintalapa, 28 mi. W., IV-9-1962 (F. D. Parker and
L. A. Stange, ASM). Villa Flores, 9 mi. N., VIII-12-1963 (F. D. Parker and
L. A. Stange, ASM). Nuevo Leon : El Alamo, IV-24-1956 (ASM). Monte-
morelos, IV- 16- 1956 (L. Martinez, ASM). San Luis Potosi : Ciudad de Valles,
VIII-8-1951 (C. J. Drake, ASM). Tamaulipas : Ciudad Victoria, XI-5-1936
(H. D. Thomas, KU, ASM) ; VIII-9- 1951 (C. J. Drake, ASM). Llera, VI-15-
1953 (ASM). Vera Cruz'. Alvarado, VII-28-1951 (C. J. Drake and F. C.
Hottes, ASM). Xico (Jico), 16 kms. S. W. Jalapa, IV-3-? (S. Meek, ASM).
EL SALVADOR: Los Chorros National Park, VIII-28-1961 (M. E.
Irwin, ASM). Quezaltepeque, 5 mi. N., VI-28-VIII-1961 (M. E. Irwin,
ASM); 2 mi. W., VIII-10-1961 (M. E. Irwin, ASM).
HONDURAS: Minas de Oro, Comayagua, 4000', IV-29-? (ASM). Guai-
maca, 10 mi. E. on Highway 3, Dept. Francisco Morazan, XI-6-1964 (J. S.
Packer, ASM).
Abedus signoreti vicinus Mayr
MEXICO: Chiapas : Comitan, VIII-30-1937 (H. D. Thomas, KU, ASM).
Jalisco : Plan de Barrancas, 3 mi. S.E., taken at black light, VII-8-1963 (F. D.
Parker and L. A. Stange, ASM). Morelos : Xochicalco Pyramid, III-29-1962
(F. D. Parker and L. A. Stange, ASM). Yautepec, taken at black light, III-26-
1962 (F. D. Parker and L. A. Stange, ASM); at black light, VII-13-1963
(F. D. Parker and L. A. Stange, ASM) . Puebla : Izucar de Matamoros, at black
light, VIII- 1-1 963 (F. D. Parker and L. A. Stange, ASM). Sinaloa'. Chupa-
deros, Highway 40, III- 19- 1962 (F. D. Parker and L. A. Stange, ASM).
1966
Records of Aquatic Hemiptera
5
The Chiapas record extends the known range of A . vicinus much farther
south and well into the range of A. signor eti s.s. The specimens are typical
A. vicinus.
Abedus signoreti sonorensis Menke
MEXICO, Sinaloa : Elota, 8 mi. S., at black light, VIII-26-1963 (F. D.
Parker and L. A. Stange, ASM). Sonora : Alamos, IX- 1-1 960 (R. L. Westcott,
ASM); 10 mi. S. E., V-22-1962 (F. D. Parker and L. A. Stange, ASM). La
Aduana, VI- 12- 1961 (A. S. Menke, ASM).
One female from the series collected at Elota, Sinaloa, has a partially
formed embolial fracture. It is interesting to note that Abedus signoreti vicinus
is characterized by the presence of this fracture, whereas A. sonorensis lacks it.
Abedus ovatus Stal
MEXICO, Morelos’. Cuernavaca, 5 mi. E., III-28-1962 (F. D. Parker and
L. A. Stange, ASM).
This is the first record of A . ovatus from Morelos, Mexico.
Abedus breviceps Stal
MEXICO, Guerrero : Iguala, 45 mi. S.W., 1-8-1956 (ASM). Mexico:
Sabinas, V-6-1962 (F. D. Parker and L. A. Stange, ASM). Vera Cruz: Orizaba,
Rio Blanca (A. J. Woolman, ASM).
The Vera Cruz record is the first for this Mexican state.
Abedus immensus Menke
MEXICO, Aguascalientes: Sierra Fria, 40 mi. S.W. Rincon de Romos,
8200', III-9-1953 (I. J. Cantrall, ASM). Durango: San Luis, 2.5 mi. W.,
8000', III-24-1953 (I. J. Cantrall, ASM).
This species appears to be restricted to the western part of the central
plateau of northern Mexico.
Abedus stangei Menke
MEXICO, Michoacan: Dos Aguas (lumber camp 22 mi. N.W. Agualilla
on Apazingan-Agualilla road), 6900 ft., VI-18-1958 (M. Mendoza, UMMZ
and ASM). Puebla: Sierra de Zacapoaxtla (ASM).
Previous records of this species were from the Mexican states of Vera
Cruz and Puebla (Menke, 1960). The Michoacan record cited here extends
the known range of A. stangei considerably westward. Like A. immensus , this
species evidently is found only at higher elevations (4000 feet and above).
6
Contributions in Science
No. 106
Literature Cited
Menke, A. S.
1960. A taxonomic study of the genus Abedus Stal. Univ. Calif. Publ. Ento-
mol., 16:393-440.
Miller, P. L.
1964. The possible role of haemoglobin in Anisops and Buenoa (Hemiptera:
Notonectidae). Proc. Royal Entomol. Soc. London, 39:166-175.
Truxal, F. S.
1949. A study of the genus Martarega. J. Kansas Entomol. Soc., 22: 1-24.
1953. A revision of the genus Buenoa. Univ. Kansas Sci. Bull., 35:1351-1523.
1957. The Machris Brazilian Expedition, Systematics of the Notonectidae. Los
Angeles County Mus., Cont. in Sci., 12:1-23.
II
LOS
ANGELES
COUNTY
MUSEUM
CONTRIBUTIONS
IN SCIENCE
Dumber 107
July 22, 1966
TWO FOSSIL BIRDS FROM THE LOWER MIOCENE
OF SOUTH DAKOTA
By Hildegarde Howard
Los Angeles County Museum of Natural History • Exposition Park
Los Angeles, California 90007
CONTRIBUTIONS IN SCIENCE is a series of miscellaneous technical papers
in the fields of Biology, Geology and Anthropology, published at irregular intervals
by the Los Angeles County Museum of Natural History. Issues are numbered sepa-
rately, and numbers run consecutively regardless of subject matter. Number 1 was
issued January 23, 1957. The series is available to scientific institutions on an ex-
change basis. Copies may also be purchased at a nominal price.
INSTRUCTIONS FOR AUTHORS
Manuscripts for the LOS ANGELES COUNTY MUSEUM CONTRIBU-
TIONS IN SCIENCE may be in any field of Life or Earth Sciences. Acceptance of
papers will be determined by the amount and character of new information and the
form in which it is presented. Priority will be given to manuscripts by staff members,
or to papers dealing largely with specimens in the Museum’s collections. Manuscripts
must conform to CONTRIBUTIONS style and will be examined for suitability by
an Editorial Committee. They may also be subject to critical review by competent
specialists.
MANUSCRIPT FORM.— (1) The 1960 AIBS Style Manual for Biological
Journals is highly recommended as a guide. (2) Typewrite material, using double
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of literature citation must conform to CONTRIBUTIONS style— see number 90 and
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Orders for additional copies should be sent to the Editor at the time corrected galley
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David K. Caldwell
Editor
TWO FOSSIL BIRDS FROM THE LOWER MIOCENE
OF SOUTH DAKOTA
By Hildegarde Howard1
Abstract: A new genus and species of raptor (Order
Falconiformes) and a new species of quail (Order Galliformes)
are described from the Sharp’s Formation of Shannon County,
South Dakota.
Since Macdonald’s (1963) significant report on the Miocene vertebrates
from the Wounded Knee area of Shannon County, South Dakota, he has
continued field work in the region with parties from the Los Angeles County
Museum of Natural History (LACM). Among the specimens collected in
1964 in the Sharp’s Formation, are two fragments of bird bones, one represent-
ing the Order Falconiformes (diurnal raptors) the other the Galliformes
(fowl-like birds). These are the first avian remains to be discovered in this
formation, from which Macdonald (1963:151-153) records 61 mammals
and 4 reptiles.
Macdonald places the Sharp’s Formation and its fauna at the bottom of
the Lower Miocene Arikaree group, and at an earlier stage in the Miocene than
the avifauna recorded by Miller (1944) from Flint Hill, Bennett County,
South Dakota.
Order Falconiformes
Family Accipitridae
Subfamily Aegypiinae
The falconiform bone is a well-preserved distal end of tibiotarsus, which
in size is comparable to this element of the Red-tailed Hawk, Buteo borealis.
But the shorter, stouter, more horizontally-placed supratendinal bridge indi-
cates not only generic, but subfamily distinction. Closest resemblance of the
South Dakota fossil is to tibiotarsi of Neogyps errans Miller and Palaeoborus
umbrosus (Cope), North American fossil members of the Old World Vulture
subfamily (Aegypiinae), in which characters are more eagle-like than in living
representatives of the group. The South Dakota tibiotarsus also displays aegy-
piine and eagle-like characters, but is sufficiently distinct from tibiotarsi of the
previously described forms to warrant establishing a new genus.
In the description to follow, comparisons are made with tibiotarsi of
Neogyps errans from the Pleistocene of Rancho La Brea, California, in the
collections of the Los Angeles County Museum of Natural History, and with
the description and illustrations of the tibiotarsus of Palaeoborus umbrosus
1Research Associate in Vertebrate Paleontology, Los Angeles County Museum of
Natural History.
1
;i HSON 1 A fii b.j'* c\
§W<nTrjmmi • mm
2
Contributions in Science
No. 107
from the Pliocene of New Mexico, as presented by Cope (1877:293-294, and
pi. 68, fig. 18).
Arikarornis, new genus
Type species: Arikarornis macdonaldi.
Diagnosis: Tibiotarsus with supratendinal bridge short and broad, thick-
ened and distinctly convex on distalmost edge; tendinal groove above bridge
deeply cut and centrally placed, with internal attachment for oblique ligament
on its sloping, internal face, well above bridge, and shaft external to groove
smoothly rounded; lateral flare from shaft to condyles very gradual, and in-
ternal condyle with only slightly more lateral thrust than external; condyles
nearly equal in anteroposterior depth, projecting at abrupt right angle from
shaft anteriorly, and having well-defined parallel borders posteriorly, with only
slight trend mediad above level of proximo-anterior border; anterior inter-
condylar fossa broad, evenly rounded, only slightly rugose, with no marked
undercutting of median borders of condyles; distal contour broad and shallow;
internal ligamental prominence a well-rounded distinct papilla, approximately
centrally located with respect to anteroposterior and proximodistal borders of
internal condyle.
Arikarornis macdonaldi, new species
Figure 1, A-D
Type: Distal end of left tibiotarsus, LACM no. 9357, collected by J. R.
Macdonald field party, June 19, 1964.
Locality and horizon: LACM loc. no. 1821 (equivalent of South Dakota
School of Mines loc. no. 5359 as recorded by Macdonald, 1963), gully on
south side of Sharp’s Cutoff Road, SW14 of Sect. 9, T. 39 N., R. 43 W.,
Sharp’s Corner Quadrangle, Shannon County, South Dakota (Pine Ridge
Reservation). Middle Sharp’s Formation, Arikaree group, lowermost Miocene.
Figure 1. A-D, Arikarornis macdonaldi n. gen., n. sp., type tibiotarsus, anterior, ex-
ternal, internal and posterior views; E. Miortyx aldeni, n. sp., type humerus, anconal
view. All figs, x 1.
1966
New Fossil Birds
3
Diagnosis: See generic diagnosis.
Comparisons of type tibiotarsus: Similar to this element in Recent genera
of Aegypiinae in short, broad supratendinal bridge, less vertical in position
than in Aquila or Buteo, and tendinal groove centrally placed on shaft above
bridge, with attachment of oblique ligament on slope of internal face of groove,
and anterior face of shaft external to groove well rounded; the short, broad,
less vertically placed supratendinal bridge is also characteristic of the tibiotar-
sus of Palaeoborus and Neogyps, and the tendinal groove is centrally placed in
Palaeoborus (slightly more lateral in Neogyps). Distinguished from Recent
Aegypiinae, and similar to Palaeoborus umbrosus and Neogyps errans in rela-
tively broad shaft and relatively short anteroposterior depth of condyles
(shorter, even, than in Palaeoborus) . Distinguished from both P. umbrosus
and N. errans in more nearly equal depth of internal and external condyles,
and more gradual lateral flare from shaft to condyles; further distinguished
from Palaeoborus in more central position of internal ligamental prominence
with respect to borders of internal condyle, and more parallel posterior bor-
ders of external and internal condyles; Cope (1877:293-294) described the
contours of the internal condyle of Palaeoborus umbrosus as “not parallel to
the exterior, but diverging backward and inward;” Neogyps is closer to Ari-
karornis in posterior contours of the condyles, but the postero-internal con-
tour in the Pleistocene form slopes much more abruptly mediad at the level of
the proximo-anterior border.
Measurements: See Table 1 .
Remarks: The North American record of the Aegypiinae comprises the
following eight species (the tibiotarsus is known only in those species marked
with an asterisk) :
Neophrontops vetustus Wetmore, Middle Miocene, Nebraska
Neophrontops dakotensis Compton, Lower and Middle Pliocene, South
Dakota and Oregon
Neophrontops vallecitoensis Howard, Middle Pleistocene, California
* Neophrontops americanus L. Miller, Upper Pleistocene, California and
Mexico
Palaeoborus rosatus A. Miller and Compton, Lower Miocene, South Da-
kota
Palaeoborus howardae Wetmore, Middle Miocene, Nebraska
* Palaeoborus umbrosus (Cope), Lower Pliocene, New Mexico
*Neogyps errans L. Miller, Upper Pleistocene, California, Nevada, and
Mexico
The skeleton of Neophrontops is markedly like that of the Recent Old
World Vulture, Neophron (see Howard, 1932) and the tibiotarsus is distinctly
different from that of Arikarornis. In its small size, however, A. macdonaldi
4
Contributions in Science
No. 107
TABLE 1
Measurements and Proportions of Tibiotarsus of
Arikarornis macdonaldi, Palaeoborus umbrosus and Neogyps errans
(Measurements in millimeters, ratios in per cent)
Arikarornis
Palaeoborus
Neogyps
Breadth of
distal end
13.2
16.0
18.0
19.4
20.8
Depth of
external condyle
9.2
12.0
11.8
12.7
13.4
Depth of
internal condyle
9.3
13.0
12.8
13.7
14.5
Ratio of depth of external
condyle to breadth of
distal end
69.6
75.0
61.5
66.0
68.3
Ratio of depth of internal
condyle to breadth of
distal end
70.6
81.3
68.0
71.0
72.5
Ratio of depth of external to
depth of internal condyle
99.1
92.4
90.0
93.6
96.3
is closer to all species of Neophrontops than to any of the other fossil aegy-
piines.
The comparisons given above show Arikarornis to have similarities with
Neogyps and Palaeoborus, but to be distinct from N. errans, the monotypic
species of Neogyps, and from P. umbrosus, the genotype of Palaeoborus. The
tibiotarsus is not known for P. howardae or P. rosatus, but the tarsometatarsus
and ulna (respectively) on which these species are based are close in size to
these elements of Neogyps errans, and therefore indicate that both species were
larger even than Palaeoborus umbrosus, hence considerably larger than Ari-
karornis macdonaldi.
The extinct Palaeohierax from the early Miocene of France (based on the
tarsometatarsus) is said by Milne-Edwards (1871, 2:456-457) to combine
characters of Gypohierax and the eagles (“Aquilides”). The genus is now
listed under tht Aegypiinae (Brodkorb, 1964:275). Lacking comparable skele-
tal elements of Palaeohierax and Arikarornis, it is impossible to draw any
conclusions as to the relationship of these two aberrant forms, other than to
state that the tarsal breadth in the single species of Palaeohierax (P. gervaisii)
indicates a much larger form than A . macdonaldi. In the present state of knowl-
edge, science is better served by maintaining generic as well as specific identity
of these birds.
The generic name, Arikarornis refers to the Arikaree group of the Lower
Miocene, in which the Sharp’s fauna occurs. The species is named in honor of
1966
New Fossil Birds
5
J. R. Macdonald whose studies have so significantly furthered the knowledge
of the Miocene of South Dakota.
Order Galliformes
Family Phasianidae
Subfamily Odontophorinae
The galliform bone is a fragment of left humerus characterized by very
deep undercutting of the head. In living galliforms this is an outstanding fea-
ture of the American quails (excepting Dendrortyx and Odontophorus accord-
ing to Holman, 1961:208). In the living quails, however, the head terminates
squarely and abruptly at the capital groove. The fossil at hand has a rounded
contour of the head as described for Miortyx teres Miller (1944:93), and it is,
therefore, assigned to the genus Miortyx. The genotypic Miortyx teres is based
on a proximal end of humerus found in the Flint Hill quarry, Bennett County,
South Dakota, of Miocene age, but younger than the Sharp’s Formation. This
specimen was lent by the University of California Museum of Paleontology
for this study.
That the Sharp’s Formation specimen cannot be allocated to Miortyx teres
is obviously attested by its markedly larger size, as well as certain qualitative
features. A distinct species is therefore established in honor of Dr. Alden H.
Miller, describer of the genus, whose untimely death, in 1965, has deprived
paleornithology of one of its ablest contributors.
In the original description, the characters of the genus Miortyx were not
separated from those of the type species. This second species makes possible
the designation of characteristics at the generic level.
Miortyx, A. H. Miller
Diagnosis (proximal end of humerus) : Anconal side of shaft below head
broadly depressed, and head deeply undercut as in Oreortyx ; descending, lip-
like anconal border of head above median crest shorter and broader than in
Oreortyx, and head less abruptly terminated internally above capital groove,
with anteroposterior depression (which in Oreortyx faces directly internally)
facing proximo-internally on the gradually rounded internal contour of the
head; capital groove well defined, with borders nearly parallel, terminating
anconally at median crest in distinct open lip; pneumatic fossa long and oval,
and anconally less markedly obscured by overhang of internal tuberosity than
in Oreortyx’, ligamental furrow on palmar surface deeply grooved.
Miortyx aldeni, new species
Figure 1, E
Type: Proximal fragment of left humerus lacking external and internal
tuberosities and deltoid crest: LACM no. 9388, collected by H. Garbani of
J. R. Macdonald field party, June 23, 1964.
6
Contributions in Science
No. 107
Locality and horizon: LACM loc. no. 1982 (equivalent of South Dakota
School of Mines loc. 5360 as recorded by Macdonald, 1963), gully beside
Sharp’s Cutoff Road, N. Vi of Sect. 17, T. 39 N., R. 43 W., Sharp’s Corner
Quadrangle, Shannon County, South Dakota (Pine Ridge Reservation). Mid-
dle of Sharp’s Formation, Arikaree group, lowermost Miocene.
Diagnosis: Humerus approximately 50 per cent larger than that of Miortyx
teres and differing also in the following qualitative characters: prominent de-
scending median border of head more anconally projected, with undercutting
of head deepened in this area; shaft below head, anconally, more broadly and
evenly depressed; external bordering ridge of depression extending at least to
level of distal terminus of pneumatic fossa (bone broken beyond this point).
Measurements: See Table 2.
Remarks: According to Brodkorb’s (1964:309-311) recent analysis of
previously described fossil Galliformes, seven extinct species of Odonto-
phorinae are recognized, as follows:
Nanortyx inexpectatus Weigel, Lower Oligocene, Saskatchewan
Miortyx teres Miller, Lower Miocene, South Dakota
Cyrtonyx cooki Wetmore, Middle Miocene, Nebraska
Lophortyx shotwelli Brodkorb, Middle Pliocene, Oregon
Colinus hibbardi Wetmore, Upper Pliocene, Kansas
Colinus suilium Brodkorb, Middle Pleistocene, Florida
Neortyx peninsularis Holman, Middle Pleistocene, Florida
TABLE 2
Measurements (in millimeters) of Humerus of
Miortyx aldeni and Miortyx teres
M. aldeni
M. teres
Breadth across proximal end from greatest extent
of bicipital crest to probable border of external
tuberosity (tuberosity broken in M. aldeni )
19.1
12.6
Breadth of depressed area of shaft from median
crest at terminus of capital groove, to ridge
bordering external edge of depression
10.0
5.4
Depth of head external to descending median border
7.6
4.8
Height of head from tip of median
border to proximal end
9.1
6.3
Comparison of the elements represented in each species with comparable
elements in the skeleton of Recent Oreortyx picta indicates that Miortyx aldeni
was outstandingly the largest of the fossil quails, with the nearest approach in
size being Miortyx teres. Besides the two species of Miortyx , only one other
quail is recognized from the Miocene. The Barstow, California Miocene spe-
1966
New Fossil Birds
7
cies described as Cyrtonyx tedfordi L. Miller, is now reallocated to the Cracidae
under the generic name Boreortalis (Brodkorb, 1964:305); this species, also,
is much smaller than M. aldeni.
European Tertiary galliforms, originally described under the genus Palae-
ortyx, in which the head of the humerus is deeply undercut as in American
quails, differ from Miortyx in deeper, more acute, and more obliquely placed
depression of the shaft anconally, and longer, narrower descending median
border of the head. As presently listed by Brodkorb (1964:298-301) the sev-
eral species involved appear under the genera Palaeortyx, Ludiortyx, Pirortyx
and Taoperdix, and are allocated to the primitive subfamily Gallinuloidinae of
the family Cracidae. According to recent personal correspondence with Brod-
korb, these European galliforms are badly in need of revision, but present
interpretation is based on the primitive condition of the carpometacarpus
which lacks the intermetacarpal tuberosity. The carpometacarpus is not known
for either species of Miortyx.
Summary and Conclusions
The first avian representation from the Lower Miocene Sharp’s Formation
of South Dakota is recorded, and two species are described: a raptor, Arika-
rornis macdonaldi, and a quail, Miortyx aldeni. Both are in ecologic agreement
with the general terrestrial aspect of the mammal and reptile fauna recorded
by Macdonald ( 1963 : 1 5 1 ) , but contribute no independent information in this
regard. Miller ( 1944:97), describing Miortyx teres, the genotype of the quail,
notes that it “is not closely enough linked with any one of the modern types to
offer a clue to its habitat!’ The same can also be said of the raptor, Arikarornis.
The scant avian representation in the Sharp’s fauna makes impossible any
critical comparison with other Miocene avifaunas. It is significant, however, to
find generic relationship between the quail, Miortyx aldeni and Miortyx teres
of the slightly later Flint Hill Miocene fauna of Bennett County, South Da-
kota (about 34 miles ESE of Sharp’s Corner). There is also a possibility of
generic relationship between Arikarornis macdonaldi and the Flint Hill aegy-
piine, Palaeoborus rosatus. Miller and Compton (1939: 156) in describing the
latter species, stated that the generic assignment was “by no means certain!’
Direct comparison of the type element (ulna) could not be made with pre-
viously described species of Palaeoborus, but parallel similarities with Pleisto-
cene Neogyps were noted. Resemblance to Neogyps is noted as well for Ari-
karornis (as described above). No parallel can be drawn between the quail,
and the raptor with regard to size trend from earliest Miocene, Sharp’s fauna,
to that of the somewhat later Miocene, Flint Hill fauna. The quail, Miortyx
aldeni, is markedly larger than M. teres (in fact the largest of the American
quails), whereas Arikarornis macdonaldi is one of the smallest of the fossil
Aegypiinae, and probably less than half the size of Palaeoborus rosatus.
8
Contributions in Science
No. 107
Acknowledgments
I am indebted to Dr. J. R. Macdonald, Senior Curator of Vertebrate
Paleontology, Los Angeles County Museum of Natural History, for the oppor-
tunity to study the avian material from the Sharp’s fauna; and to Dr. D. E.
Savage, of the University of California Museum of Paleontology, for the loan
of the type specimen of Mioriyx teres. The photographs were made by Mike
Hatchimonji, staff photographer of the Los Angeles County Museum of Na-
tural History.
Literature Cited
Brodkorb, Pierce
1964. Catalogue of fossil birds, part 2 (Anseriformes through Galliformes).
Bull. Florida State Mus., Biol. Sci., 8(3) : 195-335.
Cope, E. D.
1877. Report on the extinct Vertebrata obtained in New Mexico by parties of
the expedition of 1874. Geog. Surv. west of 100th meridian, by Geo. M.
Wheeler, vol. 4, Paleontology, 1-270, pis. 22-83.
Holman, J. Alan
1961. Osteology of living and fossil new world quails (Aves, Galliformes).
Bull. Florida State Mus., Biol. Sci., 6(2) : 13 1-232.
Howard, Hildegarde
1932. Eagles and eagle-like vultures of the Pleistocene of Rancho La Brea.
Carnegie Inst. Washington, Publ. 429:1-82.
Macdonald, James Reid
1963. The Miocene faunas from the Wounded Knee area of western South
Dakota. Bull. Amer. Mus. Nat. Hist., 125(3) : 141-238.
Miller, Alden H.
1944. An avifauna from the lower Miocene of South Dakota. Univ. Califor-
nia Publ. Bull. Dept. Geol. Sci., 27(4) :85-100.
Miller, Alden H. and Lawrence V. Compton
1939. Two fossil birds from the lower Miocene of South Dakota. Condor,
41(4) : 153-156.
Milne-Edwards, Alphonse
1871. Recherches anatomiques et paleontologiques pour servir a l’histoire des
oiseaux fossiles de la France. Paris: G. Masson, 2:1-632, with atlas.
LOS
ANGELES
COUNTY
MUSEUM
CONTRIBUTIONS
IN SCIENCE
Number 108
July 25, 1966
i Sol- 7 3 ,
(2, L. %>(& >
f
SOUNDS AND BEHAVIOR OF CAPTIVE AMAZON
FRESHWATER DOLPHINS, 1NIA GEOFFRENSIS
By Melba C. Caldwell, David K. Caldwell
and William E. Evans
II
i
Los Angeles County Museum of Natural History • Exposition Park
Los Angeles, California 90007
CONTRIBUTIONS IN SCIENCE is a series of miscellaneous technical papers
in the fields of Biology, Geology and Anthropology, published at irregular intervals
by the Los Angeles County Museum of Natural History, Issues are numbered sepa-
rately, and numbers run consecutively regardless of subject matter. Number 1 was
issued January 23, 1957. The series is available to scientific institutions on an ex-
change basis. Copies may also be purchased at a nominal price.
INSTRUCTIONS FOR AUTHORS
Manuscripts for the LOS ANGELES COUNTY MUSEUM CONTRIBU-
TIONS IN SCIENCE may be in any field of Life or Earth Sciences. Acceptance of
papers will be determined by the amount and character of new information and the
form in which it is presented. Priority will be given to manuscripts by staff members,
or to papers dealing largely with specimens in the Museum’s collections. Manuscripts
must conform to CONTRIBUTIONS style and will be examined for suitability by
an Editorial Committee. They may also be subject to critical review by competent
specialists.
MANUSCRIPT FORM.— (1) The 1960 AIBS Style Manual for Biological
Journals is highly recommended as a guide. (2) Typewrite material, using double
spacing throughout and leaving ample margins, on only one side of 8Vi x 11 inch
standard weight paper. (3) Place tables on separate pages. (4) Footnotes should be
avoided if possible. (5) Legends for figures and unavoidable footnotes should be
typed on separate sheets. Several of one kind may be placed on a sheet. (6) Method
of literature citation must conform to CONTRIBUTIONS style — see number 90 and
later issues. Spell out in full the title of non-English serials and places of publication.
(7) A factual summary is recommended for longer papers. (8) A brief abstract must
be included for all papers. This will be published at the head of each paper.
ILLUSTRATIONS. — All illustrations, including maps and photographs, should
be referred to as “figures!’ All illustrations should be of sufficient clarity and in the
proper proportions for reduction to CONTRIBUTIONS page size. Permanent ink
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photographs should be glossy prints of good contrast. Original illustrations will not
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PROOF. — Author will be sent galley proof which should be corrected and re-
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David K. Caldwell
Editor
SOUNDS AND BEHAVIOR OF CAPTIVE AMAZON
FRESHWATER DOLPHINS, INIA GEOFFRENSIS
By Melba C. Caldwell,1 David K. Caldwell2
and William E. Evans3
Abstract: Twelve types of phonations, placed in four major
categories, were recorded in 688 minutes of listening to eight cap-
tive Inia geoffrensis (Blainville) . These sounds are discussed and
sonograms of typical ones are presented. Both juveniles and
adults were studied under a variety of circumstances. In general,
the phonations are less varied, lower in intensity, and of slightly
lower frequencies than those observed in most other odontocete
cetaceans. Included among the phonations are click trains which
when correlated with observed behavior suggest an ability by this
species to echolocate. However, the use of this ability may be de-
pendent on learning. Evidence is presented to indicate that vision
is the preferred method of environmental exploration, but some
tactile sense may also be employed. Data are included to indicate
for Inia frequent and precocious sexual play, a general lack of
competitive feeding behavior, and a lower incidence of fear re-
sponses than demonstrated by the much studied Atlantic bottle-
nosed dolphin.
Introduction
Dolphins of the family Platanistidae are considered the most primitive of
the living odontocete cetaceans (Simpson, 1945: 100). For the purposes of
comparison with certain of the more advanced dolphins, of the family
Delphinidae, we were especially interested in learning something of the
phonations, and more particularly of possible echolocation ability, in the
Platanistidae. To our knowledge, only one of the four species of platanistids,
the Amazon freshwater dolphin, Inia geoffrensis (Blainville), is available
presently for study in the United States. We recorded the phonations along
with observed concurrent captive behavior of eight animals and the behavior
of two others was observed but no recordings were attempted. Other behaviors,
not necessarily related to sound production, were also studied. Amazonian
animals, one each held captive at the Toledo Zoo, Ohio (see Hofmeister, 1964) ,
and at the John G. Shedd Aquarium, Chicago, Illinois, were not studied
directly, but enough was learned of their behavior (from Max Hofmeister at
Toledo; and from William P. Braker at Chicago) to indicate that it did not
Research Associate, Los Angeles County Museum of Natural History; also Staff Re-
search Associate, Allan Hancock Foundation, University of Southern California.
2Curator of Ichthyology and Marine Mammals, Los Angeles County Museum of
Natural History; also Research Associate, Florida State Museum, and Collaborator
in Ichthyology, Institute of Jamaica.
3Research Associate, Los Angeles County Museum of Natural History; also Re-
search Marine Zoologist at the Marine Biology Facility, Point Mugu, California, for
the United States Naval Ordnance Test Station, China Lake, California.
1
2
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differ significantly from that of the animals we did study. The behavior and
sonic display of two Amazonian males held captive at Silver Springs, Florida,
was discussed by Layne and Caldwell (1964), Schevill and Watkins (1962),
Layne (1959), Allen and Neill (1957) and Phillips (1964: 95 ff.). As this
paper goes to press we have also learned that an Amazonian Inia was kept for
a short time at the Crandon Park Zoological Garden, Miami, Florida (Gordon
Hubbell, pers. comm.). From time to time other Inia have been and are held
captive at the compounds of animal importers, mostly in Florida. Most notable
of these is the Tarpon Zoo at Tarpon Springs, where, through the courtesy of
Fred Penman, individuals were observed from time to time. We have had no
reports from various observers of behavior by these miscellaneous animals not
duplicated in our own observations on the animals listed below.
The species Inia geoffrensis is found in the Amazon and Orinoco rivers of
South America and their tributaries and adjacent lakes. During times of flood,
the animals may also be found throughout the flooded forest floors and may
remain in the lakes near the rivers after the floods subside even though in some
cases a connection to the rivers no longer remains. After capture, animals to be
imported into the United States are usually held in South America for varying
lengths of time until it is determined that they are in good health and that they
will feed in captivity. The period of time that they are held in South America
varies, but usually it is at least a week and sometimes as much as several months
before they are flown to the United States. Frequently the importer does not
know the exact length of time that the animals have been held. Consequently,
even if they are observed at the moment of their arrival in the United States,
they cannot truly be called naive as they have become adjusted to captivity
to some degree, to the eating of dead fish and to the presence of humans.
None of the animals we studied had been subjected to any known rein-
forcement of vocalizations, although (as noted below) all had been trained
to take food from a human hand and some had been subjected to more
complicated training procedures.
Acknowledgments
Access to the captive animals was made through the generous cooperation
of a number of people in charge, as indicated in the list of study sites included
below. We wish to express our sincere appreciation to all of these people,
most of whom gave us considerable information on the captive history and
behavior of the animals in their care, and some of whom gave much of their
own after-hours time in making our studies more profitable. Marie Poland
Fish, William H. Mowbray and Paul Perkins of the Narragansett Marine
Laboratory, and William E. Schevill and William A. Watkins of the Woods
Hole Oceanographic Institution kindly gave us copies of recordings they had
made, independently, of captive Inia and they both also made many helpful
comments and suggestions on a late version of our manuscript. Financial
1966
Amazon Dolphin Behavior
3
support for certain phases of the work came from the National Science
Foundation (grant no. GB-1189), the National Institute of Mental Health
(grant no. MH-07509-01 ), the American Philosophical Society (grant no.
3755-Penrose), and the Museum Associates of the Los Angeles County
Museum of Natural History. Technical support for certain parts of the study
came from the Naval Ordnance Test Station, China Lake, California. William
E. Sutherland of the Lockheed-California Company, Los Angeles, provided
helpful technical advice. The photographs of the live animal are by Fred
Jenne and are used here through the courtesy of Earl S. Herald, Steinhart
Aquarium. Photographs of the sonagrams are by Armando Solis and Mike
Hatchimonji, Los Angeles County Museum of Natural History.
Study Sites and History and Description
of Animals Studied
1. One juvenile male (47.5 inches, 121 cm., in snout to caudal-notch length
on first recording session; 52.5 inches, 133 cm., in snout to caudal-notch
length on second recording session). Recorded and observed through the
courtesy of Earl S. Herald, Robert P. Dempster and Thomas Green at the
Steinhart Aquarium, California Academy of Sciences, San Francisco, Cali-
Figure 1. Inia geoffrensis. Juvenile male (“Whiskers”) from the Amazon River drain-
age near Iquitos, Peru, at the Steinhart Aquarium in late 1964.
4
Contributions in Science
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fornia, on 30 September 1964 (six days after its arrival there) and on 5 May
1965. This animal was also observed on several intervening and subsequent
occasions when the recording of phonations was not attempted. The animal
was originally captured in the Amazon drainage near Iquitos, Peru, and was
held in Florida for some six weeks before it was received in San Francisco.
On our first recording session, it had not been subjected to any training other
than to take dead food fish from the hand. By the second recording session
this training had been supplemented with exposure to a large and a small
ball and a small hoop. Recordings were also made in April, 1965, by Dr.
Fish and her associates. Some historical and behavioral observations on this
animal (Figs. 1 and 10) were presented by Herald and Dempster (1965),
Dempster (1965), Richardson (1965), and Caldwell, Caldwell and Evans
{In press ) .
2. One juvenile male (49.2 inches, 125 cm., in snout to caudal-notch length).
Recorded and observed through the courtesy of John A. Moore at the Monte
Vista Zoological Park, Bloomington, California, on 12 September 1964, after
being there for at least six weeks and possibly for as long as two months. The
animal was originally captured in the Amazon drainage near Iquitos, Peru,
and was flown directly to California. It had not been subjected to training
other than to take dead food fish from the hand.
3. Two subadult males (about 69 and 73 inches, 175 and 185 cm., in snout
to caudal-notch length). Recorded and observed through the courtesy of
Kent Burgess, David W. Kenney and Donald D. Zumwalt at Sea World, San
Diego, California, on 17 April 1965 (10 days after their arrival there) and
on 18 and 19 February 1966. During the first session we had a flat (± 2 db)
recording capability of 40 to 20,000 cycles per second; and during the second
(two-day) session this was increased to a flat response (±2 db) of 110,000
cps, with a useable response of 150,000 cps. The animals were originally
captured in the Amazon drainage near Iquitos, Peru, and were flown directly
to California. They had not been subjected to training other than to take
dead food fish from the hand, but on occasion they had been allowed to
play with small objects placed in their tanks. Recordings were also made in
April, 1965, by Fish, Mowbray and Perkins from the Narragansett Marine
Laboratory. Some preliminary results of our studies with these animals have
been described by Caldwell, Caldwell and Evans {In press).
4. Two subadult to adult males (69 and 76.5 inches, 175 and 194 cm., in
snout to caudal-notch length). Recorded and observed for 70 minutes through
the courtesy of Winfield H. Brady at the Aquarium of Niagara Falls, New
York, on 8 April 1966 after being in captivity for approximately five months.
The animals were originally captured in the Amazon river about 60 miles
from Manaos, Brazil, and were flown directly to Niagara Falls. They had not
been subjected to training other than to take dead food fish from the hand.
1966
Amazon Dolphin Behavior
5
These animals were contained in a tank with a 49.3-inch (125-cm.) male
Sotalia sp. from near Manaos. Consequently, inasmuch as we cannot be sure
which animals produced which sounds, this 70-minute listening period is not
included in Table 1. However, no sounds were recorded, which we suspected
originated from the Ini a, that we had not recorded elsewhere under uncon-
taminated conditions. Schevill and Watkins had recorded these same three
animals under these conditions about one week previous to our visit, with
similar results. The behavior of the two Ini a at Niagara Falls was similar to
that observed for captive Ini a elsewhere.
5. One adult male (about 85 inches, 216 cm., in snout to caudal-notch
length) and one adult female (about 75 inches, 191 cm., in snout to caudal-
notch length). Recorded and observed through the courtesy of Lawrence
Curtis and Gary T. Hill at the James R. Record Aquarium, Fort Worth
Zoological Park, Fort Worth, Texas, on 26 June 1965. The male had been
captive there for 34 months and the female for 37 months, and both had
been held in Florida for an unknown period of time prior to their arrival
at Fort Worth. The animals originally were captured in the Amazon near
Leticia, Colombia. They had been subjected to simple training procedures,
which included taking dead food fish from the hand, jumping clear of the
water in a vertical manner for food, and jumping and grasping a ball in order
to raise a flag. Most of the activity other than simple feeding was performed
by the male. Some historical and behavioral data on these animals were
presented by Curtis (1962), Walker (1964: 1089), Phillips and McCain
(1964), Hill (1965), and Caldwell, Caldwell and Evans (In press).
6. One adult and one juvenile of undetermined sex and size (very approx-
imately, about 60 and 70 inches, 152 and 178 cm., in snout to caudal-notch
length) were observed but no recordings were attempted on 28 June 1964, at
Homosassa Springs, Florida. The animals were originally captured in the
Amazon near Leticia, Colombia. They had not been subjected to any training,
as far as we could determine, other than to take dead food fish.
Phonations
All of the recordings resulting in Figures 2 through 9 and in Table 1
were made at a tape speed of 7.5 inches, 19 cm., per second with a Uher
4000 Report-S recorder, which at that tape speed had a flat frequency response
of 40 to 20,000 cycles per second. An Atlantic Research Corporation model
IX- 5 7 hydrophone was used, with a special preamplifier designed and built
for the system by William E. Sutherland of the Lockheed-California Company.
Sonagrams (sound spectrograms) were prepared on a Kay Sona-Graph model
6061 A Sound Spectrum Analyzer calibrated from 85 to 8000 cps. When the
recorded tape speed is reduced by half, and then fed into the analyzer, the
FREQ. (KC.)
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Contributions in Science
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TIME (SEC.)
Figure 2. Phonation of Inia geoffrensis. Echolocation-like run on solid object (hydro-
phone) by a large adult of undetermined sex. Clicks emitted in darkness at the Fort
Worth Zoo, June 26, 1965.
response of the latter is increased to 16,000 cps. The effective filter band
width used in all of the analyses was 600 cycles.
Sounds were recorded with this system when the animals were resting
or swimming leisurely, when swimming rapidly, during feeding both in isola-
tion and in competitive situations, when both strange and familiar objects
0 0.1 02 0.3 0.4 0.5
TIME (SEC.)
Figure 3. Phonation of Inia geoffrensis. “Grate!’ No stimulus observed. Emitted in
daylight by an isolated juvenile male at the Steinhart Aquarium, September 30, 1964.
1966
Amazon Dolphin Behavior
7
Figure 4. Phonations of Inia geoffrensis. “Squawks!’ No stimulus observed. Emitted
in daylight by an isolated juvenile male at the Steinhart Aquarium, September 30,
1964.
were presented, during exposure to sudden loud noises and to lights flashed
out of darkness, in isolation and with another animal of the same species of
the same or opposite sex, in light and darkness, and with another Inia of the
same sex as well as another animal of the same sex (all males) belonging to
a different cetacean family ( Sotalia sp., family Delphinidae) .
0 0.1 02 0.3 0.4 0.5 0.6 0.7 0.8
TIME (SEC.)
Figure 5. Phonation of Inia geoffrensis. “Screech!’ No stimulus observed. Emitted in
daylight by an isolated juvenile male at the Steinhart Aquarium, September 30, 1964.
8
Contributions in Science
No. 108
Figure 6. Phonations of Inia geoffrensis. “Barks!’ No stimulus observed. Emitted in
daylight by an isolated juvenile male at the Steinhart Aquarium, September 30, 1964.
All of the Inia phonations we have observed consist of trains or bursts
of impulsive broad-band clicks, characteristic of most of the odontocetes
recorded to date. The major difference in Inia clicks, versus those of other
delphinids, is the apparently limited frequency content of individual clicks
(little energy above 10 KC). In contrast, Steno bredanensis clicks contain
16
14
312
5 10
6 8
u 6
oc 9
u. 4
2
0
0 OJ 02 0.3 0.4 0.5 0.6 0.7 0.8
TIME (SEC.)
Figure 7. Phonations of Inia geoffrensis. “Whimpers!’ No stimulus observed. Emitted
in daylight by an isolated juvenile male at the Steinhart Aquarium, September 30,
1964.
1966
Amazon Dolphin Behavior
9
16
14
3«
510
6 8
LJ c
K 6
u. 4
2
0
0 0.1 02 0.3
TIME (SEC.)
Figure 8. Phonation of Inia geoffrensis. “Crack!’ Emitted in the dark when a bright
light was suddenly flashed into the eyes of an adult animal, sex not observed, at the
Fort Worth Zoo, June 26, 1965. This “crack” immediately followed a train of clicks.
energy at frequencies in excess of 100 KC (Norris and Evans, 1966). Whether
this lower frequency limit is due to a characteristic of the species or an
instrumental limitation remains to be tested.
The Inia clicks recorded were of three types: click trains at repetition
rates of 30 to 80 clicks per second, single intense clicks, and sounds of the
Figure 9. Phonation of Inia geoffrensis. Jaw “snap” or “click!’ Made in daylight by a
mature animal, sex not observed, as it caught a small live goldfish at the Fort Worth
Zoo, June 26, 1965.
10
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burst-pulse type described by Watkins (1966). Because of the extremely fast
repetition rates involved in the latter type of phonation and the resolution
limits of the analyzer determined by the filter band width used (600 cps) this
group of sounds is characterized on the sonagrams by having a complicated
harmonic structure. The complexity of this structure is indicative of repetition
rates involved (Figs. 4, 5, 6, 7). The sounds which we have listed in Table 1
as “squawk,” “squeal,” “squeaky-squawk,” “screech,” “bark,” and “whimper”
are all of this burst-pulse type, but vary only in repetition rate and frequency
(in KC) of energy, and particularly of greatest energy. The click trains shown
in Figures 2 and 3 are representative of those with repetition rates of 30 to 80
per second. The pulses in Figure 3 have emphasis at different frequency bands.
As suggested by Schevill (1964) these can possibly be ascribed to uneven
response of instrumentation or reflect effects of the environment and structure
in the actual sound representing a species or individual characteristic (voice).
Although described by a variety of different adjectives, e.g., “echolocation-
like run,” “grate,” “squeal,” “squawk,” “screech,” “bark,” etc., all of these
audible sounds consist of trains or bursts of clicks which occur at various
repetition rates. Clicks occurring at rates of 10 to 20 per second can be resolved
by the observer to consist of individual pulses or clicks and thus have a grating
or clicking quality. Clicks occurring at faster repetition rates (40 per second
and more) are not recognized by the human ear as separate clicks but rather
the whole train takes on a tonal quality and thus becomes an “echolocation-like
run,” “creaking door,” “buzz,” or a “screech,” depending on the click rate.
This same explanation holds for “squawks” and “barks” which are short
bursts of clicks (0.05 to 0.3 second duration) at relatively high click repetition
rates (150 per second and up). “Squeals” (as referred to by Schevill and
Watkins, 1962; Schevill, 1964) and “whistles” (as referred to, for example,
by Evans and Prescott, 1962; Evans and Dreher, 1962; Lilly, 1962; Dreher
and Evans, 1964; Caldwell and Caldwell, 1965) are tones, pure and most
often with a simple harmonic structure, that cannot be resolved into individual
clicks. Signals of this latter type have not been observed to be produced
by Inia .
In considering the numbers of audible emissions in each category de-
scribed (Table 1) it is well to note two facts. First, that the phonations of
Inia are of such low sound level that they are not as readily audible as those
observed by us in several species of marine dolphins {e.g., Tursiops truncatus,
Tursiops gilli, Stenella plagiodon, Globicephala scammoni, Pseudorca crassi-
dens, Lagenorhynchus obliquidens, and Steno bredanensis) . Thus some sounds
may be lost in the ambient noise of the tank, and our counts may err on the
low side. In addition, it has been shown that the sound field of certain delphinids
is extremely directional, and therefore if the animals making them happened
to be facing away from our non-directional hydrophone, the sound might
not have been recorded at its full intensity, if at all. William E. Schevill and
William A. Watkins (pers. conversation, April, 1966) have found that there
1966
Amazon Dolphin Behavior
11
is a marked decrease in low-frequency sound intensity when soniferous indi-
viduals of the killer whale, Orcinus orca, turn away from the hydrophone.
Norris and Evans (1966) have observed a similar effect with high-frequency
sounds produced by Steno bredanensis, Stenella atienuata and Stenella sp.4
Keeping these facts in mind, however, Table 1 does represent a reasonable
picture of the types and relative numbers of the audible sounds emitted by
Inia geoffrensis as we observed them under a variety of captive conditions.
The maximum number of emissions per animal per hour observed with this
species was 52.5. This is quite low in comparison with sound emission rates
in Tursiops truncatus, which in many cases will exceed 180 emissions per hour,
and somewhat lower than the approximately 88 emissions per animal per
hour recorded in a 47-minute session for captive belugas, Delphinapterus
leucas, by Fish and Mowbray ( 1962), although we have observed the general
behavior of captive Inia and Delphinapterus to be quite similar. It should also
be noted that sound emission in some species (e.g., Tursiops truncatus , Stenella
atienuata, and Stenella sp.4) has been found by Powell (In press) to be very
periodic with quite regular cycles of vocalization and nonvocalization. It is thus
difficult to quantify the “vocalness” of Inia in comparison to marine dolphins,
but in general it is safe to say that Inia is less vocal, at least in the audible range,
than most marine delphinids observed to date.
As noted above, no pure-tone “whistles” were recorded, nor have any of
the attendants with whom we have talked reported an audible whistle from any
of the Inia in their charge. In this regard it is interesting to note that two
delphinids which are considered by many workers to be the more primitive
members of the family, or even in a separate family, also have been reported
not to produce sounds other than bursts and trains of clicks. Busnel, Dziedzic
and Andersen (1963, 1965) and Busnel and Dziedzic (1966) reported on
recordings of captive Phocoena phocoena and Evans (unpublished findings)
has recorded captive Phocoenoides dalli.
A “screech,” (Fig. 5), a harsh raucous sound, was recorded on only one
occasion. This is a burst-pulse type sound with a high pulse repetition rate
which on a sonagram forms a contour similar to, but not directly related to, the
pure-tone whistle contours of many delphinids. Similar “click contours” have
been illustrated for Phocoena phocoena by Busnel, Dziedzic and Andersen
(1963) and Busnel and Dziedzic (1966: figs. 45, 47 and 49). The “screech”
that we figure here was recorded with the gain on the recorder turned up suffi-
ciently high to pick up the faint sounds emitted by the species. When this single
loud sound was recorded, therefore, the system was somewhat overloaded.
4We have not applied a specific name to the small long-snouted Hawaiian spinner
porpoise discussed here. However, F. C. Fraser ( pers . comm, to D. K. Caldwell,
1965) has suggested that the name Stenella roseiventris (Wagner) be applied. Pub-
lished precedence for the use of this name, also based on Fraser’s personal remarks to
the authors, may be found in Brown, Caldwell and Caldwell (1966) and Morris and
Mowbray ( 1966) .
Table 1. Types and relative frequency of occurrence of non-extraneous phonations by six captive Inia geoffrensis. See
text for details of animals and study sites. Two additional animals were recorded for 70 minutes at the Aquarium of
Niagara Falls, but contamination in the form of a young Sotalia sp. was present and thus we have not tried to include the
possible Inia sounds here. Numbers in parentheses indicate number of each phonation per animal per hour.
12
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No. 108
Total number of
Phonations
Crack3 (Fig. 8)
Whimper3 (Fig. 7)
Bark3 (Fig. 6)
Screech3 (Fig. 5)
Squeak3
Squeaky-squawk3
Squawk2 (Fig. 4)
Grate1 (Fig. 3)
Creaking Door1
Echolocation-like
Run1 (Fig. 2)
Number of Minutes
Recorded
Number of Animals
Study Site
S
248
52.5
30
(6.71
69
14.8
376
v-H
*
<N 4
<N
w
/— N
O
©
VO ^
NO
w
O
^ oo
w
m
/—N
_ ON
o
r-
cn
ON •
T-H
m vo
w
r- i'-
t-*
o r i
o
T“l <N
1
w
— s
VO tN
00
^ oo
67
• 1
'w'
w •
VO ®
pi Tf;
<N rt
<N VO
<N
'-i ri
VO
o
m
©
NO
vo
NO
w
ro
o
»/->
o
OO
oo
NO
Tf
pH
<N
NO
-
-
CA
NO
G
o
X2
Steinhart
Bloomingt
Sea World
Fort Wort
Totals
Contain audible pulses at Vs recorded tape speed.
2 Although at recorded tape speed these all sounded the same, at Vs recorded tape speed some did and some did not contain audible
pulses.
3Do not contain audible pulses at Vs recorded tape speed.
1966
Amazon Dolphin Behavior
13
An Inia “squeal” was reported to us by Earl Herald and by Lawrence
Curtis, usually made when the animals in their charge were taken out of the
water. A “squeal” was also reported to Layne and Caldwell (1964: 102),
made by an animal out of water during transport in an airplane, and by an
animal (possibly the same one) in water at Silver Springs, Florida. While we
have no recordings of any of these “squeals,” it is possible that the sound
which we have described as a “screech” is involved. We had first used the term
“squeal” to describe it in our notes, and the fact that it has a somewhat con-
toured quality on a sonagram may be further evidence that the same sound
was heard by the observers noted above.
Schevill and Watkins (1962) did not attempt to apply word descriptions
to the Inia sounds they included on their record. However, upon listening to
the record we hear the sounds which we list in the present report as “echoloca-
tion-like run,” “whimper,” and “bark.” The sonagram which Schevill and
Watkins (1962: fig. 4) included appears to be the lower portion of a “bark”
(see our Figure 6) . Although they only show frequencies to four kilocycles per
second, it appears that the frequency range for the sound illustrated by Schevill
and Watkins actually extends higher, as it does in the “barks” we recorded
(Fig. 6).
Adults: For the two adults at the Fort Worth Zoo we were able to cor-
relate three audible sounds with observed behavior. These were low-frequency
click trains made on the unfamiliar hydrophone (Fig. 2) and on live food fish;
indistinct low-frequency clicks followed by a loud “crack” elicited by suddenly
flashing a bright light in their eyes out of darkness (Fig. 8) ; and a jaw “snap”
or “click” that was produced when one of the animals caught and bit down
suddenly on a live goldfish (Fig. 9). This last was not made when the Inia
chewed, but only as they caught the fish in one quick jaw snap.
Pulsed sounds have been experimentally demonstrated to be echolocation
devices in only one, or possibly two, cetaceans: Tursiops truncatus (see
Kellogg, 1961; Norris, et al, 1961) and Phocoena phocoena (see Busnel,
Dziedzic and Andersen, 1965). However, such sounds are known to be pro-
duced by, and are strongly suspect of being echolocation devices in many
other odontocetes (see Norris, 1964) . We were interested then in knowing first
whether this pulsed sound as recorded by Schevill and Watkins (1962) is
characteristic of Inia geoffrensis and secondly to learn whether it is utilized as
an echolocation device by this primitive species. Layne (1958: 16) suggested
that this may be so because of the ability of wild Inia to avoid nets. Layne and
Caldwell (1964: 95) included behavioral observations on two captive animals
that suggested their use of echolocation in environmental exploration. Based
on observations and correlations with behavior, our conclusion is that the low-
frequency pulsed sounds described as “echolocation-like runs,” “creaking
door,” or “grate” are probably utilized by the adult Inia in echolocation. Both
males and females demonstrated this ability. The use of the more rapidly-
pulsed sounds such as the “squawk” (Fig. 4) for echolocation is more doubtful.
14
Contributions in Science
No. 108
The low-frequency echolocation-like runs were made most frequently as the
animal approached the hydrophone (Fig. 2). We were able to observe this
visually in both daylight and artificial light. The aquarium in which we worked
with the adults was constructed so that we could obtain almost total darkness,
and therefore we were able to record the adults under this condition also. We
could not see the animals at the time, but the tapes make it apparent that the
audible trains of clicks almost invariably precede the sound of the animals
rubbing against the listening gear. When we introduced live fish into the tank
in darkness, audible click bursts, and the subsequent jaw “click” associated
with fish captures were also produced.
Trains of audible clicks were emitted both in light and darkness. Quanti-
tatively, however, the number produced in darkness was 25.3 per animal per
hour, as opposed to 6.9 per animal per hour in daylight or artificial light.
The “crack” elicited under stress (Fig. 8) is virtually the same as that
produced by Tursiops truncatus under the same fright stimulus (Caldwell,
Haugen and Caldwell, 1962). However, we cannot state positively that this is
not a jaw clap as we heard it only momentarily as the light was flashed.
The jaw “click” or “snap” produced (Fig. 9), apparently by the teeth
hitting together, when an animal caught a small fish, is loud enough to be heard
by a diver underwater, as the trainer, Gary Hill, reported having noted this
sound when he was doing underwater feeding of the Fort Worth animals.
No audible phonations accompanied sexual behavior that resulted in an
erection by the male. None occurred when the Inia were presented with
familiar objects.
Inasmuch as no Inia whistles have been recorded, it is therefore likely that
Inia makes use of clicks in communication as well as in echolocation, much
like Physeter and Phocoena.
A puzzling loud “gurgling growl,” obviously not of extraneous (to the
tank) origin, was recorded several times from these adults. Evans (unpub-
lished) has recorded these sounds emanating from captive Tursiops truncatus
during feeding and accompanying defecation, and it is suggested that they are
the sounds of digestive processes within the animal and the nature of the sound
lends itself to this explanation.
Subadults and Juveniles: Although the two subadult males at Sea World
were capable of creating a train of audible pulses or clicks (Table 1 ) , we were
unable to elicit them, either in daylight or dark, on a specific stimulus such as
feeding dead fish, or by the presentation of strange or familiar objects. Even
when obviously startled by loud noises or having objects suddenly thrown at
their heads they did not vocalize audibly but only jerked and swam violently
away. None of the audible sounds by these animals listed in Table 1 were
correlated with any stimulus that we could note. Dr. Fish and her associates
recorded audible clicks from these two animals when the water was murky. In
addition, those workers recorded audible sounds described by them later as
1966
Amazon Dolphin Behavior
15
“chirps” and “squawks.” Upon listening to copies of their tapes, we would have
listed the “chirps” as “squeaks” and the “squawks” we would have listed as
“squeaky squawks.”
The audible vocalizations of the young males at Bloomington and at the
Steinhart Aquarium were similarly unrelated to observable stimuli. When the
Steinhart animal was 52.5 inches (133 cm.) long, we worked with him in-
tensively at night attempting to obtain a positive correlation of audible click
trains and feeding. We were totally unsuccessful, although we were able to
work in a dark building with only low-intensity ambient light from a nearby
window. Dead fish were first thrown into the tank with a splash and varying
amounts of time allowed to elapse before turning on the lights to see whether
the fish had been found. The animal usually found the fish in six seconds or
more, but failed in four seconds or less. No phonations were heard and we
have no explanation for his ability to locate the sinking fish in the dark after
the initial cue of the splash unless his hearing is so acute that he could actually
hear it falling through the water, his tactile sense so well developed that cur-
rents generated by the falling fish could be detected at close range after the
animal had generally located the fish from the initial splash, or he (and all
Inia ) has the ability to echolocate in the high-frequency (inaudible to humans)
range.
Layne and Caldwell (1964: 96) discussed the probable sensory function
of the structurally-complicated snout bristles of Inia, and the young animal
under study was so well endowed with them that he was given the name
“Whiskers.” Gustatory cues are probably ruled out by the fact that if the fish
were silently either slipped into the tank and allowed to drop or if quietly hand
held in the tank, the animal failed to find it in several minutes. If in fact he was
able to hear or “feel” the fish falling through the water after the attention-
getting cue of the initial splash when a fish was thrown, then in the latter
experiments without the splash it follows that although the fish falling through
the water would make the same sound or generate the same currents as before,
the splash must be necessary to draw his attention. The fact that this animal is
obviously able to emit audible sounds that we normally consider echolocation
bursts, but did not do so in these experiments, makes for a puzzling picture.
One possibility that suggests itself is an investigation of the factor of learning
in this species. Although audible pulsed phonations are obviously present in
the very young, i.e., the four-foot (122 cm.) male at Bloomington, California,
the use of these same phonations to echolocate objects may require experience.
Fish, Mowbray and Perkins told us in early 1966 that they recorded good
audible click trains (over 60 in a 22-minute period) from the Steinhart
Aquarium young male. These observers suspected that the click trains were
emitted in response to spectator activity, insertion of the hydrophone and fish,
movement of large garfish (Lepisosteus spatula ) in an adjacent and connected
enclosure, movements of the scientific investigators, and perhaps sponta-
16
Contributions in Science
No. 108
neously. However, like us, these investigators apparently did not record click
trains produced concurrently with a specific situation which would seem to call
for the use of echolocation.
The possibility that the animal was echolocating in the very high-
frequency ranges, inaudible to the human ear, cannot be discounted, but our
studies on this possibility have been inconclusive (see Caldwell, Caldwell and
Evans, In press ) .
Visual Acuity
The eyes of Inia are so small that doubt has been raised as to their being
functional. However, Layne (1958: 16) concluded that vision, at least above
the surface of the water, seemed good in this species in the wild. Layne and
Caldwell (1964: 93) suggested that in captive animals it is also good under-
water as well as above. The behavior of the eight animals that we observed
intensively leads us to believe that vision is not only acute but is the preferred
or primary device for environmental investigation. Although animals tend to
use any sensory device available, there is usually a tendency to rely on one in
preference to the other if the preferred sensory input is available. Since our
work was done in aquaria, good visibility by the animals was usually present
during daylight hours. All of the eight animals visually inspected any change
in their environment, including new sounds, when they visually inspected the
source. They visually inspected food offered them, and if the food was dropped
or thrown into the tank, they searched for it visually. Only rarely in the two
adults at Fort Worth were click trains added to visual inspection in daylight.
Tiny bits of left-over food were found visually. As mentioned earlier, no
echolocation-like bursts either day or night were heard from the juveniles or
subadults during observed episodes of environmental investigation. They were
also rare in the adults if sufficient light was available for us to observe. It was
noted that the eyes of the adults at Fort Worth demonstrated a pink eye shine
in the dark, which is indicative of good vision in low light intensities (Walls,
1963). Unfortunately, the degree of night vision has not been investigated in
any cetacean, but our evidence suggests that in Inia it is excellent.
Good vision in this species is apparently less hampered by the reduced
size of the external opening of the eye (the eye itself is comparatively large and
well-enervated) than by the large bulging cheeks. These cheeks are so enlarged
that they apparently prevent the animals’ seeing much below the horizontal
plane of the eye. This problem is solved by their turning over and swimming
upside down, whereby a good field of vision is opened up below the animal.
The small male at Steinhart Aquarium always visually checked the bottom of
his tank after feeding by swimming upside down around the tank and recover-
ing the small bits of fish debris lost during feeding. If he dropped a fish on the
bottom he immediately turned over and swam upside down around the tank
until he apparently saw it and then recovered it. He also swam upside down
when pursuing live fish near the bottom of his tank. Upside down swimming
1966
Amazon Dolphin Behavior
17
in this species in semi-wild conditions has been noted as occurring often (Layne
and Caldwell, 1964: 88f.; Allen and Neill, 1957: 328) and increased visibility
to the Inia is the probable explanation. Although we did not always see them
actually find food or other objects on the bottom, we observed this upside down
swimming in all of the animals we studied, including the two at Homosassa
Springs, Florida, and the two at the Aquarium of Niagara Falls. Mr. Brady
told us that more of this upside down swimming occurred when the latter ani-
mals were first captured than when we observed them some five months later,
suggesting that it is a natural behavior that for some reason began to be
abandoned by the captives.
Layne (1958: 19) noted that on one occasion the gaping jaws of a wild
Inia appeared above the surface of the water under circumstances which
suggested that it was feeding. That this was probably the case is suggested by
our underwater observations of the Fort Worth Inia during a sequence in
which they were being fed live goldfish. The small fish swam near the surface,
and in slowly pursuing them, the Inia positioned themselves at about a 45°
angle just under the surface with the jaws extending out of the water. In
attempting to catch the fish, which they eventually did, the Inia opened and
closed their jaws in a manner similar to that described by Layne. As the upside
down swimming position seems to permit better vision for objects on the
bottom, so did the upright angled position appear to make vision over the
bulging cheeks more practical for observing a small target just beneath the
surface and just ahead of the Inia — more so than would a direct horizontal
head-on approach to the target. We have also noted that Inia fed from the
surface at the side of their tank also assume a similarly angled attitude, in
which they are obviously visually observing the feeder, and then open and close
their jaws as if begging from the attendant.
Miscellaneous Behavior
Curiosity and Manipulation: It is impossible not to compare the striking differ-
ence in the intensity and duration of fear in this species with what we have
noted in the Atlantic bottlenosed dolphin. A naive animal of this latter species
requires many hours or days to acclimate to strange objects (McBride and
Hebb, 1948; Caldwell and Caldwell, 1964). Although young animals and
captives of long duration may adjust more quickly to strange objects than the
recently captured adults, it usually takes several days for any individual
Tursiops to approach unusual objects in its environment without reinforce-
ment of approach behavior.
None of the Inia ever showed any fear of the hydrophone. Within minutes
after introduction of the listening gear they often were using it as a play object,
tactual stimulant or sex object. Loud tapping against the walls of the aquaria
caused immediate approach toward the direction of the noise. With the initial
fear so evident in the bottlenosed dolphin, the curiosity and playfulness of Inia
18
Contributions in Science
No. 108
became immediately evident. The small male at the Steinhart Aquarium
demonstrated a duration of three hours almost continuous play with the
hydrophone, which he had approached immediately on its being introduced.
We have never observed such behavior by any Tursiops of any age, sex or
acclimation to captivity. The play by the Inia was discontinued only when the
hydrophone was removed.
Sexual play: Although the Steinhart Aquarium animal was only 52.5 inches
(133 cm.) long at the time just noted above, he had several erections while
playing with the gear when it rubbed against his genital area. This same animal
had previously been observed masturbating against the net webbing which
divided his tank (Earl S. Herald, pers. conversation, 1965, and motion picture
films in his and the Caldwells’ files) , and along the corner of the tank where the
wall met the floor (Edward Mitchell, pers. conversation, 1965; observations by
the Caldwells May, 1966).
The animal also once displayed an erection that lasted several seconds
immediately following defecation.
Mitchell {pers. conversation, 1965) also reported observing the same
animal in January, 1965, at the Steinhart Aquarium as it tried to “eat” drops
of water which were falling into its tank from an unseen source high above.
Mitchell noted that the Inia attempted to “eat” the drops about once every
minute and that the dolphin became progressively more agitated on each
attempt. After the procedure continued for about five minutes, the Inia pulled
away from the area of the dripping water and clumsily bumped into a wall of
netting that divided his tank. At this time he had an erection and proceeded to
masturbate against the corner of the tank as noted above. Mitchell reported
that while masturbating the Inia would rub the tip of its snout against the net
and sporadically rotate its body and thus its snout while still in contact with the
net. The dolphin’s eyes reportedly were open during all of the observed display.
This behavior was essentially the same as that recorded in the film noted above
which was made at about the same time. After the sexual display observed by
Mitchell, which lasted for about 10 to 15 minutes, the Inia lay on the bottom
of the tank on his right side or upside down without actively moving, or else
slowly swam around the tank and surfaced to breathe very slowly and appar-
ently with his eyes closed.
We have also observed this animal assume the upside-down stance while
it was resting or sleeping on the bottom of the tank. The position (Fig. 10) is
so unusual that we were startled when we first saw it, and a number of visitors
to the aquarium have been overheard to comment that the animal seemed to
be dead. Such a position must be associated with the apparent reduced fear
responses of Inia, because in nature it would seem to leave the animal open and
vulnerable to attack. However, according to available reports, when one
considers that in nature Inia apparently has no predators other than man on
rare occasions, and that it normally lives in an oft-times shallow and rather
1966
Amazon Dolphin Behavior
19
Figure 10. Inia geoffrensis. Juvenile male (“Whiskers”) in a typical upside-down
sleep position at the Steinhart Aquarium in late 1964. Also note the bulging cheeks.
protected and less hostile environment than that provided by the open sea,
this attitude of rest should be advantageous because it would seem to permit a
deeper sleep, if only for a brief period, than that achieved by marine cetaceans
which apparently only lightly doze while resting near the surface or while
actually on the move.
We suggest that the reduced fear response toward strange objects may
also be related to the conditions in the natural environment of Inia. It lives in
areas where it might encounter many more strange objects than a marine
dolphin. Not only does Inia venture into small streams, lakes and even into
flooded forest areas where there might be much debris as well as standing
vegetation, but the main stream of the Amazon itself is noted for the great
amount of floating debris that it continually carries. Evans (unpublished) has
noted a similar lack of fear toward strange objects by harbor seals (Phoca
vitulina) in California. Like Inia, Phoca lives in areas such as lagoons, coastal
rivers and bays where there may be more strange objects than more oceanic
marine mammals normally encounter.
With further regard to sexual behavior, on one occasion during our
observations at the Aquarium of Niagara Falls, the small male Sotalia in the
20
Contributions in Science
No. 108
tank with the two male Inia frequently rubbed the top of its head as if trying to
scratch it. This behavior included rubbing against the sides of the tank, and
swimming upside down to rub the top of its head along the floor of the tank.
During this same period, one of the Inia was seen rubbing his genital region
on several projecting pipes near one bottom corner of the tank. Shortly there-
after, the three animals were swimming together in a normal upright position
when the Sotalia rose beneath the same Inia just noted and began to rub the
top of its head against the underside of the Inia, including the genital region
of the latter. The Inia had a full erection shortly thereafter which lasted for
nearly a minute while the two animals remained together. During this time, and
while both were swimming in a regular circular pattern around the tank, the
Inia appeared to use the Sotalia as a sexual stimulus and even tended to force
the smaller animal toward the bottom in an apparent attempt to masturbate
against it. The Sotalia apparently had only accidentally stimulated the erection
by the Inia when it rubbed against the underside of the larger animal, and it
immediately appeared to try to escape the attentions of the Inia when the
erection occurred.
In no case have we seen one of the masturbating Inia effect an ejaculation.
As noted above, we observed one attempt at intromission by the adult
male Inia with the adult female at Fort Worth. The technique was similar to
that we have often observed in captive marine delphinids. The male approached
in an essentially upside down position beneath the female, at about a 30- to
45-degree angle to her, and attempted intromission from the inverted position.
Lawrence Curtis, Gary Hill, and Gary K. Clarke, the latter now Director of
the Topeka (Kansas) Zoological Park, have observed the breeding behavior
of the Fort Worth pair in much greater detail and have photographed much of
it. We understand that these observations are to be prepared for publication.
Competitiveness
With one exception, we observed none of the competition for food be-
tween the two sets of subadult males, at Sea World and at Niagara Falls, or the
adult male and female, at Fort Worth, that is so prominent in Atlantic bottle-
nosed dolphins. (T. truncatus) during feeding. However, the number of Inia in
each of the competitive situations was limited to two, and they were at all times
well fed. Because of the expense of air-shipping these animals from South
America, they are treated with even more exquisite care than the bottlenosed
dolphins in that they are not subjected to difficult training programs or show
procedures in order to secure their food. The dominant male at Fort Worth, in
fact, jealously dominates the simple show there and will no longer allow the
smaller female to perform. However, this does not appear to be a matter of
competition for food because both animals are always fed to satiation.
The one observed exception to lack of food competition took place on 9
January 1966 at Sea World in San Diego. One of the subadult males, the least
1966
Amazon Dolphin Behavior
21
active one, had just taken a dead food fish from the hand of a feeder and was
slowly swimming away with the limp fish trailing outside the side of its mouth.
The other subadult male, usually more active, slowly swam up alongside the
first Inia, head to head going in the same direction, and very deftly snipped olf
the trailing fish about midway the length of its long jaws and very close to the
jaws of the first animal. This incident took place early in the day, before the
animals had received much food from the public which is allowed to feed the
Inia, and such behavior was not even suggested later on in the day after the
animals had been well fed by the public. Earlier in the day when the competi-
tion had been observed, both the animals had very actively “begged” for food
by swimming up to the side of the tank and opening their jaws to the observers
standing there. While the Inia continued to accept food later in the day, they
did not appear to actively “beg” for it and their general attitude was one of
much less interest in food than it had been earlier.
Layne and Caldwell (1964: 103) noted two brief instances of food
competition between a large and a small male; in one case the activity was very
similar to that noted above at Sea World.
No food or other competition was noted between the two male Inia at
Niagara Falls or between them and the smaller male Sotalia housed with them
in a relatively small tank, although the Sotalia seemed to be hesitant about
feeding from the hand of the attendant while the Inia were being fed.
On the other hand, Kent Burgess told us in May, 1966, that he has
observed apparent competition for human affection between the two male
Inia at Sea World.
Summary
Adult Inia geoffrensis have an audible pulsed phonation that is concurrent
with investigation and search situations. This apparent echolocation device is
more frequently employed when visibility is poor but may be employed when
visibility is good. Juveniles and subadults of the same species have a similar
audible pulsed phonation, but in work on four animals of this class, we have
not been able to demonstrate a correlation of the audible phonation and a
situation that would indicate that it was used as an echolocation device.
Although an echolocation device apparently is available to at least the
adults of the species, in our observations in aquaria it was not the sensory
system of choice. Vision was apparently the preferred sensory device.
No pure tone whistles were recorded, but several other phonations are
included in their repertoire.
Fear is less easily precipitated in this species than in the Atlantic bottle-
nosed dolphin, and it is of shorter duration. Curiosity, playfulness, and early
and frequent sexual play are also characteristic of this primitive species.
Competitive feeding behavior so familiar in the Atlantic bottlenosed
dolphin was not usual in the Inia studied.
22
Contributions in Science
No. 108
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LOS
ANGELES
CONTRIBUTIONS
COUNTY
MUSEUM
IN SCIENCE
UMBER 109
October 27, 1966
A NEW HALIOTID FROM GUADALUPE ISLAND, MEXICO
( MOLLUSC A : GASTROPODA )
By Robert R. Talmadge
|i
Los Angeles County Museum of Natural History • Exposition Park
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CONTRIBUTIONS IN SCIENCE is a series of miscellaneous technical papers
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less specially requested, page proof will not be sent to the author. 100 copies of each
paper will be given free to a single author or divided equally among multiple authors.
Orders for additional copies should be sent to the Editor at the time corrected galley
proof is returned; appropriate forms for this will be included when galley is sent.
David K. Caldwell
Editor
A NEW HALIOTID FROM GUADALUPE ISLAND, MEXICO
( MOLLUSC A : GASTROPODA )
By Robert R. Talmadge1
Abstract: A new subspecies of Haliotis corrugata Gray
1828, is described from Guadalupe Island, Baja California, Mex-
ico. It differs from the nominate race in size, shape, and sculp-
ture, and is endemic to the island.
During October, 1965, the Los Angeles County Museum of Natural His-
tory sponsored an expedition to Guadalupe Island off the Pacific Coast of Baja
California, Mexico. Among the specimens obtained was a series of shells of
Haliotis corrugata Gray, 1828. This species had been known to occur at the
island, and it was also known that specimens differed in shape and size from
mainland examples. However, the few previously available specimens from
Guadalupe Island were so heavily encrusted that details of the shell sculpture
could not be examined. Based upon the original specimens it was anticipated
that a large clean series would exhibit differences warranting a subspecific
ranking. The large series of shells collected by R. S. Owen, Lawrence Thomas,
and Dean Tyler substantiated this hypothesis.
The necessary scientific collecting permit was arranged by Lie. Jorge
Echaniz R. of the Dirrecion General de Pesca e Industrias Conexas, de la
Secretaria de Industria y Comercio. I am especially grateful to Dr. James H.
McLean of the Los Angeles County Museum of Natural History for making
these specimens available for study and for providing aid in the preparation of
the manuscript. Photographs are by Mr. Armando Solis, Museum photogra-
pher.
Haliotis corrugata oweni, new subspecies
Figures 1 and 2
Description : shell auriform, deeply arched, nearly circular with the inset
apex lower than the dorsal surface. On the holotype there are three open
siphonal pores, each on an elevated projection. The sculpture is very dense,
consisting of rather coarse cording having small, sharply formed nodes, which
form a diagonal series of ridges across the surface of the shell. The exterior
coloration is a dull pinkish-tan, similar to weathered brick. The nacreous in-
terior is tinted with pink, green, and blue, but often with an overwash of yellow.
The holotype measures: long. 146, lat. 125, alt. 55 mm.
As in specimens of the typical Haliotis corrugata, juvenile specimens (less
than 45 mm. in length) neither exhibit the deep arching, nor the dense nodes.
Most subspecies in the genus Haliotis have juveniles that differ but slightly
(Talmadge, 1962). The diagnostic features appear when the animals are at
billow Creek, California: Field Associate, Department of Invertebrate Zoology,
California Academy of Sciences.
1
2
Contributions in Science
No. 109
Figure 1. Haliotis corrugata oweni, new subspecies, holotype (LACM). Long. 146,
lat. 125, alt. 55 mm.
1966
New Gastropod Mollusk
3
Figure 2. Haliotis corrugata oweni, new subspecies, paratype (LACM). Long. 105,
lat. 84, alt. 42 mm.
4
Contributions in Science
No. 109
least subadult, as the muscle scar begins to develop. In shells of this subspecies
the muscle scar begins to form when the shell attains a length of 65 to 70 mm.
At a length of 80 to 90 mm., the muscle scar is well formed and the shell has a
mature appearance.
Type Material : Holotype, Los Angeles County Museum of Natural His-
tory No. 1147 (Fig. 1); paratype, No. 1148 (Fig. 2). Additional paratypes to
be deposited and numbered by recipients in the collections of: San Diego Nat-
ural History Museum, Stanford University, California Academy of Sciences,
United States National Museum, and the collections of R. S. Owen (Pescadero,
California), and Robert R. Talmadge (Willow Creek, California).
Type Locality : Guadalupe Island, Baja California, Mexico, between a
small offshore reef and the shore at the west anchorage, depth 20 feet, collected
by Mr. R. S. Owen, October 27-29, 1965.
Discussion : This subspecies appears to be endemic to Guadalupe Island,
as the diagnostic features (dwarfed, more highly arched, denser sculpture)
have not been noted in populations from the mainland or other coastal islands.
The soft parts were identical to those of other populations in the same species.
In the haliotids, the recognized subspecies have similar or identical soft parts
(Talmadge, 1964).
Haliotis corrugata oweni is dwarfed, possibly due to scarcity of food, or
perhaps due to lack of living space— the shore of Guadalupe Island is steeply
sloping, leaving a rather limited intertidal and shallow subtidal area. Owen’s
field notes indicate that the subspecies is more prevalent on the western or
exposed side of the island, where stronger wave action may create a more
favorable habitat.
The dimensions of Guadalupe Island populations were compared to those
of mainland populations by determining ratios. The ratios of the Guadalupe
Island shells based upon length = 1000 are: length, 1000; width, 850; height.
350. Mainland populations of similar size can not be compared because they
are not mature and thereby are more depressed. Adult specimens from the
mainland, twice the length of the Guadalupe Island specimens, yielded a ratio
of: length, 1000; width, 800; height, 300. The greater height of the Guadalupe
Island population is apparent.
Literature Cited
Talmadge, R. R.
1962. The Linnaean Haliotis varia in Australia. Mem. Natl. Mus. Melbourne,
No. 25: 233-241, text Figs.
1964. The races of Haliotis fulgens Philippi (Mollusca: Gastropoda). Trans.
San Diego Soc. Nat. Hist., 13(18) : 369-376, Fig. 1.
LOS
ANGELES
CONTRIBUTIONS
COUNTY
MUSEUM
IN SCIENCE
UMBER 1 10
October 27, 1966
GALEUS
PIPERATUS, A NEW SHARK OF THE FAMILY
SCYLIORHINIDAE FROM THE GULF OF CALIFORNIA
By Stewart Springer and Mary H. Wagner
Los Angeles County Museum of Natural History • Exposition Park
Los Angeles, California 90007
CONTRIBUTIONS IN SCIENCE is a series of miscellaneous technical papers
in the fields of Biology, Geology and Anthropology, published at irregular intervals
by the Los Angeles County Museum of Natural History. Issues are numbered sepa-
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GALEUS PIPERATUS, A NEW SHARK OF THE FAMILY
SCYLIORHINIDAE FROM THE GULF OF CALIFORNIA
By Stewart Springer1 and Mary H. Wagner2
Abstract: Three specimens trawled from a depth of ap-
proximately 275 m. midway between Tiburon and Angel de la
Guardia Islands in the Gulf of California, the only known
specimens of the genus Galeus from the eastern Pacific, con-
stitute the type series of a new species which is described and
illustrated.
Among specimens collected at Station No. 64A2-16 of the R/V Alaska,
operated by the California Department of Fish and Game, were five small
sharks. Two of these were lost or misplaced but the collectors, John E. Fitch
and Robert J. Lavenberg, sent us the three remaining specimens for study.
These sharks, the first of the genus Galeus reported from the eastern Pacific
Ocean (Lavenberg and Fitch, 1966), represent a new species which is de-
scribed here. The name proposed for the new species is from the Latin
piperatus and refers to the peppered appearance of the ventral and lateral
surfaces.
Galeus piperatus, new species
Figures 1 and 2
Holotype: Los Angeles County Museum of Natural History No. 7552, adult
female, 302 mm. total length; R/V Alaska Station No. 64A2-16, at 28°55/N,
112°50.5/W, in the Gulf of California; 6 April 1964.
Paratypes: Los Angeles County Museum of Natural History No. 8818,
immature female, 256 mm. total length; U.S. National Museum 200413, adult
female, 296 mm. total length; both paratypes from R/V Alaska Station
No. 64A2-16.
Comparative Material Examined: The holotype of Galeus eastmani
(Jordan and Snyder) 1903, and four paratypes of G. sauteri (Jordan and
Richardson) 1909, from the western Pacific; series of G. arae (Nichols) 1927,
and G. cadenati Springer 1966, from the western Caribbean Sea; and G. melas-
tomus (Rafinesque) 1810, and G. polli Cadenat 1959, from the eastern
Atlantic. We have not seen specimens referable to G. murinus (Collett) 1904,
G. jenseni (Saemundsson) 1922, from the northeastern Atlantic, or G. hert-
wigi (Englehardt) 1912, from Japan.
Diagnosis: Galeus piperatus, although not easily separated on the basis
of one character alone, is distinct in that it has the following combination of
1Research Associate, Los Angeles County Museum of Natural History; Fishery
Biologist, U.S. Bureau of Commercial Fisheries, Stanford, California.
2Biologist, U.S. Bureau of Commercial Fisheries, Stanford, California.
1
2
Contributions in Science
No. 110
Figure 1. Galeus piperatus, new species, lateral view and ventral aspect of head of
holotype, LACM 7552.
1966
New Shark of the Genus GALEUS
3
characters that distinguish it from all other species of scyliorhinid sharks:
small size (females reach maturity at 30 cm.); a narrow crest of enlarged
denticles on the dorsal edge of the caudal fin but no enlarged denticles on the
leading edge of the lower caudal lobe, small denticles on the roof of the mouth
anteriorly and on the tongue; a lack of well defined dorsal blotches in the
adult, fine pepper-like spots on the ventral and lateral surfaces, each spot
composed of a single melanophore; inside of mouth black.
Description of holotype: A 302-mm. female with characteristics of Scylio-
rhinidae: caudal axis not elevated; two dorsal fins and anal fin; dorsal fins nearly
equal, origin of first just posterior to mid-point of base of pelvics; body long
and slender, head long and somewhat depressed; caudal peduncle without
precaudal pits; spiracles small, close to orbits; outer surface of nictitans en-
tirely covered with denticles; conspicuous pores on lower side of snout.
Proportional measurements in percent of total length:
Trunk at pectoral origin: width, 9.9; height, 8.9.
Snout length in front of: outer nostrils, 3.3; mouth, 6.6.
Eye: horizontal diameter, 4.0; vertical diameter, 1.3.
Mouth: width, 8.3; length, 3.6.
Nostrils: distance between nasal apertures, 2.6.
Labial furrow lengths: upper, 1.3; lower 1.7.
Gill opening lengths: first, 2.6; fifth, 1.4.
First dorsal fin: vertical height, 3.0; length of base, 4.6.
Second dorsal fin: vertical height, 3.0; length of base, 5.3.
Caudal fin: upper margin, 28.5; lower anterior margin, 8.9.
Pectoral fin: anterior margin, 12.6; width, 8.9.
Pelvic fin: overall length, 11.6.
Distance from snout to: eye, 6.0; first gill-opening, 15.6; fifth gill
opening, 21.5; first dorsal, 43.4; second dorsal, 61.9; upper
caudal, 71.2; pectoral, 20.2; pelvic, 36.7.
Interspace between: first and second dorsal, 12.2; tip second dorsal
and caudal, 2.3.
Dermal denticles small, numerous, covering entire body, three-pointed
central one longest, typical shoulder denticles about 0.4 by 0.4 mm., denticles
in crescent-shaped patch on anterior roof of mouth, 0.2 mm., leaf shaped
(Fig. 2); greatest width caudal crest 1.25 mm., originating approximately
5.0 mm. from tip of second dorsal, largest scales in crest less than 1.0 mm.
long.
Teeth small, numerous, similar in both jaws, about 3 1 rows on each side
of upper jaw, 26 on one side of lower, largest tooth height about 1.0 mm.;
upper teeth with five cusps, except tricuspid near center of jaw, middle cusp
much the longest; lower teeth similar, central cusp lower toward angles of jaw;
tooth arrangement alternate.
4
Contributions in Science
No. 110
Total number of vertebrae 121; monospondylous vertebrae 33, caudal
vertebrae 52.
General color grayish brown, dorsal surface somewhat darker, lighter
ventrally, but demarcation between upper and lower surfaces not distinct; fins
slightly darker than dorsal surface, but trailing edges of dorsals and pectorals
marked by translucent bands about 1.0 to 2.0 mm. wide; entire body sprinkled
with black melanophores 0.1 to 0.3 mm. diameter, especially noticeable on
ventral surfaces; inside of mouth black.
Comparisons: Galeus piperatus, in comparison with all the specimens of
Galeus that we were able to examine, is closest to Galeus cadenati from the
Atlantic. G. piperatus differs from this species in having a shorter interdorsal
distance, a shorter anal base, and longer gill slits. The color difference is
marked as well. In the adult G. cadenati , dark brown saddles and blotches are
present on the dorsal surface, and the dorsal fins are darker anteriorly and
dorsally; the type of G. piperatus is devoid of dark brown blotches, and the
paratypes have irregular and indistinct blotches that contrast little with the
background color.
In Galeus piperatus a crest of enlarged denticles extends along the upper
edge of the caudal fin from its base to about two-thirds the distance toward the
tip. A caudal crest is present on all members of the genus Galeus, and, among
other sharks of the family Scyliorhinidae, is also present on Figaro boardmani
I mm
I I
Figure 2. Camera lucida drawing of denticles that occur in a crescent-shaped patch
located anteriorly on the roof of the mouth of Galeus piperatus, new species. Similar
denticles are present on the tongue. We found mouth denticles in all species of
Galeus that we examined. They differ from denticles of the external surfaces in be-
ing lanceolate instead of three-pointed and in being much smaller.
1966
New Shark of the Genus GALEUS
5
Table 1
Measurements, expressed as percent of total length, of the holotype and two
paratypes of Galeus piperatus and two other species of Galeus from the Pacific.
The table was prepared to conform with the tables given by Springer (1966)
to facilitate comparison.
Species
Galeus piperatus
Galeus eastmani
Galeus sauteri
Number of specimens measured
3
1
4
Length range in millimeters
256-302
337
360-365
Tip of snout to —
-anterior nasal aperture
3. 3-4.3
3.0
3. 1-3.9
-posterior nasal aperture
5. 0-5. 9
5.0
5. 0-5. 6
-front mouth
6. 6-8. 2
6.2
6. 1-6.7
-eye
6. 0-7. 5
5.6
6. 3-7. 2
-gill I
15.6-17.1
14.8
15.6-16.7
-gill V
21.5-22.8
18.4
18.6-19.7
-origin pectoral
20.2-22.2
17.5
17.5-18.4
-1st dorsal fin
41.0-45.7
42.1
44.5-45.9
-pelvic fins
36.7-40.9
35.6
36.1-39.5
-anal fin
51.2-54.6
50.7
53.9-54.8
-2nd dorsal fin
45.4-61.9
62.3
63.3-63.9
-lower caudal fin
68.4-71.6
71.2
70.6-71.2
-upper caudal fin
68.6-73.0
71.2
72.0-75.1
-anus
40.7-44.7
38.0
40.6-43.6
Length upper caudal fin
28.3-29.7
29.7
26.7-29.5
Base 1st dorsal fin
4.6-5. 1
4.7
4. 2-5. 3
Base 2nd dorsal fin
4. 8-5. 3
5.6
4. 2-5.0
Base anal fin
12.6-13.3
11.9
11.7-13.9
Distance between dorsal fins
10.9-12.3
15.4
13.1-13.7
Internasal distance
2.6-3. 1
2.1
2. 2-3.0
Length orbit
3. 9-4. 4
4.2
3. 6-4. 2
Length lower labial fold
1. 7-2.7
1.5
1.4-1. 7
Length upper labial fold
1. 3-2.0
1.2
1.4-1. 9
Width mouth
8. 3-9. 2
6.2
6. 0-7.0
Length mouth
3. 6-4. 8
3.7
3. 8-4. 7
Height gill I
2. 3-2. 6
1.5
1. 1-1.4
Height gill V
1.2-1. 4
0.9
0.8-0. 8
Tip 2nd dorsal to caudal
1.0-2. 3
1.5
1.9-3. 1
Tip pelvic to 2nd dorsal
10.6-13.3
14.5
9.6-12.2
Tip pelvic to anal
2. 0-2. 6
5.0
2.2-3. 3
Tip pelvic to lower caudal
18.4-19.9
24.3
16.7-19.7
Length outer margin pectoral
12.6-13.0
9.5
10.1-11.1
Length anterior margin 1st dorsal
7. 8-8. 9
7.4
6. 7-7. 5
Length anterior margin 2nd dorsal
6. 8-7. 8
7.4
5. 5-6. 7
Length anterior margin anal
5. 9-7. 6
5.9
5. 3-6. 4
Distance eye to spiracle
0.6-1. 3
0.6
0.8-0. 8
6
Contributions in Science
No. 110
(Whitley) 1928, Parmaturus xaniurus (Gilbert) 1891, Parmaturus pilosus
Garman 1906, as well as on Apristurus profundorum (Goode and Bean) 1896.
Figaro is unique in having enlarged denticles not only on the upper caudal lobe
but also on the leading edge of the lower caudal lobe. The denticle crest of
A. profundorum lacks the much enlarged marginal denticles that are present
in Galeus. The two species of Parmaturus have caudal crests similar to those in
Galeus , but have shorter and thicker snouts and are generally much more
robust.
As in other species of Galeus, the marginal denticles of the caudal crest of
Galeus piperatus are larger than the central ones; the crest is structurally
similar to crests in other members of the genus, but it is proportionally some-
what smaller. In comparison with specimens of equal size, the marginal crest
denticles of G. piperatus are one-half as long as those in G. arae. The crest of
G. piperatus differs from the crest of G. sauteri and G. eastmani in being nar-
rower and having fewer denticles between the laterals.
The lining of the mouth is black or dusky in the type series of Galeus
piperatus, whereas it is white or light colored in Parmaturus xaniurus and also
in the preserved specimens of G. sauteri and G. eastmani. In some of the
specimens of G. melastomus that have been in preservative for a long time the
dark color of the inside of the mouth is entirely leached away; probably the
color of the mouth lining would not always be reliable in distinguishing pre-
served specimens. It may be a useful field character in the eastern tropical
Pacific for quick separation of Galeus and Parmaturus. Apparently the darker
color of the mouth lining characterizes fresh specimens of all the Atlantic
species of Galeus as well as G. piperatus. The mouth lining in G. murinus was
described by Collett (1904) as blackish and in G. jenseni it was described by
Saemundsson (1922) as dark violet.
The holotype of Galeus piperatus is grayish brown with the dorsal surfaces
only slightly darker than the ventral surfaces and with almost no trace of
markings. The paratypes, which are slightly smaller, show an indistinct and
irregular arrangement of darker brown blotches dorsally, and the dorsal sur-
faces are appreciably darker. Very small and numerous black spots just large
enough to be visible to the naked eye (0.1 mm.; 0.3 mm. when expanded) are
scattered generally over the skin of all of our specimens; they are especially
notable on the lower surfaces. Small, scattered spots of pigment are also found
to some extent on Galeus eastmani and G. sauteri, but they are not so numer-
ous or widely distributed. Pepper-like spots can be found on all members of
the genus Galeus, but are more noticeable to the naked eye on G. piperatus.
Unlike other species of Galeus that we have examined, G. piperatus has nar-
row, translucent bands 1. 0-2.0 mm. wide without denticles on the trailing edges
of the two dorsal fins; G. cadenati has wide colorless bands on the trailing
edges of the dorsal fins, but these borders are covered with denticles and are
not of uniform width.
Four measurements, expressed as percent of total length, indicate differ-
Vertebral characters in seven species of Galeus from x-ray examination. Data
for additional specimens of G. arae are given by Victor G. Springer and
J.A.F. Garrick (1964).
1966
New Shark of the Genus GALEUS
1
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Contributions in Science
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ences in body proportions between G. piperatus and two other Pacific species:
G. eastmani and G. sauteri. G. piperatus , with respect to these body propor-
tions, appears to be transitional between the Atlantic and the western Pacific
species, but tends to resemble more closely Galeus cadenati of the Atlantic.
The distance between dorsal fin bases, for example, is 10.9-12.3% of total
length in G. piperatus and 15.4% in G. eastmani (it is 13.1-13.7% in G.
sauteri and 12.3-14.7% in the Atlantic species, G. cadenati ). The distance be-
tween the anal-fin tip and the caudal is much less in G. piperatus than in G.
eastmani ( 1. 7-2.7% as compared with 5.6% ) . This same distance in G. sauteri
is 0.6-2.7%, putting G. piperatus close to G. sauteri with respect to this one
measurement. The width of the mouth is 8. 3-9. 2% of the total length in G.
piperatus as compared with 6.2% in G. eastmani and 6.0-7.0% in G. sauteri.
(In G. cadenati it is 6.6-9. 1% of total length). G. piperatus is not so slender
as G. eastmani and G. sauteri, and is more like the Atlantic species (trunk
height of the two adults of G. piperatus measured at the pelvic fin is 9.2 and
10.9% of total length; similar measurement of G. eastmani is 7.4%, of G.
sauteri 7. 5-8. 9% ).
From the morphometric differences between G. piperatus and two other
Pacific species, we can say that the new species has a shorter interdorsal
distance, a wider mouth, and is shorter and heavier. A shorter anal-to-caudal
fin distance also separates G. piperatus from G. eastmani.
Natural history notes: The holotype (302 mm.) and the larger paratype
(296 mm.) each have a few large eggs about 7 mm. in diameter in the func-
tional right ovary, and are considered sexually mature. The ovary of the
smaller paratype (256 mm.), in gross examination, is much smaller and shows
no indication of egg formation. This evidence suggests that Galeus piperatus
reaches maturity at about 30 cm. or less. G. piperatus is thus one of the
smaller species of the genus, only a little larger than G. arae and about the
same length when mature as G. cadenati. Two species of Galeus (G. melas-
tomus and G. hertwigi ) are reported to reach lengths of more than 60 cm.
Whether G. piperatus lays eggs in egg cases or holds the embryos in the
oviducts for development cannot be determined from the specimens examined.
Within the genus Galeus it is known that G. melastomus lays eggs in leathery
cases and G. polli bears living young (Cadenat, 1959).
Literature cited
Cadenat, J.
1959. Notes d’ichtyologie ouest-africaine. XX. Galeus polli espece nouvelle
ovovivipare de Scylliorhinidae. Bulletin de l’lnstitut Francais d’Afrique
Noire, 21(1) :395-409.
Collett, R.
1904. Four hitherto undescribed fishes from the depths south of the Faroe
Islands. Christiania Videnskabs-Selskabs Forhandlinger for 1904, No. 9,
1-7.
1966
New Shark of the Genus GALEUS
9
Engelhardt, Robert von
1912. tiber einige neue Selachier-Formen. Zoologischer Anzeiger 39(21/22) :
643-648.
Garman, Samuel
1906. New Plagiostomia. Bull. Mus. of Comp. Zool., Harvard College,
46(11) .*201-208.
Gilbert, C. H.
1892. Descriptions of thirty-four new species of fishes collected in 1888 and
1889, principally among the Santa Barbara islands and in the Gulf of
California. Proc. U.S. Natl. Mus., 14:539-566.
Goode, George Brown, and Tarleton A. Bean
1896. Oceanic ichthyology. Special Bulletin, U.S. Natl. Mus. (1895): 529 p.,
123 pis.
Jordan, David Starr, and Robert Earl Richardson
1909. A catalogue of the fishes of the island of Formosa, or Taiwan, based on
collections of Dr. Hans Sauter. Mem. Carnegie Mus., 4(4) : 159-204,
1 1 pis.
Jordan, David Starr, and John Otterbein Snyder
1904. On a collection of fishes made by Mr. Alan Owston in the deep waters
of Japan. Smithsonian Misc. Coll., 45(1447) :230-240, 5 pis.
Lavenberg, Robert J., and John E. Fitch
1966. Annotated list of fishes collected by midwater trawl in the Gulf of Cali-
fornia, March-April 1964. California Fish and Game, 52(2) :92-l 10.
Nichols, John Treadwell
1927. A new shark from the continental slope off Florida. Amer. Mus.
Novitates, (256): 1-2.
Rafinesque, Constantine Samuel
1810. Caratteri di alcuni nuovi generi e nuove specie di animali e piante della
Sicilia, con varie osservazioni sopra i medisimi. Palermo, 1810, p. 1-3,
1-105, 20 pis.
Springer, Stewart
1966. A review of western Atlantic catsharks, Scyliorhinidae, with descrip-
tions of a new genus and five new species. U.S. Fish and Wildlife Serv-
ice, Fishery Bull., 65(3) :581-624.
Springer, Victor G., and J. A. F. Garrick
1964. A survey of vertebral numbers in sharks. Proc. U.S. Natl. Museum,
1 16(3496) :73-96.
Whitley, Gilbert P.
1928. Studies in Ichthyology. No. 2. Rec. Australian Mus., 16(4) :21 1-239.
LOS
ANGELES
COUNTY
MUSEUM
Number 111
CONTRIBUTIONS
IN SCIENCE
November 9, 1966
A NEW SUBSPECIES OF THE AZTEC MASTIFF BAT,
MOLOSSUS AZTECUS SAUSSURE, FROM
SOUTHERN MEXICO
By Alfred L. Gardner
Los Angeles County Museum of Natural History
Los Angeles, California 90007
Exposition Park
CONTRIBUTIONS IN SCIENCE is a series of miscellaneous technical papers
in the fields of Biology, Geology and Anthropology, published at irregular intervals
by the Los Angeles County Museum of Natural History. Issues are numbered sepa-
rately, and numbers run consecutively regardless of subject matter. Number 1 was
issued January 23, 1957. The series is available to scientific institutions on an ex-
change basis. Copies may also be purchased at a nominal price.
INSTRUCTIONS FOR AUTHORS
Manuscripts for the LOS ANGELES COUNTY MUSEUM CONTRIBU-
TIONS IN SCIENCE may be in any field of Life or Earth Sciences. Acceptance of
papers will be determined by the amount and character of new information and the
form in which it is presented. Priority will be given to manuscripts by staff members,
or to papers dealing largely with specimens in the Museum’s collections. Manuscripts
must conform to CONTRIBUTIONS style and will be examined for suitability by
an Editorial Committee. They may also be subject to critical review by competent
specialists.
MANUSCRIPT FORM.— (1) The 1960 AIBS Style Manual for Biological
Journals is highly recommended as a guide. (2) Typewrite material, using double
spacing throughout and leaving ample margins, on only one side of 8V2 x 11 inch
standard weight paper. (3) Place tables on separate pages. (4) Footnotes should be
avoided if possible. (5) Legends for figures and unavoidable footnotes should be
typed on separate sheets. Several of one kind may be placed on a sheet. (6) Method
of literature citation must conform to CONTRIBUTIONS style — see number 90 and
later issues. Spell out in full the title of non-English serials and places of publication.
(7) A factual summary is recommended for longer papers. (8) A brief abstract must
be included for all papers. This will be published at the head of each paper.
ILLUSTRATIONS. — All illustrations, including maps and photographs, should
be referred to as “figures!’ All illustrations should be of sufficient clarity and in the
proper proportions for reduction to CONTRIBUTIONS page size. Permanent ink
should be used in making line drawings and in lettering (do not type on drawings);
photographs should be glossy prints of good contrast. Original illustrations will not
be returned unless specifically requested when the manuscript is first submitted.
PROOF. — Author will be sent galley proof which should be corrected and re-
turned promptly. Changes after the paper is in galley will be billed to the author. Un-
less specially requested, page proof will not be sent to the author. 100 copies of each
paper will be given free to a single author or divided equally among multiple authors.
Orders for additional copies should be sent to the Editor at the time corrected galley
proof is returned; appropriate forms for this will be included when galley is sent.
David K. Caldwell
Editor
A NEW SUBSPECIES OF THE AZTEC MASTIFF BAT,
MOLOSSUS AZTECUS SAUSSURE, FROM
SOUTHERN MEXICO
By Alfred L. Gardner1
Abstract: A new subspecies of the Aztec mastiff bat is
described from Chiapas, Mexico. Included also are additional
records of the species in western Mexico and the first report of
another species of mastiff bat for the Republic of Mexico. The
latter closely resembles Molossus pygmaeus Miller, and was
earlier confused with Molossus aztecus Saussure.
Since Saussure’s original description of Molossus aztecus from Ameca-
meca, Mexico, Mexico, in 1860, only four reports of additional specimens
from Mexico of this small mastiff bat have been published. Allen (1906: 260)
reported two specimens from Los Masos, Jalisco, under the name M. obscurus
Geoffroy. Miller (1913: 91) reported having examined specimens of M.
aztecus from Huehuetan, Chiapas. Later, Dalquest (1953: 68) reported five
specimens from Rio Verde, San Luis Potosi. Finally, Alvarez and Avina
(1964: 250) reported three specimens, one from each of the following locali-
ties: Juchitan, Oaxaca; Pueblo Juarez, Colima: 3 km. north of El Limon,
Tamaulipas.
Recent collecting activities in western and southern Mexico and the
examination of comparative material indicate the presence of a heretofore
undescribed subspecies of M. aztecus that may be known as :
Molossus aztecus Iambi, new subspecies
Holotype : An adult male, skin with skull, Los Angeles County Museum
of Natural History No. 27001 (UA 9525), collected by A. L. Gardner (origi-
nal No. 5139) from 11 km. northwest from Escuintla, Chiapas, Mexico, ca.
100 feet elevation, 18 November, 1962.
Allotype : An adult female, skin with skull, LACM No. 27002 (UA
9519), collected by A. L. Gardner (original No. 5053) from 11 km. northwest
from Escuintla, Chiapas, Mexico, ca. 100 feet elevation, 15 November, 1962.
Diagnosis'. Size, small for species; two color phases present, dark phase
blackish, approaching Fuscous and the light phase approaching Sudan Brown
(capitalized color terms after Ridgway, 1912) ; light basal portion of bicolored
fur short; flight membranes and ears black.
Description : Both the holotype and the allotype are in the dark color
phase. Measurements in millimeters of the holotype followed by measurements
of the allotype: Total length, 98, 92; tail, 36, 33; hind foot, 9, 8; ear from
department of Zoology, University of Arizona. (Present address: Museum of
Zoology, Louisiana State University, Baton Rouge, Louisiana 70803.)
1
Molossus coibensis $ * X 34.91 16.58 14.32 3.78 10.55 8.67 7.54 6.11
Panama R 33.5-36.5 16.1-17.1 13.7-14.6 3.6-3.9 10.3-10.9 8.3-8.9 7.3-7.9 5. 8-6.3
Contributions in Science
No. 1 1 1
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Taxon and Locality
Sex and Age
Forearm Length
Greatest Length of
Skull (inc. incisors)
Condylobasal Length
(not inc. incisors)
Interorbital
Constriction
Zygomatic Breadth
Breadth of Brain Case
Breadth across M2-M2
Length Maxillary
Tooth Row
Selected measurements in millimeters of Molossus a. Iambi, Molossus a. aztecus, and Molossus cf. pygmaeus from Mexico and Molossus
coibensis from Panama. The mean (X), range (R), and number (N) are given for each character. Samples include adults only (ad), or
adults and subadults (*).
1966
New Subspecies of Mastiff Bat
3
notch, 15, 14; tragus, 5, 4; forearm, 36.50, 35.15; weight, 17 gr., 15 gr.;
greatest length of skull, 18.25, 17.20; condylobasal length, 15.40, 15.00; inter-
orbital constriction, 4.00, 3.90; zygomatic breadth, 11.10, 10.85; mastoid
breadth, 1 1.20, 10.65; breadth of brain case, 9.10, 9.00; depth of brain case,
6.25, 6.10; breadth across M2-M2, 7.40, 7.50; breadth across canines, 4.50,
4.10; length of maxillary tooth row, 6.20, 6.10; length of mandible, 12.50,
12.20; mandibular tooth row, 6.85, 6.60.
Paratypes : Eleven additional specimens from the type locality are desig-
nated paratypes. The males (UA Nos.) are: 9517 ad.; 9520 juv.; 9522 ad.;
9524 ad.; 9527 ad.; 9528 subad.; 9529 juv. The females (UA Nos.) are: 9518
ad.; 9523 subad.; 9526 subad.; 9530 ad. All specimens except UA 9526 are in
the dark color phase; UA 9526 is in the light color phase.
Specimens Examined : Specimens examined and referred to M. a. Iambi
are from the following localities: 13 mi. SW Las Cruces (KU, 2); 15 mi. SW
Las Cruces (KU, 1); 12 mi. E Ortiz Rubio on Villa Elores Rd. (UA, 1); 20
km. SE Pijijiapan (LACM, 4; UA, 1); 11 mi. NW Escuintla, ca. 100 ft.
(LACM, 2; UA, 11).
Range: Presently known from southwestern Chiapas and the Pacific
slope of the Sierra Madre del Sur de Chiapas, Mexico.
Comparisons : This bat differs from the Mexican Plateau form, Molossus
aztecus aztecus, in generally smaller size, darker color, two color phases, and
lack of smoky-white wash over fur, especially ventrally. In these respects
Iambi compares favorably with M. coibensis Allen of Panama, a much smaller
bat (see Table 1 and Remarks). M. a. Iambi was compared with seven speci-
mens of a small Molossus , three from Puerto Madero and four from Hue-
huetan, Chiapas. Huehuetan is on the railroad about 18 miles airline to the
southeast from the type locality of Iambi and Puerto Madero is about 20 miles
airline farther south on the Pacific Coast of southern Chiapas. The specimens
from Huehuetan and Puerto Madero differ markedly from M. a. Iambi in fur
quality and color pattern: longer and thinner than in M. a. Iambi but approach-
ing the lighter color of the more northern highland populations of M. a. aztecus.
In general, they differ from Molossus aztecus in longer forearm; smaller, nar-
rower skull; narrower, lower rostrum; infraorbital foramen situated lower,
close to maxillary tooth row; sub-posterior border of infraorbital foramen not
expanded and expansion not continuous with maxillary portion of zygomatic
arch; palate narrow and not broadly domed; separation between sphenoidal
pits narrower. See Table 1 for comparative measurements.
Remarks’. In basic cranial proportion and configuration plus color pat-
tern, M. a. Iambi is very similar to the much smaller M. coibensis. Future work
may reveal that M. a. Iambi represents a northern intermediate between
M. a. aztecus and M. coibensis, in which case M. coibensis Allen (1904:227)
would be a synonym of M. aztecus Saussure (1860) .
The four specimens from Huehuetan, previously reported by Miller
(1913:91) as M. aztecus (a female adult, FMNH 44254; a male adult, USNM
4
Contributions in Science
No. 1 1 1
77661; two adult females, USNM 77662 and 77663) and the three specimens
from Puerto Madero (a subadult male, KU 68780; a juvenile female and an
adult female KU 68781 and 68782 respectively) all compare favorably with a
northern South American form closely resembling the description of Molossus
pygmaeus Miller ( 1900:162) and to which, at this time reserving a re-examina-
tion of their identity, I tentatively assign them.
The age classification referred to in this work is based upon the ossification
of epiphyseal cartilages and cranial sutures. Specimens are classed as juvenile
if the epiphyseal cartilages of the digital elements of the wing are not com-
pletely ossified. This is usually readily visible through the skin in the prepared
specimen. This lack of ossification coincides with the unossified sutures be-
tween the presphenoid and basisphenoid, and between the basisphenoid and
basioccipital bones of the skull. Specimens are designated subadult if the epi-
physeal plates of the wing elements appear ossified but the suture between the
basisphenoid and basioccipital bones is still open or incompletely ossified in
the midline of the skull. Adults are those which have no prominent unossified
sutures in the skull.
I collected nineteen of the 22 specimens of M. a. Iambi in mist nets placed
across pools or over streams at road crossings. Late one afternoon, in camp at
the type locality, I noticed several bats fly out of a hole, high in a tall fig tree
and swoop down over the hill toward the river. One of the bats, which proved
to be M. a. Iambi, became enmeshed in a mist net already in place at the edge
of a deep pool of water in a gravel pit left by bridge construction workers.
Three of five females collected at 20 mi. SW Pijijiapan between 13 March and
27 March, 1961, were pregnant, each with a single embryo (crown-rump meas-
urement of one, 1 1 mm.) and two contained no embryos. Another female col-
lected 12 mi. E Ortiz Rubio on Villa Flores Rd., 26 June, 1964, contained one
embryo measuring 14.1 mm. crown-rump. Males collected at the type locality,
between 13 and 18 November, 1962, showed some sign of sexual activity.
Three males had enlarged testes measuring 2 mm. x 5 mm., 3 mm. x 5 mm., and
4 mm. x 6 mm. respectively. Three additional males had testes not enlarged.
Two of the 22 specimens at hand show the light color phase. Two others
show a partial mixing of the color phases; both, apparently in the lighter phase,
are molting and the new fur is dark.
Molossus aztecus Iambi is named in honor of the late Chester C. Lamb,
a veteran field collector and naturalist who was very well acquainted with the
Mexican state of Chiapas.
Additional Specimens Examined : Comparative material was kindly made
available by the American Museum of Natural History (AMNH), the Field
Museum of Natural History (FMNH), the Museum of Natural History, Uni-
versity of Kansas (KU), the Los Angeles County Museum of Natural History
(LACM), and the United States National Museum (USNM). Specimens in
the mammal collection, Department of Zoology, University of Arizona, are
indicated by UA. Molossus aztecus aztecus from Colima: Tlapeixtes, 4 km.
1966
New Subspecies of Mastiff Bat
5
ENE Manzanillo (AMNH, 2). Jalisco: Pena Colorada, Rio de Talpa, ca.
10 km. N Talpa de Allende (UA, 2); Rancho de los Ocotes, ca. 6 km. N
Talpa de Allende (UA, 1 ) ; Rio de Talpa, Talpa de Allende (UA, 6) ; Los Masos
(AMNH, 2). Oaxaca: 5 mi. N Juchatengo, ca. 4000 ft. (UA, 2); 3 mi. S
Nejapa (KU, 1); Tehuantepec (AMNH, 1). Sinaloa: Alisos, ca. 50 km. NNE
(by Rd.) from Badiraguato (UA, 1). Molossus cf. pygmaeus from Chiapas:
Huehuetan (FMNH, 1; USNM, 3); 1 mi. SE Puerto Madero, (KU, 3).
Acknowledgments
I am indebted to the various institutions which made comparative material
available, and to Dr. E. L. Cockrum of the University of Arizona, under whose
direction this description was prepared.
I wish to express appreciation to Dr. Hernandes Corzo, Prof. Jose A. Davila
C., and the late Sehor Ing. Luis Macias A. of the Direccion General de Forestal
y de Caza, Mexico, D.F., under whose auspices my collecting permits were
obtained.
Literature Cited
Allen, J. A.
1904. New bats from Tropical America, with note on species of Otopterus.
Bull. Amer. Mus. Nat. Hist., 20 : 227-237.
1906. Mammals from the states of Sinaloa and Jalisco, Mexico, collected by
J. H. Batty during 1904 and 1905. Bull. Amer. Mus. Nat. Hist., 22:
191-262.
Alvarez, T. and C. E. Avina
1964. Nuevos registros en Mexico de la familia Molossidae. Revista de la
Sociedad Mexicana de Historia Natural, Mexico, 25:243-254.
Dalquest, W. W.
1953. Mammals of the Mexican state of San Luis Potosi. Louisiana State
Univ. Studies, Biol. Ser., 1:1-229.
Miller, G. S., Jr.
1900. A second collection of bats from the Island of Curacao. Proc. Biol.
Soc. Wash., 13:159-162.
1913. Notes on the bats of the genus Molossus. Proc. U.S. Natl. Mus., 46:
85-92.
Ridgway, R.
1912. Color Standards and Color Nomenclature. Washington, D.C.
LOS
ANGELES
COUNTY
MUSEUM
CONTRIBUTIONS
^ IN SCIENCE
UMBER 112
October 27, 1966
THE TAXONOMY AND NOMENCLATURE OF SOME
NORTH AMERICAN BEES OF THE GENUS CENTR1S
WITH DESCRIPTIONS OF NEW SPECIES
( Hymenoptera : Anthophoridae)
By Roy R. Snelling
Los Angeles County Museum of Natural History • Exposition Park
Los Angeles, California 90007
CONTRIBUTIONS IN SCIENCE is a series of miscellaneous technical papers
in the fields of Biology, Geology and Anthropology, published at irregular intervals
by the Los Angeles County Museum of Natural History. Issues are numbered sepa-
rately, and numbers run consecutively regardless of subject matter. Number 1 was
issued January 23, 1957. The series is available to scientific institutions on an ex-
change basis. Copies may also be purchased at a nominal price.
INSTRUCTIONS FOR AUTHORS
Manuscripts for the LOS ANGELES COUNTY MUSEUM CONTRIBU-
TIONS IN SCIENCE may be in any field of Life or Earth Sciences. Acceptance of
papers will be determined by the amount and character of new information and the
form in which it is presented. Priority will be given to manuscripts by staff members,
or to papers dealing largely with specimens in the Museum’s collections. Manuscripts
must conform to CONTRIBUTIONS style and will be examined for suitability by
an Editorial Committee. They may also be subject to critical review by competent
specialists.
MANUSCRIPT FORM.— (1) The 1960 AIBS Style Manual for Biological
Journals is highly recommended as a guide. (2) Typewrite material, using double
spacing throughout and leaving ample margins, on only one side of 8V2 x 11 inch
standard weight paper. (3) Place tables on separate pages. (4) Footnotes should be
avoided if possible. (5) Legends for figures and unavoidable footnotes should be
typed on separate sheets. Several of one kind may be placed on a sheet. (6) Method
of literature citation must conform to CONTRIBUTIONS style — see number 90 and
later issues. Spell out in full the title of non-English serials and places of publication.
(7) A factual summary is recommended for longer papers. (8) A brief abstract must
be included for all papers. This will be published at the head of each paper.
ILLUSTRATIONS. — All illustrations, including maps and photographs, should
be referred to as “figures!’ All illustrations should be of sufficient clarity and in the
proper proportions for reduction to CONTRIBUTIONS page size. Permanent ink
should be used in making line drawings and in lettering (do not type on drawings);
photographs should be glossy prints of good contrast. Original illustrations will not
be returned unless specifically requested when the manuscript is first submitted.
PROOF. — Author will be sent galley proof which should be corrected and re-
turned promptly. Changes after the paper is in galley will be billed to the author. Un-
less specially requested, page proof will not be sent to the author. 100 copies of each
paper will be given free to a single author or divided equally among multiple authors.
Orders for additional copies should be sent to the Editor at the time corrected galley
proof is returned; appropriate forms for this will be included when galley is sent.
David K. Caldwell
Editor
THE TAXONOMY AND NOMENCLATURE OF SOME
NORTH AMERICAN BEES OF THE GENUS CENTRIS
WITH DESCRIPTIONS OF NEW SPECIES
(Hymenoptera: Anthophoridae)
By Roy R. Snelling1
Abstract: In the current paper representatives of six sub-
genera are considered. Trichocentris : The female characteristics
of this subgenus are given, based upon Centris vanduzeei, which is
redescribed. Paracentris : C. pallida, C. tiburonensis, C. Cali-
fornia, C. hoffmanseggiae, C. subhyalina and C. mexicana
are recorded from new areas. The males of C. californica and
C. subhyalina are fully described and C. birkmanii is synony-
mized with the latter. Previously considered by me to be a syno-
nym of C. lanosa, C. lanosa resoluta is reinstated as a subspecies.
The following new species are described: C. angustifrons
(United States), C. zacateca (Mexico, United States), C. an-
thracina (Mexico to Ecuador) and C. laevibullata (Mexico). A
key to North American species is given. Centris s. str .: C. inermis
pallidifrons is synonymized under C. i. gualanensis. C. eisenii, C.
decolorata and C. flavofasciata are recorded from the United
States for the first time. Xanthemisia : C. aethiops is assigned to
this subgenus and C. armillatus formally synonymized with it.
C. carolae is described as a new species from Mexico. Hemi-
siella : C. lanipes subtar sata is considered a subspecies of C.
trigonoides, and C. transversa is recorded from the United States.
Melanocentris : three aberrant species are described as new:
C. strawi (Mexico), C. ruthannae (United States) and C. anom-
ala (Mexico).
During the course of my preliminary studies on these bees, many new
and significant distributional records have been discovered. The primary
objective of this paper is to make these data available for the forthcoming
Second Supplement to the Catalog of Hymenoptera of America north of
Mexico. The opportunity is taken, also, to present the descriptions of several
new species as well as observations on the taxonomy and nomenclature of
certain other species.
Although some of the material recorded below is from my personal col-
lection (now in the Los Angeles County Museum of Natural History, and
indicated by LACM), the bulk of it has been made available from the collec-
tions of several institutions. For their generosity in making these specimens
available to me, I wish to express my deep gratitude to the following: W. L.
Brown, Jr., Museum of Comparative Zoology (MCZ); G. D. Butler, Jr., Uni-
versity of Arizona (UA); H. R. Dietrich, Cornell University (CU); P. D.
Hurd, Jr. and J. A. Powell, California Insect Survey, University of California,
Entomology Section, Los Angeles County Museum of Natural History.
1
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Berkeley (CIS); K. V. Krombein, U.S. National Museum (USNM); A. T.
McClay, University of California, Davis (UCD); C. D. Michener, University
of Kansas (KU); T. B. Mitchell, North Carolina State College (NCSC);
J. G. Rozen, American Museum of Natural History (AMNH) ; P. H. Timber-
lake, Citrus Experiment Station, University of California, Riverside (UCR).
I am especialy indebted to J. S. Moure for reviewing the manuscript and mak-
ing certain corrections based on his own extensive knowledge of these bees,
much of which is still unpublished.
Subgenus TR1CHOCENTRIS Snelling
The present definite assignment of Centris vanduzeei Cockerell to this
subgenus has made it possible to provide a diagnosis of the female of Tricho-
centris.
Probably the most noticeable and interesting features of the females are
the primitive nature of the basitibial plates and the complete absence of
modified setae on the front and middle tibiae. These two characters, coupled
with the sparse scopa and pubescent abdomen seem to confirm the view that
this is a primitive group.
The following combination of characters should aid in recognizing the
females of Trichocentris : Mandibles long, slender, quadridentate; first flagellar
segment longer than scape; front and middle basitarsi lacking modified setae;
basitibial plate with poorly developed secondary plate; scopal hairs con-
spicuously shorter and sparser than in Paracentris; abdominal tergites with
long, erect pubescence, not concealing surface; apices of tergites with distinct
pubescent fasciae.
Centris morsei Cockerell, provisionally assigned by me (Snelling, 1956:
3) to Trichocentris should be placed in Paracentris since it shows undeniable
relationships with C. ( Paracentris ) caesalpiniae Cockerell.
Centris ( Trichocentris ) vanduzeei Cockerell
Centris vanduzeei Cockerell, 1923. Proc. Calif. Acad. Sci. (4) 12:75-76.
$ $. Snelling, 1956. Pan-Pacific Ent. 32:4. $ $.
Recently I was able to critically examine a cotype female and a male of
this species, and confirmed my earlier conjecture (Snelling, 1956:4) that C.
vanduzeei should be assigned to this subgenus.
Since Cockerell failed to designate an allotype when he described this
species, I am taking this opportunity to do so. I have selected and designated
as Lectoallotype an original cotype male from San Jose Island, Gulf of Cali-
fornia, Mexico, May 28, 1921 (E. P. Van Duzee) . This male has been returned
to the California Academy of Sciences. In his original description Cockerell
(1923:75) remarked that the pubescence of the female labrum was brownish,
but it is whitish and therefore concolorous with the rest of the facial pubescence.
The following redescription is based upon these cotypes and is intended
to give a better picture of the many peculiarities of this species.
1966
The Taxonomy and Nomenclature of Bees
3
FEMALE: Integument of head, thorax, legs and abdomen black; all basi-
tarsi and hind tibiae dark ferruginous; tibial spurs, medio- and distitarsi, and
tarsal claws light ferruginous; tegulae ferruginous; wings hyaline, yellow-
tinged, veins and stigma ferruginous. Pubescence of head, thorax, legs and ab-
domen pale whitish; that of thoracic dorsum tinged pale fulvous; anterior tarsi,
apical fourth of middle tibiae, middle tarsi, scopa of hind tibiae and basitarsi,
apical tarsal segment and broad apical band on fifth tergite with bright orange-
fulvous pubescence. Abdominal tergites with abundant erect, whitish pubes-
cence (not obscuring surface), forming distinct apical fasciae on first four
segments; ventrites II to IV with whitish apical fasciae; discs of ventrites I to
III, and VI, with long erect fulvous hairs, IV and V, with whitish hairs on discs.
Head : Broader than long, mandibles long, slender, the two inner teeth
much smaller than the outer, and on a plane at right angles to that of the outer
teeth. Maxillary palpi five-segmented, third segment longest, penultimate seg-
ment longer than basal, but shorter than apical. Labrum rugosely punctate,
with shining interstices between punctures; apex broadly pointed. Clypeus
slightly protuberant, apical middle slightly flattened, apex evenly, broadly con-
cave; shiny, with broad median area tapering slightly toward apex with a few
scattered punctures, lateral areas with punctures rather coarse, close. Frons
rather finely, sparsely punctate, with impunctate areas before anterior ocellus
and laterad of posterior ocelli; vertex and genae sparsely punctate, the punc-
tures a little finer than those of frons. First flagellar segment longer than scape,
slightly longer than following five segments combined. Eyes converging slightly
above. Distance between posterior ocelli greater than distance between ocelli
and eyes; distance from ocelli to posterior margin of vertex greater than dis-
tance between posterior ocelli. Clypeus half as long as lower transfacial dis-
tance (at level of extreme lateral angle of clypeus).
Thorax : Mesoscutum rather coarsely, closely punctate, the punctures
about one-half times their diameters apart; meso- and metapleura a little more
finely punctate, with the punctures about their own diameters apart; meso-
scutellum with punctures equal to those of mesoscutum, becoming distinctly
closer apically until they are almost touching; metanotum impunctate, indis-
tinctly tessellate; basal area of propodeum shiny, impunctate; lateral areas
finely, sparsely punctate. Anterior margin of third submarginal cell about one-
third as long as posterior margin. Basitibial plate elongate, with poorly differen-
tiated secondary plate, with longitudinal median depression; scopal pubescence
rather short and sparse, not forming heavy compact mass, not concealing sur-
face of tibiae and basitarsi; distitarsi teardrop shaped, one-third as broad as
long, shorter than preceding segments combined.
Abdomen : With fine, piliferous punctures on discs of all segments;
pubescence forming distinct, white, apical fasciae on tergites one to five.
Measurements : Body length (front of vertex to apex of second tergite),
10.0 mm.; forewing length, 9.5 mm.
MALE: Integument black, except all tarsal segments, tibial spurs, and
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apical half of tegulae. Tibial spurs and all tarsi light ferruginous, the tarsal
claws fuscous. Apical half of tegulae yellowish. Pubescence mostly pale whit-
ish; that of tarsi and seventh tergite and ventrite fulvous. Wings lighter than in
female, veins and stigma ferruginous. Pubescence very long and dense on face
and abdomen, erect on the latter.
Head : Broader than long. Mandibles long, slender, tridentate; inner tooth
smallest and on a different plane than the two outer teeth. Labrum moderately
shiny, finely, closely punctate, the punctures almost touching; the surface
obscured by long dense pubescence; apex broadly pointed. Clypeus strongly
protuberant, slightly flattened on each side of median line; shiny, with a median
longitudinal impunctate area, the lateral punctures a little coarser than those
of labrum, more widely spaced, with shining interstices. Frons shiny, finely
punctate, the punctures a little closer than those of clypeus; an impunctate
area in front of anterior ocellus and posterior ocelli. Vertex finely and closely
punctate, the punctures about their own diameters apart, except posteriorly
where they become a little less crowded. Genae finely punctate, the punctures
about their own diameters apart, except posteriorly where they become a little
more crowded. Antennal scape short, about half as long as first flagellar seg-
ment, which is slightly shorter than the following five segments combined.
Distance from lower margin of anterior ocellus to base of clypeus about two-
thirds the breadth of face at level of base of clypeus; distance between antennal
sockets much greater than distance between sockets and eyes; distance between
posterior ocelli much greater than distance between sockets and eyes; distance
between posterior ocelli much greater than distance between ocelli and eyes,
the ocelli about twice an ocellar diameter apart and removed from eyes by
about one-third an ocellar diameter; distance between posterior ocelli about
two-thirds distance from ocelli to posterior margin of vertex. Vertex slightly
concave between tops of eyes as seen from front.
Thorax : Mesoscutum and mesoscutellum dull, closely, finely, punctate,
the punctures separated by less than a puncture diameter. Mesopleura finely,
sparsely punctate anteriorly, the punctures becoming coarser, closer posteri-
orly. Metanotum shinier than mesoscutum, impunctate tessellate. Entire pro-
podeum shiny, finely, sparsely punctate, the punctures for the most part sepa-
rated by two or more times their diameters. Anterior half of tegulae finely,
closely punctate, the posterior half with punctures much finer and somewhat
sparser. Front and middle distitarsi as long as mediotarsi. Posterior femora
strongly swollen, about half as broad as long. Wings as in female.
Abdomen : Dull, all segments finely, closely, piliferously punctate. Sixth
tergite with a dense brush of long pubescence on each side of the median line,
concealing seventh which has a small patch of short pubescence on each side,
and its apex slightly emarginate.
Measurements’. Body length (front of vertex to apex of second tergite),
1 1 .5 mm.; forewing length, 1 1 .3 mm.
The only additional male known is from Ruffo Ranch, Isla Ceralbo, Gulf
1966
The Taxonomy and Nomenclature of Bees
5
of California, Mexico, March 22, 1953 (P. H. Arnaud; Sefton Orca Expedition
to Gulf of California; LACM) . Many maps show this as “Isla Cerralvo!’
Subgenus PARACENTRIS Cameron
Padre J. S. Moure has informed me ( personal communication ) that the
correct name for this subgenus, formerly known as Penthemisia Moure, is
Paracentris Cameron (1903:235; type species: Paracentris fulvohirta Cam-
eron). He has examined Cameron’s type and found it to be close to C. ( Pen-
themisia) bucholzi Friese.
Centris ( Paracentris ) pallida Fox
Centris pallida Fox, 1899. Proc. Acad. Nat. Sci. Phila. 51:66 9 . Snelling,
1956. Pan-Pacific Ent. 31:6.$$
Centris pallida callognatha Cockerell, 1923. Proc. Calif. Acad. Sci. (4)
12:78. 9 . Synonymy of Snelling, 1956. Op. cit. supra.
Centris trichosoma Cockerell, 1923. Proc. Calif. Acad. Sci. (4) 12:78.
$ . Synonymy of Snelling, 1956. Op. cit. supra.
Dr. P. D. Hurd provided a single female of this species collected on Santa
Catalina Island, Los Angeles County, CALIFORNIA, June 7-9, 1954, by R.
Ryckman, R. Lee and C. Christianson (CIS). This seems at first an unusual
record, but when one considers that a number of anthophorids and other bees,
common to the desert regions on the mainland, occur there also, it is not too
surprising. The fact that the species has not been collected there before may
be due, in part, to the natural swiftness and alertness of these bees, coupled
with the fact that this species probably is not common on the Island.
Four males from Arizona are noteworthy in that the hind femora are
greatly swollen, being one-half as broad as long in the more extreme cases. This
condition is superficially similar to the subgenus Trichocentris. Because of this
femoral development, and certain differences in punctation and the hidden
ventrites, this form at first was regarded as a new species. However it is now
evident that these variations are met within the range of variation of these
characters in C. pallida, although the cline from the typical form, with slender
femora, to the swollen form is not complete.
Centris pallida is an anomalous form of Paracentris, and is very close to
the more primitive Trichocentris, as evidenced by the abundant pale pubes-
cence of the abdomen, four-segmented maxiliary palpi, lack of modified
setae on the anterior basitarsi of the female, and the occasionally swollen hind
femora of the males. However, because of the Paracentris- like basitibial plate
and scopa of the female, and the genitalia of the male, it seems best for the time
being to retain C. pallida in Paracentris. An additional reason for the present
treatment is the relationship with C. tiburonensis Cockerell, a more typical
Paracentris. Separation of these species into different subgenera would serve
only to obscure the relationship between them, and to render more tenuous
the division between the two subgenera.
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No. 112
Centris ( Paracentris ) tiburonensis Cockerell
Centris tiburonensis Cockerell, 1923. Proc. Calif. Acad. Sci. (4) 12:78,
9. Snelling, 1956. Pan-Pacific Ent. 31:6. $ 8 .
A recent collection extends the range of this species to Arizona.
NEW RECORDS. BAJA CALIFORNIA: 2 $ $ Sierra de los Cocopas,
35 kilo. SW Mexicali, July 6, 1953 (R. R. Snelling: LACM) ; on Dalea spinosa.
CALIFORNIA: Imperial Co.: 25 9 9,4 8 8 , 4.5 mi. E. Coyote Wells, June
10, 1956 (R. R. Snelling: LACM), on D. spinosa. Riverside Co.: 1 $ Palm
Springs, June 25, 1941 (E. C. Van Dyke; CAS) ; 2 8 8 , Palm Springs, June 24,
1952 (P. H. Timberlake; UCR), on D. spinosa ; 1 9,2 8 8 , same data (R. H.
and L. D. Beamer, W. LaBerge, A. Wolf, C. Liang, C. Weiner; KU) ; 16 9 9 ,
6 8 8 , Indio, June 7, 1956 (M. Wasbauer; CIS), on D. spinosa', 15 9 9,1
8, 20 mi. E. Indio, June 21, 1958 (C. D. MacNeill; CAS); 1 9, Desert
Center, June 16, 1947 (G. H. and J. L. Sperry: LACM) : 1 9,1 8 , 20 mi. W.
Blythe, June 10, 1956 (M. Wasbauer; CIS), on D. emoryi; 16 9 9,9 8 8,
Shaver’s Well, June 15, 1963 (R. R. Snelling; LACM), on D. spinosa. San
Bernardino Co.: 1 9,3 8 8, 8 mi. SE Needles, June 5, 1938 (P. H. Timber-
lake; UCR), on D. spinosa', 4 9 9,2 8 8,5 mi. W. Essex, June 30, 1952 (R.
H. and L. D. Beamer, W. LaBerge, A. Wolf, C. Liang, C. Weiner; KU); 5
9 9, 10 8 8, Essex, July 2, 1953 (E. S. Ross; CAS), 3 9 9,2 8 8 on Dalea.
ARIZONA: Mojave Co.: 87 9 9,7 mi. NE Topock, June 24, 1959 (R. R. &
M. D. Snelling; LACM), on Dalea.
Centris ( Paracentris ) lanosa resoluta Cockerell
Centris cockerelli resoluta Cockerell, 1923. Proc. Calif. Acad. Sci. (4)
12:76-77. 9 8.
Centris lanosa resoluta, Michener, in Muesebeck, et al., 1951. U. S. D. A.,
Agr. Monog. 2:1241.
Centris lanosa, Snelling, 1956. Pan-Pacific Ent. 32:7 (in part).
In my paper on California Centris (Snelling, 1956:7), the form described
by Cockerell as C. cockerelli resoluta was placed in synonymy with C. lanosa
Cresson, because of the occurrence of certain individuals in Arizona and
Sonora, Mexico, which were similar to typical specimens from Texas and New
Mexico. At the time of my original decision, adequate material from the latter
states was not available. The intermediate specimens are from an overlapping
area in the range of the two subspecies which leads me to regard them now as
hybrids. The range of C. lanosa lanosa (with yellow clypeus in the female) as
now understood includes Texas, New Mexico and Tamaulipas, Nuevo Leon
and Chihuahua, Mexico, while C. /. resoluta (with ferruginous clypeus in the
female) inhabits Arizona, California, Nevada and Sonora, and Baja California,
Mexico. The intermediate specimens are found in western New Mexico and
Chihuahua and eastern Arizona and Sonora in certain areas where the ranges
of the two forms overlap.
1966
The Taxonomy and Nomenclature of Bees
7
Centris ( Paracentris ) calif ornica Timberlake
Centris calif ornica Timberlake, 1940. Pan-Pacific Ent. 16:139. 9. Snel-
ling, 1956. Pan-Pacific Ent. 31:7. $.
Shortly after the proofs for the above paper were returned to the printer,
I secured a single male of this rare species in Kings County, CALIFORNIA.
During subsequent seasons, additional specimens of both sexes were taken in
other areas.
The male of C. californica is very similar to that of C. ho ff manse ggiae
Cockerell. A few slight differences have been noted in the eighth and ninth
ventrites, but these seem to intergrade so completely as to be worthless. How-
ever, the structural differences in the females seem to be great enough to war-
rant separation on the species level. The males appear to be best separated by
the more densely pubescent abdomen with whitish apical fringes on the tergites,
the entirely light pubescence of the apical tergites, and maculate antennal scape
of C. californica. In the following description of the C. californica male, no
structural characters are used, since in all important respects the two species
seem to be identical.
MALE: Black, with creamy-yellow macula as follows: outer side of
mandibles mostly; labrum; clypeus, except extreme upper lateral margins;
triangular supraclypeal mark; linear mark on underside of antennal scape
(greatly reduced in some specimens). Legs, except darkly ferruginous tarsi,
dark rufescent, tarsal claws darkly rufescent, with apices lighter; tibial spurs
light ferruginous. Tegulae lutescent. Wings clear hyaline, veins and stigma fer-
ruginous. All pubescence whitish, except on inner side of tarsal segments where
it is light ferruginous. Tegulae with very short, dense, fine appressed pubescence
in addition to the usual longer, sparser pubescence. Discs of tergites with dis-
tinct whitish apical fasciae of slightly longer denser pubescence; ventrites with
discal pubescence sparser, longer, with distinct apical fasciae on segments two
to five.
Measurements : Body length (front of vertex to apex of second tergite),
9.5 to 12.5 mm.; forewing length, 8.5 to 11.5 mm.
The distribution of this species overlaps that of C. hoff manse ggiae in the
Mojave and Great Basin areas of California. There is, however, a discrete
separation of the two in flight periods and floral visitations; C. hoff manse ggiae
flies from mid-April to mid-June, frequenting chiefly leguminous plants. The
flight period of C. californica apparently extends from late June to early
October, and the species appears to restrict itself to the plants of the family
Capparidaceae, if the “mustard” record of the holotype be disregarded.2
2The probability is strong that the holotype was actually collected on Wislizenia
rather than mustard. The two are somewhat similar in general appearance. Further-
more, the residents of the area around Kerman, Fresno County, commonly refer
to Wislizenia as “alkali mustard” or merely “mustard!’ I have collected specimens
of C. californica in the Kerman area, all on Wislizenia, even though mustard was
available at most sites.
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Contributions in Science
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New records for the distribution of calif ornica are: CALIFORNIA:
Tulare Co.: 1 9,1 $ , 6 mi. N. Alpaugh, Sept. 17, 1959. Kings Co.: 1 $ , 12
mi. S. Flanford, Sept. 28, 1955. Fresno Co.: 10 $ 9, 82 $ $, 5 mi. E. San
Joaquin, Aug. 22, 23, 30, 1960; 4 9 9,8 $ S, Kerman, Aug. 25, 31, 1960;
4 $ $ , 8 mi. W. Kerman, Aug. 22, 1960; 4 9 9,8 $ $, Raisin City, Aug. 24,
1960; 10 9 9, 68 $ $, Helm, Aug. 24, 29, 1960; 1 9, Mendota, Aug. 25,
1960 (all R. R. Snelling; LACM), all on Wislizenia refracta; 4 9 9, 10 $ $,
10 mi. SW. Carruthers, Aug. 31, 1960 (S. M. Fullerton, in the collection of
Mr. Fullerton), on W. refracta. Madera Co.; 2 9 9,9 mi. SW. Madera, Aug.
30, 1960; 6 9 9 , 14 mi. SW. Madera, Sept. 2, 1960 (all R. R. Snelling; LACM)
all on W. refracta. Merced Co.: 1 9, Stevenson, Aug. 31, 1960; 1 9,4 $ S,
4 mi. SE. El Nido, Sept. 4, 6, 1956 (all R. R. Snelling; LACM), all on W.
refracta. Inyo Co.: 3 9 9, Olancha, Sept. 3, 1956 (R. M. Bohart; UCD).
NEVADA: Washoe Co.; 1 $ Southern Pyramid Lake, July 27, 1957 (D. C.
Rentz; LACM). Humboldt Co.: 1 9, Winnemucca, Aug. 30, 1956 (T. R.
Haig; UCD). Pershing Co.: 1 9,3 $ $ , Lovelock, Aug. 30, 1956 (T. R. Haig;
UCD) ; 1 9 , 8 mi. S. Lovelock, Aug. 4, 1956 (T. R. Haig; UCD) .
Centris ( Paracentris ) hoffmanseggiae Cockerell
Centris hoffmanseggiae Cockerell, 1897. Annals and Magazine of Natural
History (6) 19:395. $ (not 9). Snelling, 1956. Pan-Pacific Ent. 38:8. 9 $.
Centris hoffmanseggiae var. davidsoni Cockerell, 1904. Bui. So. Calif.
Acad. Sci. 3:160. $ . Synonymy of Snelling, 1956. Op. cit. supra.
A single female of this species collected at Yuma, ARIZONA, May 4,
1955, on flowers of alfalfa by D. Tuttle (UA), apparently is the first recorded
occurrence of the species on the lower Colorado Desert. An additional Arizona
specimen is from the Santa Rita Range Reserve, April 10, 1957 (A. W. Wood-
row; UA), one male on Cercidium.
Centris ( Paracentris ) subhyalina Fox
Centris subhyalina Fox, 1899. Proc. Acad. Nat. Sci. Phila. 51:69. 9.
Centris birkmanii Friese, 1900. Termeszetrajzi Fiizetek, 23:44. $ 9.
NEW SYNONYMY.
Centris lanosa lanosa, Mitchell, 1962. N. C. Exp. Sta. Tech. Bui. 152:
334-335. 9, $. (misidentification) .
In discussing this species, Fox (1899:69) stated that it might eventually
prove to be the female of C. lanosa, which, however, is now known to be a
very different species only remotely related to C. subhyalina. On the other
hand, C. subhyalina matches perfectly the species which has been called C.
birkmanii, also described from Texas. In the First Supplement to the Catalog
of Hymenoptera (Krombein, 1958:257), this species is indicated as the op-
1966
The Taxonomy and Nomenclature of Bees
9
posite sex of C. lanosa lanosa Cresson, on the authority of P. H. Timberlake. I
have discussed {in lift.) this matter with Mr. Timberlake, and he has indicated
that the synonymy is probably not correct. He further concurs with my view
that C. birkmanii is a probable synonym of C. subhyalina.
Mitchell ( 1962:334-335) has recorded and described this species, as C. /.
lanosa Cresson, from Florida. I have examined two of the males, from Inver-
ness, Florida, Charles Robertson Collection, and find them inseparable from
C. subhyalina. Admittedly, the seventh and eighth tergites are not perfectly
typical, but these appear to fall within the range of variation which I attribute
to this species. It should be pointed out that Mitchell’s illustration of the geni-
talia of this species (1962:336, Fig. 97) is in error, failing to show the branched
setae on the gonocoxites; the specimen from which the figure was made has
the setae conspicuously branched.
Although little is presently known of the distribution of this species, I
regard the Inverness record as questionably valid. Since the easternmost valid
record seems to be at Giddings, Washington County, Texas, almost 500 miles
west of Inverness, in a very different habitat, I feel that the Florida record
should be considered a result of mislabeled specimens. However, it should be
pointed out that according to Prof. Mitchell {personal communication) other
material, similarly labeled, represents species of bees known to occur in central
Flordia.
The male of this species is described here in order to distinguish it from
the superficially similar C. lanosa, from which it differs by the more protu-
berant clypeus, presence of a clearly defined, very narrow median impunctate
line on the clypeus, the presence of lateral face marks and different apical
ventrites.
MALE: Integument black. Bright lemon-yellow maculae as follows:
mandibles basally; labrum; clypeus; lateral face marks, ending at level of
antennal sockets; narrow transverse supraclypeal mark; underside of antennal
scape. Apex of first flagellar segment, entire second segment, dull ferruginous;
remainder of antennae brownish. Tegulae dull yellow. Wings hyaline, veins and
stigma brownish to piceous. Coxae, trochanters, femora, middle and hind tarsi
darkly rufescent; anterior tibiae and tarsi ferruginous; tibial spurs and tarsal
claws darkly rufescent to blackish. Pubescence of head, thoracic dorsum, pro-
podeum, upper half of meso- and metapleurae, anterior legs mostly, outer
surface of middle tibiae, narrow postero-basal fringe on hind tibiae, first ab-
dominal tergite, all pale fulvous; that of inner side of anterior tarsi bright
fulvous, appearing almost golden; pubescence of legs, except as noted above,
darkly fuscous-brown; that of abdomen, except first tergite, fuscous-brown to
black. Abdominal tergites reflecting deep, dull blue-black, apical margins of
tergites and ventrites testaceous. Pubescence of underside of thorax fuscous.
Head : Mandibles tridentate, very similar to C. lanosa. Labrum rugosely
punctate, with shining interstices between the rather close punctures; apex
broadly rounded; apical brush poorly developed to almost absent. Punctures of
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Contributions in Science
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clypeus slightly finer than of labrum, with shining interstices between, some-
what sparser laterally; median impunctate longitudinal line elongate, very
narrow. Frons, sides of face, and genae shiny, finely closely punctate, the
punctures barely separated. Punctures of vertex posteriorly coarser than of
frons, dense, laterally about a puncture diameter apart; areas laterad of pos-
terior ocelli and in front of anterior ocellus shiny, impunctate. Posterior ocelli
about twice an ocellar diameter apart, separated from eyes by slightly more
than an ocellar diameter, separated from posterior margin of vertex by about
two and two-fifth times an ocellar diameter. Distance from anterior ocellus to
base of clypeus equal to breadth of face at level of face of clypeus. First flagel-
lar segment about as long as scape, slightly longer than following two segments
combined; median segments of flagellum about two-thirds as wide as long.
Thorax : Punctures of mesoscutum, mesoscutellum, metanotum and pro-
podeum comparable in size to those of vertex, separated by about one-third to
three-fifths a puncture diameter, integument between slightly tessellate, mod-
erately shiny; punctures of meso- and metapleura about same as those of ver-
tex, crowded, almost touching, interstices moderately shiny; tegulae impunc-
tate, minutely tessellate. Second submarginal cell receiving first recurrent
vein slightly before middle; third submarginal cell receiving second recurrent
at its apex.
Abdomen : All tergites and ventrites with dense, piliferous punctures, with
moderately shiny interstices between; apical margins impunctate. Pubescence
of discs of tergites dense, somewhat obscuring surface, becoming progressively
longer caudally; of ventrites, a little less dense. Pseudo-pygidial area poorly
defined, almost hidden by dense pubescence on both sides.
Measurements : Body length (front of vertex to apex of second tergite),
8.0 to 10.5 mm.; forewing length, 9.0 mm.
New records of distribution of this species are as follows: KANSAS: 5
9 9, Morton County, June, 1902 (F. H. Snow; KU). TEXAS: 1 9 , Lee
County (NCSC) ; 1 9, Lee County, May 25, 1906, Malvacea (NCSC); 1 9,
Fedor, April 27, 1909 (NCSC); 1 9, Reeves County, June 15, 1940 (T. B.
Mitchell; NCSC); 1 9,1 $, Bexar County, May 1, 1929 (H. B. Parks;
NCSC); 1 8, Giddings, May 15, 1953 (R. H. Beamer; KU), on Brazoria
truncata; 1 9,1 $, Giddings, May 9, 1954 (R. H. Beamer; KU), 9 on
Monarda, $ on Gaillardia.
Centris ( Paracentris ) mexicana F. Smith
Centris Mexicana F. Smith, 1854. Cat. Hym. Brit. Mus. 2:378. 9.
A male of this species has been submitted by G. D. Butler from the
Chiricahua Mountains, Cochise County, ARIZONA, 7000-8000 feet, Septem-
ber 7, 1953 (G. D. Butler; UA), on thistle.
This is the first record of C. mexicana in the United States, although it is
a very common species during the summer months in Sonora and Chihuahua.
1966
The Taxonomy and Nomenclature of Bees
11
Centris (Paracentris) zacateca Snelling, new species
Figure 1, a and f
Diagnosis : Although this species is superficially similar to C. mexicana,
both sexes of C. zacateca may be recognized by their smaller size and the pale
pubescence on the vertex and pronotal lobes. The female has the clypeus with
a definite median impunctate line, while that of C. mexicana has the clypeal
punctures sparse, but evenly distributed so that no median impunctate line is
formed. In the male of C. zacateca the first flagellar segment is from 2.59 to
3. 10 times the length of the second; in C. mexicana it is from 3.83 to 3.87 times
the length of the second.
FEMALE: Integument of head, thorax and abdomen black, that of ab-
domen without bluish reflections; of legs, very dark rufescent appearing black,
tarsi a little lighter; tibial spurs and basitibial plates dark rufescent; tarsal claws
rufescent to dark ferruginous; tegulae lutescent; pygidial plate dark rufescent;
antennae dark rufescent, a little lighter beneath. Pubescence black except pale
fulvous as follows: top of vertex, not extending beyond lateral ocelli; occipital
margins, immediately behind vertex; dorsum of mesoscutum; mesoscutellum;
post-scutellum; upper lateral corners of propodeum; lateral lobes of pronotum
and immediately adjacent upper mesopleura.
Head : Mandibles quadridentate, apical tooth longest, teeth progressively
smaller toward the inner. Maxillary palpi five-segmented, second and third
segments subequal, fourth about \A as long as third, fifth about 2A as long as
fourth. Scape and first flagellar segment subequal; second and third flagellar
segments combined 3/s as long as first; labrum rounded apically, with very
dense preapical brush of hairs; punctures a little finer than those of clypeus,
with shining interstices. Clypeus shining, with broad impunctate median line,
tapering slightly toward apex, ending slightly before apical rim, punctures
becoming denser laterally; disc bare medially, with abundant moderately long
pubescence laterally; paraocular areas and sides of face finely punctate, the
punctures about IV2 times their diameters apart; supraclypeal area medially
impunctate, laterally like sides of face; frons a little more densely punctate,
the punctures becoming larger above; usual impunctate triangular area before
anterior ocellus; area before lateral ocelli very finely, rather closely punctate;
vertex between eyes and ocelli sparsely punctate to impunctate except narrow
band close to eyes; vertex finely, sparsely punctate posteriorly. Antennal
sockets about twice their diameter apart, removed from eyes about IV2 times
their diameter. Distance from anterior ocellus to base of clypeus about 4/7 of
distance between eyes at base of clypeus; inner orbits parallel. Distance be-
tween lateral ocelli slightly greater than that between ocelli and eyes; lateral
ocelli separated from posterior margin of vertex by about twice an ocellar
diameter.
Thorax : Punctures of mesoscutum a little larger than those of sides of
face, about their own diameters apart; the interstices moderately shiny, not
12
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No. 112
tessellate. Punctures of mesopleura slightly larger, distance between punctures
averaging about twice their diameters; interstices moderately shiny, not tes-
sellate. Punctures on anterior one-fourth of mesoscutellum as on mesoscutum,
interstices becoming slightly tessellate. Post-scutellum moderately shiny, im-
punctate, tessellate. Upper medial two-thirds of propodeum moderately shiny,
impunctate; punctures appearing laterally, equal in size and spacing to those of
mesopleura, interstices a little less shining. Basitibial scoop of middle legs
present, well-developed.
Abdomen : Moderately shiny, tessellate, setigerously punctate; apical
ventrite without median carina; pygidial plate apically truncate.
Measurements : Body length (front of vertex to apex of second tergite),
12.5 to 14.5 mm.; forewing length, 10.8 to 12.2 mm.
MALE: Pubescence, integument and wings colored as in female.
Head : Clypeus evenly rounded basally, apical third of disc slightly flat-
tened; punctation essentially as in female, with distinct median longitudinal
impunctate line. Punctures of frons and vertex as in female. Ocellar-clypeal
distance 0.59 to 0.60 times transfacial distance at level of clypeal base. Scape
and first flagellar segment subequal; second and third segments combined 0.67
to 0.68 times first flagellar. Distance between lateral ocelli a little greater than
distance between eyes and ocelli; lateral ocelli separated from posterior margin
of vertex by about twice an ocellar diameter.
Thorax and abdomen as described above for female.
Measurements : Body length (front of vertex to apex of second tergite),
1 1.4 to 13.7 mm.; forewing length, 11.2 to 12.4 mm.
Holotype male and allotype female (Los Angeles County Museum of
Natural History), 32 miles west of Pinos, 7100 feet, Zacatecas, MEXICO,
August 9, 1958 (R. M. Straw, #1486), on Penstemon tenuifolius. Paratypes :
5 8 8 , 1 $ , same data as holotype; 1 8 , Zacatecas, Zac., July 16, 1954 (J. W.
MacSwain: CIS); 1 8 , Guadalupe, Zac., June 28, 1953 (C. & P. Vaurie; D.
Rockefeller Mex. Exp., AMNH) ; 8 $9,9 mi. S. Fresnillo, Zac., Aug. 9, 1954
(E. G. Linsley, J. W. MacSwain, R. F. Smith; CIS); 2 9 9, Penudas, Aguas
Calientes, July 17, 1954 (J. W. MacSwain; CIS); 1 8,1 9, Encarnacion de
Diaz, Jalisco, Aug. 18, 1953 (C. & P. Vaurie; D. Rockefeller Mex. Exp., 1953,
AMNH) ; 1 9 , Ojuelos, Jalisco, June 25, 1953 (C. & P. Vaurie; D. Rockefeller
Mex. Exp., 1953, AMNH); 1 8, 21-23 mi. W. Ojuelos, 7000 feet, Jalisco,
July 29, 1958 (R. M. Straw, #1446; LACM); on Penstemon tenuifolius’, 5
8 8, 5 9 9 , 20 mi. S. Durango, 6300 feet, Durango, Aug. 12, 1958 (R. M.
Straw, #1514; LACM), on P. tenuifolius ; 1 9, 9.5 mi. N. Chihuahua, 5000
feet, Chih., Aug. 18, 1952 (C. D. MacNeill and E. E. Gilbert; CIS); I 9,
Llano de Rio Santa Clara, 27 mi. W. Parrita, Chih., Aug. 12, 1950 (R. F.
Smith; AMNH) ; 6 8 8, Canon de Santa Clara, 5 mi. W. Parral, Chih., July 6,
1954 (J. W. MacSwain and E. I. Schlinger; CIS), on Baccharis; 1 9, Rodeo,
Hidalgo Co., N. Mex., Aug. 22, 1958 (R. M. Bohart; UCD).
1966
The Taxonomy and Nomenclature of Bees
13
Centris (Paracentris) angustifrons Snelling, new species
This species, currently known only from the unique type, does not seem
to be closely related to any species known to me. Superficially it somewhat
resembles C. subhyalina, with which it has in common a black clypeus and
similarly colored pubescence, but differs from that, and all similar species, in
the much narrower face. Until the male is discovered, the affinities of C. angus-
tifrons will remain uncertain.
FEMALE: Integument blackish-ferruginous, abdomen without irides-
cent or metallic reflections; legs ferruginous, tibial spurs, apical tarsal segments
and tarsal claws darker. Wings hyaline, with distinct brownish tinges, veins and
stigma blackish-brown. Pubescence of head, dorsum of thorax, upper half of
meso- and metapleurae, propodeum (except sides) and first tergite ochraceous,
that of thoracic dorsum slightly tinged with fulvous at the tips; elsewhere light
to rather dark brown (as in C. subhyalina) .
Head'. Mandibles quadridentate, apical tooth longest, median teeth ap-
proximately equal in size to each other, inner tooth a little longer, broader than
median teeth. Maxillary palpi five-segmented, second segment the longest,
third a little shorter, but longer than combined length of last two, apical seg-
ment the shortest. Labrum moderately shiny, disc coarsely rugoso-punctate;
apical margin broadly rounded. Clypeus duller than labrum, with raised
median impunctate area, laterad of raised portion with a few, variably spaced,
coarse punctures. Punctures of frons fine, dense, integument somewhat shin-
ing; punctures of sides of face conspicuously coarser than those of frons, but
finer than those of clypeus, mostly separated by a puncture diameter or more,
especially between the eyes and ocelli where they are quite sparse. Punctures
of vertex, behind ocelli, fine, dense. Facial quadrangle slightly longer than
broad. Distance from anterior ocellus to clypeal base 0.53 times breadth of
face at level of clypeal base; distance between lateral ocelli about 2.1 times an
ocellar diameter, about 1.6 times distance between ocelli and eyes; distance
between ocelli and posterior margin of vertex about 2.7 times an ocellar diame-
ter. First flagellar segment slightly longer than scape, longer than following
three segments combined.
Thorax: Punctures of mesoscutum finer than those of mesopleurae, sepa-
rated by about a puncture diameter on both areas, mesopleural punctures equal
in size to those of sides of face; mesoscutellar punctures a little larger, denser,
than those of mesoscutum, interstices more distinctly tessellate; propodeum
shiny, posterior face with scattered fine punctures, lateral faces with punctures
a little coarser, separated by slightly more than puncture diameter. Anterior
basitarsi with two or three elongate, spatulate setae on posterior ventral margin
(lacking in all other species of Paracentris) ; the usual row of coarse, flattened
setae present on anterior ventral margins of anterior and middle basitarsi.
Secondary basitibial plate poorly defined (evidently due to wear).
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Contributions in Science
No. 112
Abdomen : Punctures of tergites rather dense, finest on basal segments,
becoming progressively coarser on succeeding segments.
Measurements : Body length (front of vertex to apex of second tergite)
10.3 mm.; forewing length, 10.6 mm.
Holotype : Female (Los Angeles County Museum of Natural History),
Huachuca Mountains, Arizona, August 19, 1903 (Oslar).
Centris (Para centris) anthracina Snelling, new species
Figure 1, b and g
Centris clypeata, Cockerell, 1948. Proc. U.S. Natl. Mus. 98:474. $ 8.
(— clypeata Friese, 1900?)
This is the species which has long been known as C. clypeata Friese; C.
clypeata Friese is a junior homonym of C. clypeata Lepeletier. A new name
is therefore necessary for Friese’s species. Unfortunately two species have been
confused as Friese’s and until the type can be critically examined it cannot be
determined which of the two was before him when he described C. clypeata.
I have therefore elected to describe both of the species involved as new, since
one is presumably identical with Friese’s species and the other is undescribed.
In the event that either proves to be a synonym of C. clypeata Friese (a junior
homonym) the name applied here, although a synonym, then becomes avail-
able for the species as the next validly proposed name. This method is advan-
tageous in that type specimens (C. anthracina or C. laevibullata, as the case
may be) are then located in American museums available to those who are
most directly concerned with the genus.
Diagnosis’. This handsome species belongs to a small group which in-
cludes C. nigerrima Spinola and more remotely, C. laevibullata Snelling, new
species. From C. laevibullata, the male of which is unknown, the female
differs by having the first flagellar segment 5.14 to 5.28 times the length of the
second (3.5 times the second in C. laevibullata) , the narrower face (TFD
2.26 X OCD in C. anthracina, TFD 1.87 to 2.02 X OCD in C. laevibullata )
and by the median impunctate line which extends to the clypeal apex (Median
impunctate line extending about % of distance toward apex in C. laevibullata .)
FEMALE: Integument of head, thorax and abdomen black, that of
abdomen with vaguely bluish reflections; of medio- and distitarsi, rufescent;
tibial spurs black; basitibial and pygidial plates very darkly rufescent. Tegulae
dark brownish. Wings light brownish, reflecting dull violaceous tints, stigma
and veins dark brownish. Long, erect black or blackish pubescence as follows:
Face, vertex except laterally, genae, thorax except propodeum, legs and ab-
domen (progressively longer on tergites, reaching greatest length on fourth,
that of fifth shorter, not so dense, sixth sparsely pubescent on basal half; discs
of ventrites with pubescence sparser, apical margins of last four segments
with long, erect pubescence). Tegulae with sparse, erect black pubescence.
1966
The Taxonomy and Nomenclature of Bees
15
Figure 1. a-e, ventrite VIII and f-j, ventrite IX, respectively of males: a & f, C.
{Paracentris) zacateca ; b & g, C. ( P .) anthracina; c & h, C. ( Melanocentris ) strawi;
d & i, C\ (M) ruthannae\ e & j, C. ( M .) anomala.
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Contributions in Science
No. 112
Head : Mandibles similar to those of C. nigerrima, apical tooth yellow
except at apex. Maxillary palpi five-segmented, two apical segments slightly
longer than basal, third longest. First flagellar segment longer than scape, about
equal to following three combined. Labrum narrowly rounded apically, rather
coarsely, rugosely punctured, with a few long, seta-like hairs at apex, in strong
contrast to the shorter, plumose hairs of the disc. Clypeus shiny, sparsely,
coarsely punctate, with slightly raised median impunctate line, ending about
one-third an ocellar diameter from the narrow, impunctate testaceous apical
rim; pubescence sparse on disc, more abundant laterally and basally. Paraocu-
lar and supraclypeal areas, interantennal area, frons and vertex behind and
between ocelli densely punctate; vertex laterally sparsely punctate, shiny, the
punctures larger than of frons, slightly smaller than of clypeus. Distance be-
tween antennal sockets distinctly greater than distance from sockets to inner
margin of eyes. Posterior ocelli about two and one-half times an ocellar diame-
ter apart; distance between posterior ocelli slightly greater than distance be-
tween ocelli and eyes, distinctly greater than distance between ocelli and
posterior margin of vertex. Eyes strongly converging above; ocellar-clypeal
distance less than transfacial at base of clypeus.
Thorax : Mesoscutum, mesoscutellum, metanotum and pleura moderately
shiny, with abundant punctures intermediate in size between those of clypeus
and of frons; propodeum somewhat smaller, more sparsely punctate. Tegulae
abundantly, minutely punctate. Claws of hind tarse minutely dentate.
Abdomen : Punctured as in C. nigerrima. Pygidial plate with faint median
carina along apical third, broadened basally, lateral margins slightly raised,
apex truncate. Apical ventrite with weak, longitudinal carina medially.
Measurements : Body length (front of vertex to apex of second tergite),
13.0 to 13.5 mm.; forewing length 13.5 to 14.0 mm.
MALE: Integument black, similar to female. Labrum and clypeus (except
laterally), and transverse supraclypeal mark shining creamy-yellow; apical
tooth of mandible largely yellowish. Pubescence essentially as in female.
Head : Broader than long, eyes strongly converging above. Mandibles
tridentate, apical tooth longest. Maxillary palpi as in C. nigerrima. Labrum
broadly rounded at apex, shiny, irregularly and sparsely punctate, with distinct
sparse, erect, short black pubescence. Clypeus shining, rounded when viewed
from side, with moderately large punctures, apical middle slightly flattened;
with rather sparse, long, black pubescence laterally. Supraclypeal area duller,
almost impunctate. Para-ocular and inter-antennal areas, frons and vertex
duller than clypeus, closely punctate, the punctures about equal in size to those
of clypeus; the usual impunctate area in front of anterior ocellus and vertex
between eyes and ocelli present. First flagellar segment much longer than scape,
longer than following four segments combined, about equal to distance be-
tween eyes at vertex. Greatest facial breadth less than distance between anterior
ocellus and base of clypeus; posterior ocelli about twice an ocellar diameter
apart, separated from eyes by slightly more than an ocellar diameter.
1966
The Taxonomy and Nomenclature of Bees
17
Thorax : Punctured as in female, but tegulae more sparsely punctate. Hind
femora about one-third as broad as long.
Abdomen : Pubescence and punctation as in female.
Measurements'. Body length (front of vertex to apex of second tergite),
12.0 to 12.7 mm.; forewing length, 13.4 to 14.2 mm.
Holotype : Male (Museum of Comparative Zoology); Uyaca Mtn.,
HONDURAS, March 23 (R. Williams). Allotype : Female (Museum of
Comparative Zoology); Antigua, GUATEMALA, December 26 (A. Pelen).
Paratypes : 2 $ 8, same data as Allotype (MCZ, AMNH); 1 9, Antigua,
GUATEMALA, December 17 (A. Pelen; USNM); 1 $, Antigua, GUATE-
MALA, no date (W. P. Cockerell; USNM)3; 1 9, MEXICO, July, 1935
(C. F. Baker colln.; USNM) ; 1 $ , Santa Tecla (= Nuevo San Salvador), EL
SALVADOR, November 11, 1955 (“P. A. B” #648.47; USNM): 1 $ , San
Mateo, COSTA RICA, May 21, (collector unknown; AMNH, #25614); 1 $,
San Mateo, COSTA RICA, December, 1920 (collector unknown; AMNH,
#25614); 12 $ $, ECUADOR, no date (C. F. Baker colln.; USNM). Three
paratypes have been retained by the author, the remainder returned to their
respective collections.
Centris (Paracentris) laevibullata Snelling, new species
Diagnosis'. This species is based, as the preceding, upon a specimen deter-
mined by Cockerell as C. clypeata Friese. However, it is a very different species
and is readily distinguished from all other species of Paracentris by the clypeal
structure and rather strongly metallic blue color of the abdomen. For addi-
tional characters see discussion under C. anthracina.
FEMALE: Integument and pubescence as described for C. anthracina.
Wings very dark brownish, stigma and veins almost black. Abdomen with
rather strong dark metallic blue reflections.
Head: Maxillary palpi, mandibles and labrum as in C. anthracina. Clyp-
eus with median, impunctate, slightly swollen area on basal half, in strong
contrast to duller, coarsely, rugosely punctate apical and lateral areas; median
apical area with rugose, elongated punctures. Remainder of facial punctation
about as in C. anthracina. Facial breadth at level of base of clypeus 1.8 times
distance from anterior ocellus to base of clypeus. Posterior ocelli separated by
about twice an ocellar diameter, distance between ocelli distinctly greater than
distance between ocelli and eyes, slightly greater in distance between ocelli and
posterior margin of vertex. Distance between antennal sockets much greater
than distance from sockets to eyes. Antennae as in C. anthracina.
3 Apparently all of the above type material was recorded by Cockerell (1949) as
C. clypeata Friese. In that paper he records a single male from the type locality as
having been taken on the flowers of Wigandia. Whether or not this is the specimen
here selected as type of C. anthracina is not known, as there is nothing on the labels
to indicate this. He also records a female from Escuintla, Guatemala, which I have
not seen.
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Contributions in Science
No. 112
Centris laevibullata is otherwise similar to C. anthracina, but the pygidial
plate has the sides converging to a broadly rounded apex, and the surface is a
little more roughened.
Measurements'. Body length (front of vertex to apex of second tergite),
13.5 mm.; of forewing, 13.0 mm.
Holotype : Female (Los Angeles County Museum); Orizaba, Veracruz,
MEXICO, June 29, 1933. Paratype : female, 14 mi. NW. Zitacuaro, Micho-
acan, MEXICO, Aug. 24, 1959 (L. A. Stange, A. S. Menke; UCD).
The following key may prove useful in identifying the species of Para-
centris now known to occur in America north of the Panama Canal Zone. Body
lengths are measured from the posterior margin of the anterior ocellus to the
apical margin of the second tergite, with the head and abdomen in normal
position. The ocellar-clypeal distance (OCD) is measured from the base of the
clypeus to the anterior margin of the anterior ocellus; the transfacial distance
(TFD) is the breadth of the face measured at the level of the base of the
clypeus.
KEY TO NORTH AND CENTRAL AMERICAN SPECIES OF
PARACENTR1S
Antennae with twelve segments; abdomen with six segments . . . females
Antennae with thirteen segments; abdomen with seven segments . . males
FEMALES
1. Pubescence of head, thorax and legs entirely or predominantly black (that
of mesopleura entirely black) 2
Pubescence of head, thorax and legs entirely or predominantly pale (that
of mesopleura light in part or entirely so) 7
2(1). Pubescence of thoracic dorsum entirely black 3
Pubescence of thoracic dorsum pale, at least in part 5
3(2). First flagellar segment 3.5 times length of second (median impunctate
area of clypeus extending about % of distance toward apex; TFD 1.87
to 2.02 x OCD) (Mexico) laevibullata Snelling
First flagellar segment at least 5.0 times second 4
4(3). First flagellar segment 6.0 times second; TFD 1.12 to 1.23 x OCD;
median impunctate line of clypeus extending toward apex as a very nar-
row, slightly raised line (ending about Vi an ocellar diameter from apical
margin) (s. Ariz., n. Mex.) aterrima F. Smith
First flagellar segment 5.14 to 5.28 times second; TFD 2.26 x OCD;
median impunctate area of clypeus broad over entire length (s. Mex. to
Ecuador) anthracina Snelling
1966
The Taxonomy and Nomenclature of Bees
19
5(2). Clypeus protruding very nearly as far in front of eye as eye is wide when
viewed in profile; pubescence of thoracic dorsum usually dark fox-red
(Calif.) rhodomelas Timberlake
Clypeus no more than half as wide as eye when viewed in profile, usually
much less; pubescence of thoracic dorsum whitish 6
6(5). Large species, 14.5 to 18.5 mm. long; pubescence of vertex and pronotal
lobes black; clypeal punctures sparse, becoming obscure apically; apical
middle of clypeus slightly roughened (s. Ariz., n. Mex.)
mexicana F. Smith
Smaller, 12.5 to 14.5 mm. long; pubescence of vertex and pronotal lobes
pale; clypeus with median impunctate line widest above, punctures ar-
ranged in oblique rows (s. N.M., n. Mex.) zacateca Snelling
7 ( 1 ). Maxillary palpi four-segmented and pubescence mostly pale whitish;
surfaces of tergites obscured by short, appressed pubescence 8
Maxillary palpi five-segmented; if four-segmented, abdominal tergites
beyond second with pubescence entirely dark, not obscuring surface . . 9
8(7). Mandibles tridentate; abdominal ventrites three to five with distinct
apical fringes of moderately long white pubescence (N. Mex., Ariz.,
Calif., Nev., n. Mex.) pallida W. Fox
Mandibles quadridentate; ventrites without pale apical pubescent fringes
(nw. Mex., Calif., Ariz., Nev.) tiburonensis Cockerell
9(7). Clypeal integument entirely black, immaculate 10
Clypeal integument at least partially yellow, orange or red 13
10(9). Clypeus sparsely punctate, with broad median impunctate line; pubes-
cence of second tergite variable 11
Clypeus coarsely, closely punctate, median impunctate line, when present,
very narrow, sharply defined, slightly raised; no pale pubescence on
second tergite (Tex., Kans.) subhyalina W. Fox
11(10). Pubescence of second tergite and lower half of mesopleura pale, at
least in part; facial quadrangle at least as broad as long, usually slightly
broader 12
Pubescence of second tergite and lower half of mesopleura dark; facial
quadrangle slightly longer than broad (Ariz.) .... angustifrons Snelling
12(11). Eye, viewed laterally, wider than gena; clypeus distinctly bulging
basally; pubescence of second tergite entirely pale; ventrites three to five
with pale apical fringes; vernal to early summer (N. Mex., Ariz., Calif.,
Nev., n. Mex.) hofjmanseggiae Cockerell
Eye, viewed laterally, no wider than gena; clypeus weakly bulging basally;
at least some discal pubescence of second tergite black; ventrites three to
five lacking pale fringes; late summer to autumnal (Calif., Nev.)
calif ornica Timberlake
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No. 112
13(9). Large species, 15.5 to 18.5 mm.; clypeal punctures sparse, fine; TFD
2.14 to 2.17 x OCD (Tex., Colo., N. Mex., Ariz., n. Mex.)
caesalpiniae Cockerell
Smaller, 8.5 to 13.5 mm.; clypeal punctures denser, separated by less than
two times a puncture diameter 14
14(13). Inner orbits distinctly convergent above; small species, 8.5 to 10.5
mm.; TFD 1.71 to 1.74 x OCD; integument of legs dark rufescent to
fuscous, never ferruginous 15
Inner orbits barely, if at all, convergent above; size variable; TFD at
least 1.90 x OCD; integument of legs variable, frequently ferruginous . .
16
15(14). Clypeal maculation pale yellow (Tex., N. Mex., e. Ariz., ne. Mex.) . .
lanosa lanosa Cresson
Clypeal maculation yellow-orange to light ferruginous (Ariz., Nev.,
Calif., nw. Mex.) lanosa resoluta Cockerell
16(14). TFD 1.90 to 1.95 x OCD; sternal pubescence dark brown or blackish,
contrasting sharply with ochraceous pubescence of mesopleurae; pubes-
cence of middle and hind legs entirely blackish; inner orbits diverging
above; (first flagellar segment shorter than following three combined)
(Tex., N. Mex., Ariz., Calif., n. Mex.) atripes Mocsary
TFD at least 2.0 x OCD; sternal pubescence variable, but never so
strongly contrasting; inner orbits almost parallel 17
17(16). Clypeus sparsely punctate, with median area pale yellowish; legs dark
rufescent; paraocular areas black; TFD 2.10 to 2.21 x OCD (N. Mex.,
Ariz., Nev., Calif., n. Mex.) hoffmanseggiae Cockerell
Legs and clypeus bright ferruginous; paraocular areas and stripe along
inner orbits yellowish; TFD 2.09 to 2.13 x OCD (Tex., N. Mex., Ariz.,
Calif., Nev., n. Mex.) rhodopus Cockerell
MALES
1. Clypeus either entirely black, or with two small, widely separated apical
maculae 2
Clypeus largely or entirely yellow, white or orange-yellow 5
2(1). Pubescence of thoracic dorsum white or fulvous 3
Pubescence of thoracic dorsum black aterrima F. Smith
3(2) . Pubescence of posterior pronotal lobe, and usually entire lateral pronotal
area, pale; first flagellar segment 2.6 to 3.1 times second
zacateca Snelling
1966
The Taxonomy and Nomenclature of Bees
21
Pubescence of lateral pronotal area, including pronotal lobe, black; first
flagellar segment 3.8 to 3.9 times second mexicana F. Smith
4(1). Pubescence of head and thorax at least partially pale 5
Pubescence of head, thorax and abdomen black .... anthracina Snelling
5(4). Maxillary palpi four-segmented; antennal scape black beneath; ab-
dominal pubescence pale, dense, suberect 6
Maxillary palpi five-segmented; if four, then tergites with pubescence
sparse, dark, or antennal scape with yellow stripe beneath 7
6(5). Face narrow, inner orbits strongly convergent above; distance between
eyes and ocelli about Vi diameter of lateral ocelli pallida W. Fox
Face broader, inner orbits almost parallel; distance between eyes and
ocelli about equal to diameter of lateral ocelli . . . tiburonensis Cockerell
7(5). Face broad, inner orbits not or scarcely convergent above; TFD at least
1.82 x OCD 8
Face narrower, inner orbits usually strongly convergent above; TFD no
more than 1.72 x OCD 10
8(7). Integument of posterior femora and tibiae ferruginous; pubescence of
hind basitarsi black, contrasting with pale tibial pubescence; paraocular
area and underside of scape bright lemon-yellow; apical margins of tergites
with thin, medially interrupted pubescent fasciae (TFD 1.82 x 2.0 x
OCD) rhodopus Cockerell
Integument of posterior femora and tibiae black or darkly rufescent; pu-
bescence of hind basitarsi and tibiae concolorous; paraocular areas and
underside of scape black; tergites without apical pubescent fasciae ... 9
9(8). Large species, 13.5 to 12.4 mm.; TFD 1.92 to 1.97 x OCD; posterior
femora with one-half or more of pubescence pale
caesalpiniae Cockerell
Smaller, 10.0 to 12.4 mm.; TFD 1.82 to 1.88 x OCD; posterior femora
usually with pubescence all dark, occasionally with a light stripe along
posterior margin atripes Mocsary
10(7). Clypeus not at all strongly protuberant, only half as far in front of
mandibular base as eye is wide; first flagellar segment length variable . 1 1
Clypeus strongly protuberant, as far in front of mandibular base as eye is
wide when viewed laterally; first flagellar segment shorter than following
four combined (pubescence of thoracic dorsum fulvous to fox-red) ....
rhodomelas Timberlake
11(10). Abdominal tergites with abundant erect or suberect pale discal pubes-
cence 12
Abominal tergites beyond first without pale erect discal pubescence . . 13
Contributions in Science
No. 112
12(1 1 ) . TFD 1.35 to 1.38 x OCD; underside of scape with pale spot or narrow
stripe; pubescence of tergites IV— VI entirely pale; late summer and
autumnal californica Timberlake
TFD 1.49 to 1.53 x OCD; underside of scape black; pubescence of
tergites IV— VI usually at least partly fuscous; vernal to early summer . .
hoff manse ggiae Cockerell
13 ( 1 1). Paraocular area and underside of scape yellow; clypeus coarsely,
closely punctate, with narrow median impunctate line
suhhyalina W. Fox
Paraocular area and underside of scape black; clypeus sparsely punctate,
with very broad median impunctate area lanosa Cresson
Subgenus CENTRIS Fabricius
Centris ( Centris ) inermis gualanensis Cockerell
Centris inermis gualanensis Cockerell, 1912. Annals and Magazine of
Natural History (8) 9:568. Schwarz, 1934. Amer. Mus. Nov. 722:12. Miche-
ner, 1954. Bui. Amer. Mus. Nat. Hist. 104:138-139.
ICentris inermis Friese, 1900. Annalen des K. K. Naturhistorischen
Hofmuseums, Wien, 15:314 (part). Cockerell, 1928. Psyche, 35:173; 1949.
Proc. U. S. Natl. Mus. 98:479.
Centris inermis pallidifrons Cockerell, 1949. Proc. U. S. Natl. Mus., 98:
479. $ . NEW SYNONYMY
The form described from Zamorano, Honduras, by Cockerell as C. i.
pallidifrons does not seem sufficiently distinct from his C. i. gualanensis to
warrant separation. The duller clypeus of the former seems to be a matter of
personal appreciation and is not consistent throughout any series. Two C. i.
pallidifrons cotypes have the clypeus much brighter than Cockerell’s descrip-
tion would allow. Furthermore, the two forms are not geographically separa-
ble, and so it seems best to reduce this form to the synonymy of C. i. gualanen-
sis.
The male recorded by Cockerell in the same paper as C. inermis has the
inner tooth of the mandibles obscurely bidentate, which would seem to be the
reason for his statement that they are quadridentate. Normally the males of this
species have tridentate mandibles.
Centris (Centris) eisenii W. Fox
Centris eisenii Fox, 1894. Proc. Calif. Acad. Sci., 4:22. 9. Cockerell,
1923. Proc. Calif. Acad. Sci. (4) 12:75. 9.
This species is recorded from the United States for the first time.
NEW RECORDS: ARIZONA: 1 $ , Nogales, August (Oslar; E. P. Reed
colln.; CAS); 2 9$, Nogales, June 1, 1903 (Oslar; CU); 1 9, Picacho, Pinal
Co., June 18, 1961 (F. D. Parker; UCD). SONORA: 1 9 , San Bernardo,
1966
The Taxonomy and Nomenclature of Bees
23
Rio Mayo, July 6, 1935 (H. S. Gentry; LACM); 2 2$, Guaymas, April 11,
1921 (E. P. Van Duzee; CAS); 1 2, Guaymas, July 2, 1952 (W. H. Lange;
UCD). MORELOS: 1 2, 18 $ $ , Lake Tequesquitengo, 2800 it., March 16,
22, June 5, 1959; 1 2, 3-6 mi. S. Cuernavaca, 4000 ft., April 1, 1959; 1 $,
Huajintlan, 2800 ft., April 11, 1959; 1 $ Canon de Lobos, Yautepec, 4000 ft.,
April 13, 1959; 1 2 , 3 mi. N. Alpuyeka, 3400 ft., April 18, 1959 (all H. E.
Evans; CU). PUEBLA: 6 2 2, 11 mi. SE Acatlan, July 10, 1952 (E. E. Gil-
bert, C. D. MacNeill: CIS). SINALOA: 1 2, 14 mi. SE Elota, May 5, 1953
(R. C. Bechtel, E. I. Schlinger; CIS); 3 $ $ ,2 mi. N. San Miguel, June 17,
1956 (R. P. Allen; CIS).
Centris ( Centris ) decolorata Lepeletier
Centris decolorata Lepeletier, 1841. Historie Naturelle des Insectes, Hy-
menopteres, II, p. 160. $ . Friese, 1900. Annalen des K. K. Naturhistorischen
Hofmuseum, Wien, 15:325-326. $ 2.
Material now at hand indicates that this species occurs in the extreme
southern portion of Texas, on the off-shore islands.
NEW RECORDS: TEXAS: Cameron Co.: 1 2,2 $ 8, Pt. Isabel, June
23-27, 1956; 1 $ , Boca Chico, June 26, 1956; 5 $ $, Padre Isl., June 25, 1956
(all H. E. Evans and E. G. Matthews; CU). VERACRUZ: 2 $ $, Vera Cruz,
June 12, 1959 (H. E. Evans; CU); 1 2,2 $ $, Vera Cruz, no date (C. F.
Baker colln., 1 $ with #8257; CU) .
Centris ( Centris ) flavofasciata Friese
Centris flavifrons var. flavofasciata Friese, 1899. Termeszetrajzi Fuzetek,
22:46. $. 1900, Annalen des K. K. Naturhistorischen Hofmuseums, Wien,
15:318. $.
Centris flavofasciata Michener, 1954. Bull. Amer. Mus. Nat. Hist., 104:
137. $.
This species is added to the United States fauna for the first time.
NEW RECORDS: ARIZONA: 2 2 2, Nogales, July 10, 1903 (Oslar;
CU). MORELOS: 6 $ $, L. Tequesquitengo, March 16, 22, 1959 (H. E.
Evans; CU). GUERRERO: 1 2, Xalitla, 1500 ft., March 19, 1959 (H. E.
Evans; CU). SONORA: 1 2, 16 mi. S. Empalme, May 7, 1953 (E. I.
Schlinger; CIS).
Subgenus XANTHEM1S1A Moure
Centris (Xanthemisia) aethiops Cresson
Centris aethiops Cresson, 1865. Proc. Ent. Soc. Phila. 4:193. 2. Friese,
1900. Annalen des K. K. Naturhistorischen Hofmuseums, Wien, 15:268, 2.
Centris armillatus Cresson, 1865. Trans. Amer. Ent. Soc. 2:298. $.
Friese, 1900. Annalen des K. K. Naturhistorischen Hofmuseums, Wein, 15:
268-269. $ . NEW SYNONYMY.
24
Contributions in Science
No. 112
NEW RECORDS: CUBA: 1 8, no further data (AMNH, No. 26426);
1 8, Guantanamo, June, 1921 (C. T. Ramsden; AMNH); 1 $, San Carlos
Est., Rio Seca, Guantanamo, April 17, 1914 (C. T. Ramsden; AMNH).
Although the synonymy of C. armillata with C. aethiops has been known
for many years, there seems to have been no formal publication of this fact,
and I am taking the opportunity to do so at this time. This species is here as-
signed to Xanthemisia on the basis of the mandibular structure of the female
and the ventrites and genitalia of the male. Although the males of this and the
following species have entirely black faces, unique for the subgenus, this char-
acter is not considered significant. The female of C. aethiops differs most obvi-
ously from other Xanthemisia in having the thoracic pubescence entirely dark.
Centris (Xanthemisia) carolae Snelling, new species
MALE: Integument black; that of legs rufescent; abdominal tergites dull
metallic greenish-bronze, the apical margins lutescent; ventrites dull metallic
greenish-bronze, with very broad lutescent apical margins. Pubescence mostly
black or very dark brownish; anterior one-fourth and posterior one-fourth of
mesoscutum, the mesoscutellum and metanotum with pubescence bright
lemon-yellow; of abdomen mostly golden, suberect, becoming longer and
denser on successive segments, very dense on four apical segments, especially
laterally; ventrites with pubescence long, golden. Labrum densely pubescent,
but clypeus mostly nude, except laterally. Tegulae with pubescence sparse, very
short, erect. Longest hairs of hind tibiae much longer than greatest tibial width:
longest hairs of hind basitarsi almost equal to length of that segment. Tegulae
black; tibial spurs blackish; tarsal claws yellowish, with rufescent apices. Seg-
ments two to basal half of thirteen of flagellum ferruginous beneath, flagellum
otherwise darkly infuscated. Mandibles black, with reddish preapical mark.
Head : Mandibles tridentate, apical tooth long and slender, middle tooth
slightly larger than inner; two inner teeth acute, triangular, both well separated
from apical. First flagellar segment distinctly longer than scape, about equal to
the following three combined. Labrum broadly rounded apically, rugosely
punctate, moderately shining. Clypeus much duller than labrum except in basal
middle; disc with punctures large, shallow, well separated, becoming denser
laterally; median line from apex to base broadly, shallowly concave, nearly
impunctate. Paraocular area, supraclypeal area, frons (except shiny impunc-
tate triangular area in front of anterior ocellus) densely, rather coarsely punc-
tate, but the punctures distinctly smaller than of clypeal disc, the punctures
somewhat less approximate in ocellar-ocular area. Distance from anterior
ocellus to base of clypeus about equal to breadth of face at level of base of
clypeus; posterior ocelli separated from eyes by slightly more than diameter of
ocelli; distance between ocelli and eyes much less than distance from ocelli to
posterior margin of vertex; eyes slightly converging above.
Thorax : Mesoscutum, mesoscutellum and metanotum very closely punc-
1966
The Taxonomy and Nomenclature of Bees
25
tate, punctures about equal to those of lateral margins of clypeus; pleurae
closely, more coarsely punctate; propodeum less closely punctate, the punc-
tures about equal to those of mesoscutum; basal area large, tessellate, sparsely
punctate. Wings strongly infuscated with brownish, less strongly so beyond the
cellular area, reflections violaceous; stigma and veins blackish. Hind femora
about one-half as broad as long, with weakly developed longitudinal carina
beneath; greatest width of hind tibiae about one-fourth their length.
Abdomen : Discs of segments very sparsely punctate, surface moderately
shining, the few punctures piliferous. Pseudopygidial area distinct, apex sub-
truncate, disc slightly depressed so that lateral margins appear carinate.
Measurements'. Body length (front of vertex to apex of second tergite),
14.6 mm.; forewing length, 14.7 mm.
Holotype : Male (Los Angeles County Museum of Natural History);
Tuxtla Chico, 875 m., Chiapas, MEXICO, March 14, from the duBois collec-
tion.
I am very happy to be able to dedicate this outstandingly attractive species
to Miss Carol Bumgardner, a true and honest friend. The combination of black,
yellow and greenish-bronze is very striking, the bronze taking on subtle under-
tones beneath the golden abdominal pubescence.
Subgenus HEMIS1ELLA Moure
Centris ( Hemisiella ) trigonoides subtar sata Cockerell
Centris lanipes subtarsata Cockerell, 1949. Proc. U. S. Natl. Mus., 98:
476-477: $ $.
An examination of a long series of both sexes of this form, including two
cotypes from Honduras, indicates that this bee is distinct from C. lanipes
Fabricius, but is inseparable, structurally, from C. trigonoides Lepeletier (=
C. hoplopoda Moure). The latter is a widely distributed species in South
America, and typical material has been recorded from the Canal Zone, Pana-
ma. The male genitalia and hidden ventrites of C. subtarsata offer no characters
by which it can be separated from C. trigonoides.
The populations from Mexico, Guatemala and Honduras, however, differ
consistently from those farther south in that most of the pubescence of the
abdominal tergites is light ferruginous to yellow-ferruginous, rather than
fuscous to black. The females further differ in that the two apical maculae of
the clypeus are somewhat smaller and less approximate in the northern popu-
lations. The Guatemala males have the dark infuscations of the tergites more
restricted, and in some cases entirely lacking except on the base of the first
segment, and the legs somewhat less extensively infuscated. Therefore, I regard
C. subtarsata as a subspecies of C. trigonoides.
The following description of the chromatic characteristics is given in
order to separate this from the nominate form. For the structural characteris-
tics of the species, refer to the original description of C. hoplopoda by Moure
(1943; 160).
26
Contributions in Science
No. 112
FEMALE: Structurally inseparable from nominate form, apparently
differing principally in that the suberect pubescence of the tergites is light
fuscous to yellow-ferruginous, rather than dark fuscous; pubescence on inner
side of hind basitarsi ferruginous instead of blackish; labrum with a large
brownish median infuscation; facial macula light yellowish, clypeal marks well
separated medially; legs variable, but generally with front and middle coxae,
trochanters, femora, tibiae and basitarsi rufescent; hind legs entirely and all
post-basitarsal segments ferruginous.
MALE: Structurally inseparable from nominate form; apical tooth of
mandibles dull orange-yellow, narrow transverse ferruginous band separating
this area from the dull creamy-yellow basal two-thirds of mandible (apical
tooth dark in C. t. trigonoides ); facial maculae pale yellowish (somewhat
darker in C. t. trigonoides) ; legs lightly rufescent or brownish (darkly rufes-
cent or blackish in C. t. trigonoides)’, abdomen mainly light ferruginous (ter-
gites usually strongly infuscate in C. t. trigonoides) ; pubescence of second to
fourth tergites light fuscous (darker in C. t. trigonoides) , in some individuals
entirely pale yellow-ferruginous.
One female from Chichen-Itza, one from Santa Emilia and two from
Costa Rica have the pubescence of the tergites mostly blackish, but all have
the hairs of the inner side of the hind basitarsi ferruginous and the clypeal
maculae well separated, except in the Chichen Itza specimen in which they are
almost contiguous. The two Costa Rican specimens have the front femora
darkly rufescent, with a dorsal light ferruginous stripe from base to apex. A
female, here assigned to C. t. trigonoides, from Nova Teutonia, BRAZIL, has
the abdominal pubescence light fuscous, instead of black, but that on the inner
side of the hind basitarsi is black and the clypeal maculae are large, almost
touching medially.
Since no additional specimens of this form have been recorded since it
was originally described, the following are made known. All specimens are
from the collection of the Museum of Comparative Zoology.
NEW RECORDS: MEXICO: 1 9, Chichen-Itza, June 29; 1 9,1 8,
Acapulco (A. Agassiz). BRITISH HONDURAS: 7 8 8, Banque Viejo
(Father Stanton). GUATEMALA: 5 $9,5 8 8, Santa Emilia, Pochuta,
1000 m., Feb.-March, 1931 (J. Bequaert); 1 9, Ciudad de Guatemala (J.
Bequaert); 2 8 8, Los Amates (Kellerman); 1 8, Sanerate (Kellerman).
NICARAGUA: 14 9 9 , Polvon (J. McNeill Exped.). COSTA RICA: 2 9 9 ,
Palmar, Puntarenas.
Centris ( Hemisiella ) transversa Perez
Centris transversa Perez, 1905. Bulletin Museum Histoire Naturelle,
Paris, 11:39: 9 8.
Dr. Butler has submitted a single female of this species which he had
collected at the Boyce Thompson Arboretum, near Superior, ARIZONA, on
August 23, 1953, on flowers of tamarisk.
1966
The Taxonomy and Nomenclature of Bees
27
This record adds another subgenus to the United States, and it is cer-
tainly surprising to find a species of this group, which is so decidedly tropical
in distribution, in the southwestern deserts. I know of no records of any species
of Hemisiella in any of the northern Mexican states, and it is almost a certainty
that this is an accidental intrusion. It would be interesting to know if the
species is now established in the Nearctic Region.
For the identification of this specimen I am indebted to Padre Moure.
Subgenus MELANOCENTR1S Friese
The three new species described below are all anomalous members of this
subgenus, and each presents peculiarities which are difficult to reconcile with
current concepts of Melanocentris. In the males of all three the mandibles are
tridentate (as usual in Melanocentris) , but the upper inner mandibular carina
ends at the base of the innermost tooth, rather than at the base of the second
tooth (the usual condition in Melanocentris) . In none of the three new species
does the scutellum show any indication of the rounded lobes characteristically
present in Melanocentris. Finally, all are unique in the metallic color of the
abdomen (especially pronounced in the first two species). The female of one
species lacks the large, spatulate setae on the anterior basitarsi which are
present in all other species of this subgenus. While all three of these species
agree in their divergence from typical Melanocentris, they are not closely
related to one another.
Centris (Melanocentris) strawi Snelling, new species
Figure 1, c and h
Diagnosis : This highly distinctive species is not closely allied to any
described species of Melanocentris and may be readily recognized by the
following combination of characters: the pubescence of the thoracic dorsum
is white, black elsewhere; the abdomen has strong metallic blue reflections;
the labrum and clypeus are pale whitish; the body length is much less than
that of other Melanocentris.
MALE: Integument black, that of abdomen with strong metallic blue
reflections; labrum entirely and clypeus, except black lateral and basal borders,
pale whitish; underside of flagellar segments three to eleven lighter than
flagellum elsewhere. Pubescence largely black; of labrum, whitish, except for
marginal dark hairs; a few pale hairs on dorsum of pronotum and on pronotal
lobes; all pubescence of mesoscutum, mesoscutellum and tegulae whitish;
intermixed whitish pubescence present on summit of first tergite, more
abundant laterally.
Head'. Mandibles tridentate, upper inner mandibular carina ending at
base of third tooth. Maxillary palpi four-segmented, second and third segments
approximately equal in length, fourth about half as long as third. Labrum
28
Contributions in Science
No. 112
shining, closely, moderately coarsely punctate. Clypeus duller than labrum,
median area of disc slightly raised, disc with scattered obscure punctures
laterally. Punctures of frons and sides of face closer, finer than those of clypeus,
interspaces a little more roughened. Vertex, laterad of ocelli, shining, im-
punctate; post-ocellar area with dense punctures larger than those of frons.
Distance from anterior ocellus to clypeal base about 0.6 times breadth of face
at level of clypeal base; distance between posterior ocelli a little more than
twice the distance between ocelli and eyes, the latter equal to an ocellar
diameter; distance between ocelli and posterior margin of vertex about twice
the distance between the ocelli. First flagellar segment slightly longer than
scape, shorter than following three segments combined.
Thorax : Punctures of mesoscutum, mesoscutellum, meso- and metapleu-
rae uniformly dense, about equal to those of vertex, interstices moderately
shiny, slightly roughened; propodeal integument moderately shiny, minutely
roughened, with sparse, fine punctures. Tarsal claws slightly flattened, bifid,
posterior femora about one-third as broad as long. First recurrent vein of
forewing entering second submarginal cell at end of basal third.
Abdomen : Discs of tergites with punctures distinct, fine, separated by
about a puncture diameter; tergites I-III with narrow impunctate apical
margins; impunctate apical margins of tergites IV-VI about three times as
broad as that of tergite III; ventrites with lateral punctures finer than those
of tergites, those of discs about equal to tergal punctures. Pygidial plate bare,
apex truncate, with strong longitudinal depression.
Measurements : Body length (front of vertex to apex of second tergite),
1 1.0 mm; forewing length, 12.0 mm.
Holotype, male (Los Angeles County Museum of Natural History), 25
mi. E. San Luis de la Paz, 7300 ft., Guanajuata, MEXICO, July 31, 1958 (R.
Straw, No. 1463A), on Penstemon potosinus.
This species is dedicated to Dr. Ralph Straw, California State College at
Los Angeles, who has kindly consented to have the type deposited in the Los
Angeles County Museum.
Centris (Melanocentris) ruthannae Snelling, new species
Figure 1, d and i
Diagnosis'. Although superficially similar to the following species, C.
ruthannae may be recognized by the more pronounced metallic blue reflections
of the abdomen. The female has the mandibles tridentate, rather than quadri-
dentate as in other Melanocentris. The male mandibles have the upper inner
carina ending at the base of the innermost tooth; this, together with the metallic
abdominal reflections and the bright lemon yellow clypeus and labrum should
be sufficient for its recognition.
MALE: Integument, except as noted below, of head, thorax and ab-
1966
The Taxonomy and Nomenclature of Bees
29
domen black, that of tergites and ventrites with strong deep blue reflections.
Labrum, clypeus (except lateral infuscations along margins); paraocular areas
and transverse supraclypeal mark dull lemon-yellow. Tegulae, medio- and
distitarsi darkly rufescent. Pubescence black, except for triangular patch of
appressed pale pubescence on base of mandibles and that of labrum which is
whitish. Wings hyaline, with strong brownish infuscation, reflecting purplish,
veins and stigma darkly rufescent.
Head: Broader than long, inner orbits slightly converging above. Maxil-
lary palpi five-segmented, third segment longest, almost as long as second plus
fourth; fourth segment twice as long as fifth. Mandibles tridentate, apical tooth
longest; inner larger than middle, broadly triangular. First flagellar segment
about equal to scape, equal to following three segments combined. Labrum
dull, coarsely, closely punctate. Clypeus dull, coarsely (sometimes striately)
punctate; disc with narrow median impunctate raised line. Paraocular and
supraclypeal areas almost impunctate, shiny. Frons and vertex finely punctate,
with shining interstices, except for narrow, sparsely punctate band along inner
orbits, beginning at level of posterior ocelli and running to tops of eyes. Dis-
tance between antennal sockets about one and one-half times a socket diameter;
Ocellar-clypeal distance less than transfacial distance at level of clypeal base.
Distance from posterior ocelli to eyes equal to an ocellar diameter; posterior
ocelli separated by two and one-half times an ocellar diameter; distance be-
tween posterior ocelli about four-fifths distance between ocelli and posterior
margin of vertex.
Thorax : Mesoscutum and mesoscutellum coarsely, closely punctate, with
shining interstices. Mesopleura discally closely punctate, the punctures equal
to those of mesoscutum; punctures becoming finer and a little sparser posteri-
orly, closer, larger and shallower sternally, appearing almost rugose. Metano-
tum dull, impunctate, tessellate; basal area of propodeum dull, impunctate,
tessellate; lateral areas finely punctate dorsally, shining, ventrally roughened,
with scattered punctures. Tegulae finely, closely punctate. Distitarsi slender,
elongate, equal to first and second mediotarsal segments combined.
Abdomen'. Discs of tergites shining, with sparse, piliferous punctures.
First tergite with pubescence long, erect; discs of second to fourth with
abundant short, suberect pubescence, not concealing surface; pubescence
progressively longer on succeeding segments. Pseudopygidial area converging
slightly toward the broadly rounded apex, disc slightly concave. Ventrites with
pubescence and punctation similar to those of tergites. Terminalia as illustrated.
Measurements'. Body length (front of vertex to apex of second tergite)
12.6 to 15.0 mm.; forewing length, 13.0 to 15.0 mm.
FEMALE: Integument black, of abdomen with strong bluish reflections;
most of apical tooth, spots on other two mandibular teeth, yellow-orange.
Wings strongly infuscated with brownish, veins and stigma black. Pubescence
entirely dark brownish; coarse erect seta-like hairs shining black.
30
Contributions in Science
No. 112
Head : Mandibles long, strongly curved at apical three-fifths, distinctly
tridentate, apical tooth longest, the broad inner tooth smallest, inner mandib-
ular carina ending at base of inner tooth. Maxillary palpi four-segmented,
second segment a little shorter than third, fourth about equal to basal. Labrum
shining, densely, coarsely punctate, apical margin rounded. Clypeus with disc
dull, roughened, with large scattered punctures, apico -median portion espe-
cially strongly tessellate and dulled; lateral and basal areas shining, with coarse,
close punctures, from which arise long, slender, simple seta-like hairs, espe-
cially laterally. Face shining, with punctures finer than those of clypeus, densest
on frons, laterally separated by about a puncture diameter. Facial quadrangle
about as broad as long. Distance from anterior ocellus to clypeal base 0.5
times breadth of face at level of clypeal base; interocellar distance 2.8 times
diameter of a lateral ocellus, about 1.5 times ocellar-ocular distance; distance
from lateral ocelli to posterior margin of vertex 2.5 times an ocellar diameter.
First flagellar segment longer than scape, a little longer than following three
segments combined.
Thorax : Mesoscutum and mesoscutellum dull, the integument strongly
roughened, obscuring the coarse punctures; meso- and metapleurae shinier,
with coarse, close punctures; mesoscutellum not at all bilobed; metanotum
dull, strongly tessellate, with a few scattered punctures. Basal area of pro-
podeum tessellate, somewhat shining, with scattered punctures; disc with
coarser, denser punctures; lateral areas densely tessellate, with dense, obscure,
minute punctures. Legs normal for the subgenus. Second recurrent vein of
forewing entering second submarginal cell at end of basal third.
Abdomen'. Tergites I, apical half of II, III— V with integument shining,
with abundant piliferous punctures, the punctures becoming progressively
sparser and coarser on succeeding segments; basal half of tergite II conspicu-
ously duller, integument slightly roughened, with contiguous shallow, poorly
defined punctures from each of which arises a short, plumose hair, rendering a
somewhat velvety appearance. Ventrites dull, granulate, with moderately
coarse, dense punctures; apical margin of ventrite III slightly produced medi-
ally; ventrite VI with strong, high, longitudinal carina on apical half. Pygidial
plate essentially flat, surface dull, apex truncate.
Measurements'. Body length (front of vertex to apex of second tergite),
14.8 to 15.2 mm.; forewing length, 13.2 to 13.5 mm.
Holotype male, allotype female (Los Angeles County Museum of
Natural History), Madera Canyon [Santa Rita Mts.], ARIZONA, no date
( J. A. Comstock) . Paratypes : 1 $ , Baboquivari Cyn., west end of Baboquivari
Mts., Arizona, July 25-27, (H. B. Leech and J. W. Green; CAS) ; 1 9 , 5 mi. E.
Continental, Arizona, August 29, 1961 (F. G. Werner; UA), on Kallstroemia.
Paratypes returned to their respective institutions.
I am very happy to be able to dedicate this remarkable and distinctive new
species to my wife.
1966
The Taxonomy and Nomenclature of Bees
31
Centris (Melcmocentris) anomala Snelling, new species
Figure 1, e and j
Diagnosis : This species, although superficially similar to C. ruthannae,
differs radically from this and all other Melanocentris in that the female lacks
the long, spatulate setae on the anterior basitarsi. Both sexes may be immedi-
ately recognized by the five-segmented maxillary palpi (four-segmented in
other Melanocentris ) .
FEMALE: Integument black, of abdomen faintly reflecting dark metallic
blue. Wings strongly infuscated with brownish, reflecting purple, veins and
stigma black; all pubescence black.
Head : Mandibles quadridentate, two inner teeth about equal in size, dull
orange-yellow. Maxillary palpi five-segmented, apical segment a little shorter
than basal, second and third longest, each longer than combined length of last
two. Labrum shining, rugosely punctate, punctures close; apex broadly
rounded. Clypeus strongly protuberant, with slight bulge along median line,
apical one-fifth flattened; median basal area quite shiny, impunctate; discal
punctures coarse, elongated, well separated, with area of median bulge a little
more shiny than disc, punctures round, sparse. Punctures of face finer, sparser
than of clypeus; of frons finer than of clypeus, close, except for median im-
punctate line; supraclypeal area very shiny, sparsely punctate; punctures of
vertex close, finer than of clypeus; ocellar area shiny, sparsely punctate. Dis-
tance from anterior ocellus to base of clypeus about one-half breadth of face
at level of clypeal base; distance between posterior ocelli equal to distance
between ocelli and eyes, about twice an ocellar diameter. First flagellar seg-
ment longer than scape, longer than following three segments combined.
Thorax : Punctures of mesoscutum, mesoscutellum, meso- and meta-
pleura, uniformly close, about equal to those of vertex, interstices moderately
shiny; mesoscutum with impunctate median line which broadens slightly, in
front of posterior margin. Metanotum and basal area of propodeum moder-
ately shiny, tessellate, much more sparsely punctate than lateral areas of
propodeum, where punctures are about a diameter apart, becoming a little
denser laterobasally. Front and middle tibiae and tarsi very densely covered
with short, compact hairs, the setae of these tibiae sparse, not flattened. Scopa
of hind legs very dense and compact, completely obscuring surface; basitibial
plate twice as long as greatest breadth, disc distinctly depressed beyond the
transverse margin of the secondary plate, the depression shining, in contrast
to duller areas.
Abdomen : Punctures fine, close, piliferous, interstices moderately shiny,
discs strongly pubescent, pubescence not obscuring surface, except on last
segment. Pygidial plate narrowly rounded at apex, with triangular raised area
on basal third.
Measurements'. Body length (front of vertex to apex of second tergite),
13.5 to 15.0 mm.; forewing length, 13.5 to 15.0 mm.
32
Contributions in Science
No. 112
MALE: Integument as in female. In some males there are two separated,
cuneiform pale yellow maculae on apical margin of clypeus. Pubescence of
head, thorax and abdomen very dark brown, appearing black, except lighter
areas on genae, behind vertex, basal tergite, underside of abdomen, and on legs;
dirty-white pubescence on posterior surfaces of front and middle femora, an-
terior and posterior surfaces of hind femora; pseudopygidial area fringed with
golden brown pubescence.
Head : Broader than long, eyes slightly converging above. Mandibles long,
slender, tridentate; inner tooth broad, truncate. Labrum, clypeus and rest of
face punctured as in female. Maxillary palpi five-segmented, apical segment
about as long as basal, last two together a little shorter than second, which is
subequal to the third. Distance from anterior ocellus to base of clypeus less
than breadth of face at level of clypeal base; distance between antennal sockets
about twice distance between sockets and eyes; posterior ocelli about twice an
ocellar diameter apart, separated from eyes by about an ocellar diameter; dis-
tance between posterior ocelli slightly greater than distance from ocelli to
posterior margin of vertex.
Thorax : Punctation as in female. Posterior femora one-third as broad as
long; apical tarsi a little more than twice as long as greatest breadth. Cellular
area of wings densely pubescent.
Abdomen : Punctation as in female. Apical tergite with pseudopygidial
area distinct, well developed.
Measurements : Body length (front of vertex to apex of second tergite),
14.0 to 15.5 mm.; forewing length, 14.0 to 15.5 mm.
Holotype male, allotype female (California Academy of Sciences),
8 mi. S. Guadalajara, Jalisco, MEXICO, late September, 1954 (F. X. Wil-
liams). Paratypes : 16 8 8,1 $ $, same data as Holotype; 1 8, Tizapan,
Jalisco, MEXICO, Sept. 15, 1963 (D. H. Janzen; LACM). Paratypes are in
the collections of the California Academy of Sciences and the Los Angeles
County Museum of Natural History.
One of the males bears a label with the following note by Dr. Williams:
“These Hemisia ? bees common, flying often low and swiftly over ground and
at times alighting to seek $ !’
1966
The Taxonomy and Nomenclature of Bees
33
Literature Cited
Cameron, P.
1903. Descriptions of new species of Hymenoptera taken by Mr. Edward
Whymper on the “higher Andes of the equator!’ Trans. Amer. Ent. Soc.,
29:225-238.
Cockerell, T. D. A.
1923. Expedition of the California Academy of Sciences to the Gulf of Cali-
fornia in 1921. The bees (I). Proc. Calif. Acad. Sci. (4), 12:73-103.
Fox, W. J.
1899. Synopsis of the United States species of the Hymenopterous genus
Centris Fabr. with description of a new species from Trinidad. Proc.
Acad. Natl. Sci. Phila., 51:63-69.
Krombein, K. V., et al.
1958. Hymenoptera of America North of Mexico. Synoptic Catalog. First
Supplement. U. S. D. A., Agric. Monog. No. 2, 305 pp.
Mitchell, T. B.
1962. Bees of the eastern United States, II. North Carolina Agr. Exp. Sta.,
Tech. Bull., 152:1-557.
Moure, J. S.
1943. Abelhas de Batatais. Arquivas do Museo Paranaense, 3:145-203.
Snelling, R. R.
1956. Bees of the genus Centris in California. Pan-Pacific Ent., 32:1-8.
LOS
ANGELES
COUNTY
MUSEUM
CONTRIBUTIONS
IN SCIENCE
UMBER 113
December 28, 1966
A NEW PELOBATINE FROG FROM THE LOWER MIOCENE OF
SOUTH DAKOTA WITH A DISCUSSION OF THE EVOLUTION
OF THE SCAPHIOPUS-SPEA COMPLEX
By Arnold G. Kluge
j;
Los Angeles County Museum of Natural History • Exposition Park
Los Angeles, California 90007
CONTRIBUTIONS IN SCIENCE is a series of miscellaneous technical papers
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Editor
A NEW PELOBATINE FROG FROM THE LOWER MIOCENE OF
SOUTH DAKOTA WITH A DISCUSSION OF THE EVOLUTION
OF THE SCAPHIOPUS-SPEA COMPLEX
By Arnold G. Kluge1
Abstract: A new species of pelobatid frog, Scaphiopus
neuter, is described from the Lower Miocene of the Wounded
Knee area of South Dakota. The new species is a member of the
Pelobatinae and it appears to lie within the genus Scaphiopus.
Many of the characteristics of the fossil suggest that it is near the
point of divergence of the subgenera Scaphiopus and Spea. Skele-
tal variation in Recent species of Pelohates and Scaphiopus is de-
scribed as it relates to the taxonomic assignment of the fossil.
The taxonomic status of the North American pelobatine frogs, Scaphiopus
Holbrook (1836) and Spea Cope (1866), has been debated for many years.
The two groups of species are variously treated in the literature (usually with-
out comment) as a single genus, or as separate genera or subgenera; see Zweifel
(1956: 22) for a brief summary. The absence of related fossils from the early
Cenozoic, the rarity of specimens from the middle Cenozoic (Oligocene-
Miocene), Auffenberg (1956), and the fragmentary nature of the known fossil
material from the late Cenozoic (Pliocene-Pleistocene), Bayrock (1964),
Brattstrom (1964), Estes and Tihen (1964), Gehlbach (1965), Gut and Ray
(1963), Holman (1958, 1959a, 1959b), Lynch (1965), Mecham (1959),
Taylor (1936, 1938, 1941, 1942), Tihen (1954, 1960), and Zweifel (1956),
has hindered the interpretation of the phylogenetic history of these taxa. I
believe that the recent discovery of a fossil from the Lower Miocene, described
herein, is a significant step toward elucidating the primary dichotomy in the
complex. Because of the apparent intermediate phylogenetic position of the
specimen between the two groups of previously recognized species it is referred
to as
Scaphiopus neuter, new species
Figures 1 through 6
Holotype: LACM 9209, collected by Harley J. Garbani on June 25, 1964;
nearly complete skeleton with the exception of the distal parts of the limbs.
Type locality: LACM 1982 (= South Dakota School of Mines V5360; see
Macdonald, 1963); in the gullies on both sides of the Sharps Cutoff road in
the N. l/i of sect. 17, T. 39 N., R. 43 W, Wounded Knee area, Shannon County,
South Dakota.
department of Zoology, The University of Michigan, Ann Arbor, Michigan, and
Research Associate in Vertebrate Paleontology, Los Angeles County Museum of
Natural History.
1
2
Contributions in Science
No. 113
Horizon: Approximately 225 feet above the base of the Sharps Formation,
Arikaree Group, Lower Miocene (see Macdonald, 1963, for complete strati-
graphic analysis).
Diagnosis: Scaphiopus neuter differs from all fossil and Recent species of
the Pelobatinae in the following combination of characteristics: (1) nine free
presacral vertebrae, (2) quadratojugal absent, (3) sternum cartilaginous (?),
(4) transverse processes of vertebrae five through nine do not appear to project
Figure 1. Stereophotograph. Dorsal view of the holotype of Scaphiopus neuter.
Actual size.
strongly anteriorly (the ninth vertebra may be exceptional), (5) coccyx fused
to sacrum, (6) maxilla and squamosal not in contact, widely separated, (7)
operculum large, (8) frontoparietal, squamosals, maxillae and nasals covered
with slight to moderate amounts of encrusting dermal bone, (9) moderately
large frontoparietal fontanelle present, inner borders of which are highly
emarginate, (10) prootic foramen completely enclosed by bone, (11) palatine
absent, (12) pterygoid process of maxilla absent, (13) dorsal protuberance of
ilium very large, (14) postsacral webbing very extensive, and (15) two
postsacral coccygeal foramina present.
Description of holotype: (Figs. 1 and 2) All of the bones of the skull are
present with the possible exception of the premaxillae which appear to have
been eroded away during fossilization. The skull is broad and very deep, with
a light encrustation of dermal bone on the frontoparietals (Fig. 3A) and
1966
New Miocene Fossil Frog
3
squamosal, and a very heavy encrustation on the posterior portion of the maxil-
laries and the posterolateral part of the remaining nasal (Fig. 4M,N). The
frontoparietals are thin and emarginate (ragged) medially, thus producing a
Figure 2. Stereophotograph. Right dorsolateral view of the holotype of Scaphiopus
neuter. Actual size.
relatively large frontoparietal fontanelle (Fig. 3B). The frontoparietals are
narrow and do not show any evidence of a lateral wing-like extension of
encrusting dermal bone, nor is there any indication of a frontoparietal boss;
both the anterior and posterior openings of the frontoparietal canal can be
seen from a dorsal view in the absence of the wing-like extensions (Fig. 3C).
The dorsal surface of the prootics is very concave. A large operculum is
present. The prootic foramen (for the passage of the trigeminal nerve) is
completely enclosed in bone. The dorsal end of the squamosal extends antero-
ventrally, but it is widely separated from the maxilla. The quadratojugal is
absent. The maxillaries are toothed, but the presence of vomerine teeth can not
be demonstrated because most of the vomers have been eroded away or are
covered with matrix. The palatine bone is absent. The remaining right nasal is
in broad contact with the maxilla (Fig. 4N). The maxilla does not possess a
shelf-like extension (pterygoid process) posteromedially. The mandible is
edentulous. There are nine free (normal) presacral vertebrae (Figs. 1 and 2),
all of which appear to be procoelous. Most of the neural arches and the dorsal
spines of the vertebrae have been eroded away. The second, third and fourth
4
Contributions in Science
No. 113
vertebrae possess long diapophyses; those of the second vertebra project
anterolaterally, the third laterally and the fourth slightly posterolaterally. The
diapophyses of vertebrae five through nine have been largely destroyed, but
there is some indication that they projected laterally, with the possible exception
of those of the ninth which may have been directed slightly anteriorly. The
sacral vertebra is completely fused to the coccyx. The sacral diapophyses have
Figure 3. Dorsal view of the frontoparietal region of the holotype of Scaphiopus
neuter. The right frontoparietal is nearly complete, while the left has largely been
eroded away. A. Region of encrusting dermal bone. B. Natural emarginations of
frontoparietal bone bordering the frontoparietal fontanelle. C. Anterior and poste-
rior openings of the frontoparietal canal. Scale equals 5 mm.
1966
New Miocene Fossil Frog
5
been almost completely destroyed but the size of their proximal remnants and
the position of the intersacral-coccygeal foramen indicate that they were
greatly dilated. The postsacral webbing appears to have been extensive; this is
Figure 4. Lateral view of the right maxilla (M), nasal (N) and pterygoid (P) of the
holotype of Scaphiopus neuter. Note the heavy layer of encrusting dermal bone on
the maxilla and nasal. Scale equals 5 mm.
suggested by the length of the lateral remnants and the presence of two
postsacral coccygeal foramina. Only the more proximal portion of the coccyx
remains; it possesses a relatively low, rounded dorsal crest. The pectoral girdle
appears to be arciferal (Fig. 5); the remaining left clavicle is robust and
strongly arched anteriorly. The sternal style does not appear to be present in
the matrix which may indicate that it was cartilaginous. Both the scapula and
the suprascapula are heavy elements. The exposed distal portion of the left
humerus possesses a well developed medial epicondyle and a very prominent,
anteriorly directed crest. There is a moderately deep depression on the inner
surface of the humerus, proximal to the medial epicondyle. The pelvis is
represented by both ilia (the proximal end of both of the ilial shafts is missing)
and the anterodorsal portions of the ischia (Fig. 6). The ilial shaft is very
robust and nearly round in cross-section; the dorsal crest, if present, is repre-
sented by only a thin line on the inner aspect of the shaft. Anterior to the
acetabular fossa on the ilial shaft is a shallow depression. The dorsal protuber-
6
Contributions in Science
No. 113
ance is extremely large and almost completely occupies the region of the
supra-acetabular depression. There appears to be no indication of a dorsal
prominence. The acetabular fossa is relatively shallow and the dorsal and
anterior parts of the acetabular ridge are very low. The ventral portion of the
acetabular ridge is moderately well developed in association with the extremely
concave sub-acetabular expansion. The posterior part of the acetabular ridge,
Figure 5. Ventral view of the left clavicle (CL) and coracoid (CO) of the holotype
of Scaphiopus neuter. Scale equals 5 mm.
Figure 6. Lateral view of the right pelvis of the holotype of Scaphiopus neuter. Note
the large dorsal protuberance (DP) on the ilium. Scale equals 5 mm.
1966
New Miocene Fossil Frog
7
situated on the ischium, appears to be well developed. The posterior ischial
depression is very deep. The exposed proximal portion of the left femur
exhibits a well developed, sharp, posteriorly directed, trochanter ridge. The
distal limb elements (including both the hands and feet) are not preserved.
Measurements of holotype (in millimeters) : Total head and body length,
85; length of skull from tip of snout to occipital condyles, 24; width of skull
between maxillae (posterior extremes), 27; depth of skull from level of
frontoparietals to distal tip of quadrate, 12; depth of the deepest part of the
body of the ilium, 9. The above measurements are only approximations
because of the incompleteness of many of the bones.
Skeletal variation in Recent members of Pelobates and the Scaphiopus-
Spea complex: Numerous examples of most of the presently recognized Recent
species in the Pelobatinae were examined (with the exception of Pelobates
syriacus, P. transcaucasicus, and P. varaldii) to more accurately determine the
intrageneric relationships of Scaphiopus neuter (see specimens examined,
below). Some of the information on variation has an important bearing on the
familial assignment of the fossil as well.
Pasteur (1958) erected the genus Pseudopelobates for the species trans-
caucasicus (previously considered a member of Pelobates) on the basis of the
following characters: four dorsal vertebrae with very long diapophyses per-
pendicular to the axis of the vertebral column, and the atlas fused to the first
dorsal vertebra. Variability of these same characters in both the Megophryinae
and the Pelobatinae (Ramaswami, 1935: 66; Zweifel, 1956; also see following
discussion) leads me to consider Pseudopelobates a synonym of Pelobates.
Basoglu and Zaloglu (1964) also consider transcaucasicus to be a member of
Pelobates. The fact that the characters used by Pasteur are considerably fewer
in number and not of the same level of change (e.g., change in length of
element versus loss of an element) as those used by Zweifel (pp. 24-5, table 1 )
to characterize Pelobates, Macro pelobates, Scaphiopus, Spea and Neoscaphio-
pus (the latter three taxa were considered subgenera) further supports this
synonymy.
Classically, the Pelobatidae2 were considered to have the usual anuran
presacral vertebral number of eight; however, recent studies (with the excep-
tion of Adolphi’s early contribution in 1895) have indicated considerable
variation in the family that encompasses seven to nine vertebrae. Tihen (1960)
presented new data and summarized all published information on vertebral
variation in Recent and fossil members of the family. His paper clearly shows
that any discussion on the number of presacral vertebrae must consider the
following points: ( 1 ) what are the numbers of the vertebrae (particularly that
of the sacrum which is used as the primary reference point in the vertebral
sequence), (2) what is the amount of postsacral involvement (postsacral
webbing) in the formation of the sacrum, and (3) what is the degree of both
-See Myers and Leviton (1962) for discussion of family name.
8
Contributions in Science
No. 113
pre- and postsacral fusions with the sacrum? The definitive coccyx is formed
by the fusion of caudal vertebrae (four in Megophrys, Griffiths, 1963). Tihen
concluded from his survey that all fusions of the last presacral vertebra with
the sacrum in the Scaphiopus-Spea group involved the eighth (last presacral)
and ninth (sacral) vertebrae. He disagreed with Ritland’s (1955) interpreta-
tion that the ninth (last presacral) and tenth (sacral — usually the first post-
sacral vertebral segment embryologically) were fused in the holotype of the
extinct Neoscaphiopus noblei. The nine presacral vertebrae exhibited by
Scaphiopus neuter and a Recent specimen of Scaphiopus intermontanus
(S 2925) strongly suggest that a reconsideration of Ritland’s interpretation is
in order. Tihen added further support to Chrapliwy’s (1956) and Zweifel’s
(1956) contention that there was a greater degree of vertebral variation (fusion
of vertebrae, number of vertebrae, and degree of postsacral webbing) in the
Pliocene progenitors of the Scaphiopus-Spea complex than there is in their
descendent forms. Furthermore, he visualized the variation as normal and as
being more markedly reduced in living members of the Scaphiopus evolu-
tionary line than in the Spea group. Since Tihen’s work, Holman (1963) has
reported on the fusion of the last presacral vertebra (presumably the eighth) to
the sacrum (both were symmetrical in form and their fusion) in a specimen of
Scaphiopus holbrookii. His total sample consisted of Pelobates fuscus subsp.
(2), Scaphiopus bombifrons (2), Scaphiopus couchii (1), Scaphiopus ham-
mondii (12), and S. holbrookii subsp. (19). Estes and Tihen (1964) referred
three fragmentary sacrococcyges from the Early Pliocene Valentine Formation
of Nebraska to Scaphiopus alexanderi. Although the unique holotype of
S. alexanderi exhibits fused eighth (last presacral) and ninth (sacrum) verte-
brae, the Valentine material does not. In those Recent species that I have
studied (see list of specimens examined, below) there is the following varia-
tion: in two out of the 89 S. couchii the eighth presacral is fused to the sacrum
(symmetrical vertebral form and fusion); in one out of the 20 S. hammondii
(already referred to by Zweifel, 1956: 27) the eighth is fused to the sacrum
(symmetrical vertebral form and fusion); in one out of the 34 S. bombifrons
the eighth is fused to the sacrum (symmetrical in vertebral form, but only
partially fused along one sacral diapophysis) ; in one out of four S. inter-
montanus there is an extra presacral vertebra (ninth) which is fused to the
sacrum (symmetrical in vertebral form and fusion). The first and second
presacral vertebrae are also completely fused to each other in a single specimen
of S. couchii (no other vertebral irregularities noted) .
Other vertebral variation that was studied in the skeletal material (infor-
mation from X-ray plates was not used) involved the degree of postsacral
webbing, the size of the postsacral nerve foramina, and the shape of the sacral
cotyle. The figured examples ( Scaphiopus hammondii, A-D, and Scaphiopus
bombifrons, E-G) given by Zweifel (1956: 29, fig. 19A-G; reproduced here as
Fig. 7) were used to categorize the variation in postsacral webbing. Table 1
summarizes the information that I have obtained; the nearly random distribu-
1966
New Miocene Fossil Frog
9
tion indicates that the degree of webbing cannot be used to characterize the
Scaphiopus or Spea groups of species.
The size of the postsacral foramen was subjectively delimited as either
small or large. Pelobates fuscus subsp. (one specimen examined) had a much
Figure 7. Ventral view of sacral vertebrae, from Zweifel, 1956. A-D. Scaphiopus
hammondii, Mariposa County, California. E-G. Scaphiopus bombifrons, Wyoming.
Scale equals 10 mm.
Table 1
Variation in Postsacral Webbing1
categories
hammondii
bombifrons
taxa (specimens examined)
A
B
C
D
E
F
G
Pelobates cultripes (3)
3
Pelobates fuscus ( 1 )
1
Scaphiopus h. holbrookii (18)
15
2
1
Scaphiopus h. hurterii (2)
2
Scaphiopus couchii (7)
3
3
1
Scaphiopus intermontanus (5)
i
1
3
Scaphiopus bombifrons (33)
5
8
2
3
8
4
3
Scaphiopus hammondii (16)
4
5
1
4
2
total
32
16
3
4
16
9
5
Categories of variation, A-G, after Zweifel (1956). See Fig. 7.
10
Contributions in Science
No. 113
larger one. Scaphiopus h. holbrookii and Scaphiopus holbrookii hurterii had
small foraminae, while all Scaphiopus couchii, Scaphiopus intermontanus and
Scaphiopus hammondii, and most Scaphiopus bombifrons had a large aperture.
Only two S. bombifrons (33 individuals examined) exhibited a small foramen.
A second, more distal, and usually much smaller, postsacral foramen is present
in all those specimens that possess more extensive postsacral webbing. Beddard
( 1907a, b) proposed that the sacrum proper is formed from two vertebrae in
Pelobates and one in Scaphiopus. Apparently, he based his conclusion on the
points of exit of the spinal nerves from the vertebral column in the region of the
sacrum; I have not been able to duplicate his observations on the material that
I have examined.
The cotyle was round in outline in all of the Pelobates examined, while
all Scaphiopus h. holbrookii, Scaphiopus holbrookii hurterii, and Scaphiopus
couchii exhibited an oval depression. In Scaphiopus intermontanus, Scaphiopus
bombifrons and Scaphiopus hammondii the cotyle was extremely variable in
shape. There was an equal tendency in these species toward roundness or
ovalness, and there was no suggestion of sexual or ontogenetic change.
In many different groups of anurans, the proximal end of the columella
articulates with a calcified cartilaginous element, the operculum (following
Eiselt’s terminology, 1942), which is located within the membranous covering
of the fenestra ovalis. The calcified element is extremely variable intertaxo-
nomically, in size and shape and its attachment to the M. opercularis. Func-
tionally, Eiselt considered the columello-operculum-M. opercularis as the
“Vibrationsleitungsorgan.” In dermestid beetle prepared skeletons of the
pelobatines that I have studied (those with the membrane of the fenestra ovalis
still intact), the operculum is either present or absent. I have considered the
element to be present when the membrane was opaque (whitish in appearance
and of a typical sesamoid composition) and absent when the membrane was
transparent (nearly so in the dried state). Even in the latter condition, there
may be a small amount of calcification, as indicated by clearing and staining
preserved specimens with Alizarin red-S, in spite of the fact that the membrane
is transparent in dermestid beetle prepared material of the same species. This
may account for the discrepancies between Eiselt’s data on the occurrence of
the operculum in fluid preserved pelobatids and that presented here. By my
method of determination, the operculum is absent in all Pelobates cultripes,
Pelobates fuscus subsp., Scaphiopus h. holbrookii and Scaphiopus holbrookii
hurterii, and present in all Scaphiopus intermontanus, Scaphiopus bombifrons
and Scaphiopus hammondii. The operculum was present (62.5%) or absent
(37.5%) in Scaphiopus couchii; when present it was extremely small and in
one specimen it was absent on one side.
One of the most conspicuous features of the skull of pelobatines is the
degree of development and distribution of encrusting dermal bone. Only its
distribution on the frontoparietal, posterior portion of the maxillae, and poste-
roventral section of the nasals is considered here because of its presence on
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New Miocene Fossil Frog
11
these areas in Scaphiopus neuter. For the purpose of this discussion, the degree
of development of the encrustation has been categorized as extensive, moderate,
limited, or absent. In Pelobates, Scaphiopus h. holbrookii, Scaphiopus hol-
brookii hurterii and Scaphiopus couchii the frontoparietal is extensively cov-
ered; there may be subtle differences between the two genera, however,
they are difficult to quantify. The dermal encrustation is absent (sensu stricto)
in Scaphiopus intermontanus, Scaphiopus bombifrons and Scaphiopus ham-
mondii (see following discussion of the frontoparietal boss). The encrustation
on the maxillae and nasals was extensive in Pelobates, limited to extensive in
S. h. holbrookii, S. h. hurterii and S. couchii (modally extensive), and absent
in S. intermontanus, S. bombifrons and S. hammondii. A single adult male
S. hammondii (S 2384) from El Toro, Orange County, California, exhibited a
limited degree of encrustation; in other specimens from the same locality, the
encrustation was absent.
The trend toward reduction of encrusting dermal bone in the Pelobatinae
has a positively correlated, and thus complementary, trend in the degree of
contact between the frontoparietal and the squamosal and the squamosal and
the maxilla. In Pelobates cultripes the frontoparietal-squamosal and the
squamosal-maxilla contacts are extensive. In Pelobates fuscus subsp. the
frontoparietal and the squamosal are not joined, and the squamosal-maxilla
union is slightly reduced in width; the frontoparietal-squamosal union is also
interrupted in all Scaphiopus. In Scaphiopus couchii the width of the zone of
contact between the squamosal and the maxilla is narrower than that in
Scaphiopus h. holbrookii and Scaphiopus holbrookii hurterii. In Scaphiopus
intermontanus, Scaphiopus bombifrons and Scaphiopus hammondii the
squamosal and maxilla are widely separated from each other. The squamosal in
these three species is greatly reduced in size and does not exhibit any obvious
degree of encrustation.
As has been recognized for many years in the pelobatines, the dorsal
enlargement of the frontoparietal bone (boss) is characteristic of only Scaphio-
pus intermontanus and Scaphiopus bombifrons. A well developed boss may
give the appearance of extra-dermal bone. Hughes (1965) used the thickness
of the boss as an index to the degree of hybridization in a mixed breeding
population of Scaphiopus hammondii and S. bombifrons from Lubbock
County, Texas. My observations indicate that there is considerable ontogenetic
change in the size of the boss; absent or only poorly developed in the small
S. intermontanus and S. bombifrons to well developed in large individuals.
Because Hughes did not take into account the parameter of growth, his data
on hybridization must be considered inconclusive. My observations do not
indicate any sexual differences in the degree of development of the boss. In
comparing individuals of the same size, the boss of 5. bombifrons usually
appeared to be larger than that of S. intermontanus.
The presence of the frontoparietal fontanelle is positively correlated with
the absence of encrusting dermal bone in pelobatines; the fontanelle is present
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No. 113
in Scaphiopus intermontanus, Scaphiopus bombifrons and Scaphiopus ham-
mondii, and absent in all other species. The formation of the fontanelle appears
to be an extension of the evolutionary trend toward reduction of encrusting
bone in the subfamily (I have assumed that the trend has proceeded from a
massive type of skull, as exhibited by Pelobates cultripes, to the delicate form
of S. hammondii) . From my field experience with Scaphiopus couchii (dermal
encrustation present, fontanelle absent) and S. hammondii (dermal encrusta-
tion absent, fontanelle present) in the southwestern United States, and from a
survey of the literature, this trend in morphology does not appear to be cor-
related with any obvious change in the substrate and burrowing habits of the
species as one might suspect. The smallest fontanelle appears to be exhibited by
S. intermontanus and the largest by S. hammondii (comparing individuals of
approximately the same size). The fact that the development of the fronto-
parietal boss obscures the overall size of the fontanelle makes this very difficult
to observe, however.
In Pelobates and Scaphiopus h. holbrookii, Scaphiopus holbrookii hurterii
and Scaphiopus couchii, a slightly to moderately well developed pterygoid
process of the maxilla is present (Table 2). This process is a posteromedial
projection that borders the lateral surface of the pterygoid bone, immediately
anterior to the plane of squamosal-maxilla contact. In Scaphiopus intermon-
tanus, Scaphiopus bombifrons and Scaphiopus hammondii the process is almost
always absent (Table 2). The marked similarity of Tables 2 and 3 appears to
be coincidental.
Table 2
Variation in the Length of the
Pterygoid Process of the Maxilla
variation
taxa (specimens examined)
absent
short
long
Pelobates cultripes (3)
3
Pelobates fuscus ( 1 )
1
Scaphiopus h. holbrookii (18)
12
6
Scaphiopus h. hurterii (2)
1
i
Scaphiopus couchii (7)
7
Scaphiopus intermontanus (4)
4
Scaphiopus bombifrons (34)
32
2
Scaphiopus hammondii (16)
13
3
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New Miocene Fossil Frog
13
Table 3
Variation in the Size of the Palatine Bone
variation
taxa (specimens examined)
absent
small
large
Pelobates cultripes (3)
3
Pelobates fuscus ( 1 )
1
Scaphiopus h. holbrookii (18)
2
16
Scaphiopus h. hurterii (2)
1
1
Scaphiopus couchii (7)
7
Scaphiopus intermontanus (4)
4
Scaphiopus bombifrons (34)
32
2
Scaphiopus hammondii (16)
13
3
The optic, oculomotor and trigeminal nerves enter the posterior quadrant
of the ocular orbit from the cranial vault from anterior to posterior, respec-
tively. In Pelobates and the Scaphiopus-Spea group the optic foramen (or its
bony emargination) is extremely large, while the occulomotor foramen is
correspondingly small (not always discernible as a discrete aperture). The
prootic foramen, for the passage of the trigeminal nerve, is highly variable in
the degree of encirclement by bone; this variation involves a well defined
morphogenetic trend. In Scaphiopus h. holbrookii and Scaphiopus holbrookii
hurterii the prootic foramen is widely emarginate anteriorly; absence or near
absence of an emargination in Pelobates suggests an even earlier stage of
evolution of the prootic foramen. Scaphiopus couchii exhibits a slightly nar-
rower emargination than that characteristic of S. holbrookii, but usually wider
than the condition exemplified by Scaphiopus intermontanus, Scaphiopus
bombifrons or Scaphiopus hammondii. The latter three species indicate the
culmination of this morphogenetic trend (n = narrowly emarginate; s = split-
like emargination; c = prootic foramen completely encircled by bone); S.
intermontanus (n = 60%, s = 20%, c = 20%), S. bombifrons (n = 17.7,
s = 29.4, c = 52.9), S. hammondii (n = 46.7, s = 13.3, c = 40). No sexual
dimorphism and little ontogenetic change in this character was noted.
As Zweifel (1956: 38, fig, 24) has already pointed out, one of the most
peculiar morphogenetic trends in Pelobates and Scaphiopus is the reduction of
the palatine (as a ridge of bone fused to the maxilla) and its replacement by a
process of vomer. In pelobatines, the palatine bone appears to be fused to the
maxilla by the time of metamorphosis; a recently transformed Scaphiopus h.
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No. 113
holbrookii (S 2494) of 12 mm. snout to vent length did not have a center of
palatine ossification separate from that of the maxilla. The process of the vomer
is usually absent in Pelobates (if indicated it is very small and widely separated
from the palatine) , while in all members of the genus Scaphiopus it is extremely
long, contacts the palatine (when present), and closely approaches the medial
margin of the inner shelf of the maxilla. The length of the process of the vomer
in Scaphiopus appears to be relatively constant from species to species, how-
ever, its replacement of the palatine is more obviously seen in its increasing
depth (this contradicts that suggested by Zweifel, 1956; see his fig. 24). In
Pelobates, the palatine is narrow, and very long and shallow, relative to that in
Scaphiopus holbrookii. In S. holbrookii and Scaphiopus couchii the palatine is
very wide and deep and only slightly shorter than that seen in Pelobates (this
differs from that stated by Zweifel, 1956; table 1 ) . The initial stages of palatine
reduction appear to be in its width, more specifically along the anterior zone
of palatine-maxilla fusion ( e.g compare S. holbrookii with S. couchii). It is
only in Scaphiopus intermontanus, Scaphiopus bombifrons and Scaphiopus
hammondii that the depth of the process is affected. Table 3 graphically
presents this trend.
Table 4
Degree of Development of the Dorsal Protuberance
of the Ilium
variation
taxa (specimens examined)
absent
small
moderate
large
Pelobates cultripes ( 3 )
3
Pelobates fuscus ( 1 )
i
Scaphiopus h. holbrookii (18)
5
9
4
Scaphiophus h. hurterii (1)
1
Scaphiopus couchii (7)
3
3
1
Scaphiopus intermontanus (5)
3
2
Scaphiopus bombifrons (34)
11
17
6
Scaphiopus hammondii (15)
3
6
5
1
The dorsal protuberance of the ilium (following the terminology proposed
by Estes and Tihen, 1964: 457) is usually absent or only slightly developed in
pelobatines (Table 4). Because of the difficulty in accurately measuring the
size of the protuberance, the degree of development, relative to its protusion
1966
New Miocene Fossil Frog
15
above the surface and the dorsal border of the supra-acetabular expansion, it
was subjectively categorized as absent, small, moderate or large (see Table 4).
The degree of development of the protuberance is clearly not sexually dimor-
phic, nor does there appear to be any ontogenetic relationship; the latter finding
differs from that suggested by Estes and Tihen (1964). The information
presented in Table 4 does not suggest any interspecific differences in Scaphi-
opus. The absence of the dorsal protuberance in the four Pelobates examined
agrees with the observations of Estes and Tihen (1964: 459) .
Taxonomic position of Scaphiopus neuter: The condition of sacrococcy-
geal fusion, markedly dilated sacral diapophyses, procoelous vertebrae and an
arciferal pectoral girdle makes the assignment of Scaphiopus neuter to the
family Pelobatidae relatively certain. The presence of a free coccyx in the
following modern families removes them from being considered closely related
to the fossil: Ascaphidae, Discoglossidae, Rhinophrynidae, Pelodytidae, Lepto-
dactylidae (including Pseudinae and Rhinodermatinae) , Bufonidae, Hylidae,
Centrolenidae, Dendrobatidae and Sooglossidae.
The coccyx is not fused to the sacrum in some members of the Pipidae
and Atelopodidae, and it is usually free in the Microhylidae, Phrynomeridae,
Ranidae, Rhacophoridae and megorphryine pelobatids (intraspecifically vari-
able in Megophrys) . Griffiths (1963: 271) recently defined the Pelobatinae on
the basis of their anchylosed sacrococcyx, whereas in the Megophryinae these
elements were not considered to be fused together. The sacrum and the coccyx
are reported to be fused in Megophrys nasuta (Beddard, 1907a; Boulenger,
1908) and occasionally fused in Megophrys major (Boulenger, 1908; Rama-
swami, 1935). Zweifel (1956: fig. 5) also showed a diagram of a Megophrys
( c.f .) monticola with a fused sacrococcyx; it must be pointed out that M. nasuta
and M. monticola have recently been considered conspecific (Inger, 1954). In
the Megophryinae, this variation appears to be limited only to the genus
Megophrys, and within it to only two or three species out of approximately 24
that are currently recognized. In the Pelobatinae, Pelobates cultripes does not
exhibit a completely anchylosed sacrococcyx in the sense that it occurs in
Pelobates fuscus and in the genus Scaphiopus. The union of the sacrococcyx
in P. cultripes is usually indicated by a thin, ventral suture, although the two
elements can only be separated with difficulty in subadult-adult animals. The
sacrococcyx union in P. cultripes seems to represent an evolutionary stage
intermediate between the usual free condition in the Megophryinae and the
fused state in the Pelobatinae.
The presence of cylindrical or but slightly expanded, sacral diapophyses
in the following families also excludes them from being considered closely
allied to Scaphiopus neuter: Ascaphidae, Leptodactylidae, Dendrobatidae,
Ranidae and Rhacophoridae. The presence of procoelous presacral vertebrae
in the fossil suggests that it does not belong to the Ascaphidae (amphicoelous) ,
Discoglossidae, Pipidae or Rhinophrynidae (either opisthocoelous or bicon-
cave), Phrynomeridae (diplasiocoelous) , or Ranidae, Rhacophoridae or Hy-
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No. 113
peroliidae (almost always diplasiocoelous) . Boulenger (1908) found some
evidence of both procoely and opisthocoely in the genus Megalophrys (now
included in Megophrys ) and Griffiths (1963) noted free intervertebral discs in
a four-year-old adult Megophrys major (family Pelobatidae) . The apparent
presence of an arciferal pectoral girdle in the fossil also indicates that it should
not be considered a member of the Microhylidae, Ranidae, Rhacophoridae or
Hyperoliidae. The tendency towards the development of a firmisternal pectoral
girdle in the Atelopodidae and Dendrobatidae, and the firmisternal or partly
arciferal girdle of the Pipidae removes them from possible consideration as
well. The frequent reduction of the number of presacral vertebrae in the former
two families even further removes them from consideration. The only family
that completely agrees with those characters noted above for the fossil is the
Pelobatidae2, more specifically the subfamily Pelobatinae. The fossil does not
exhibit any characteristics that contradict its placement in the Pelobatidae.
Beddard (1907b) stated that the presence of two cricoid cartilages, the
more or less rudimentary condition of the metatarsal tubercle, the less com-
pletely webbed hind toes, the presence of a glandular patch on the thighs (absent
in Megophrys feae, Beddard, 1911), and the absence of the anterior hyoidean
cornua can be used to distinguish the Oriental pelobatids (= Megophryinae)
from Pelobates and presumably Scaphiopus (= Pelobatinae). This dichotomy
follows the lines of megophryine-pelobatine evolution recognized here, but
unfortunately these particular characters cannot be applied to fossil material.
Recent papers by Griffiths (1963: 258, 279) and Tihen (1965) have
strongly emphasized the importance of the number of free presacral vertebrae
(nine, or less) in delimiting ordinal-subordinal phylogency and evolutionary
trends in the Anura. Neither author mentioned the fact that discoglossids have
eight or nine vertebrae (although the latter number only rarely) and that seven
to nine vertebrae occur in the Pelobatidae (see previous discussion). This degree
of variation in “more primitive” frogs suggests that the number of vertebrae be
considered of less paleotelic weight ( sensu Camp, 1923) than that accorded to
it by Griffiths and Tihen. In my opinion, the presence of nine (normal) pre-
sacral vertebrae in Scaphiopus neuter, does not dictate that it should be placed
in the notobatrachid-ascaphid evolutionary line as outlined by Griffiths and
Tihen.
The general form of the pelvic girdle (see Zweifel, 1956: fig. 7), the fused
sacrococcyx, the reduced dermal encrustation on the skull and the presence of
a frontoparietal fontanelle, and the form, length and direction of the diapophy-
ses of the presacral vertebrae (see Zweifel, 1956: figs. 5, 14) strongly indicates
that the fossil is a member of the subfamily Pelobatinae. I consider the
Pelobatinae to include only the Recent genera Pelobates (including Pseudo-
pelobates as previously discussed) and Scaphiopus (and Spea ). The extinct
genera, Archaeopelobates Kuhn, Eopelobates Parker, Macropelobates Noble,
Miopelobates Wettstein-Westersheimb, Palaeopelobates Kuhn, and Pelobati-
nopsis Kuhn, all appear to be more closely associated with the megophryine
1966
New Miocene Fossil Frog
17
evolutionary line in the family rather than with the Pelobatinae (Wettstein-
Westersheimb, 1955; Kuhn, 1962; Griffiths, 1963; 271). The Upper Jurassic
genus Stremmia Nopsca, placed in the Pelobatidae by Reig (1958), is probably
not an amphibian (Hecht, 1960). The Lower Cretaceous pelobatids [?] from
Israel (Nevo, 1956) have not been studied in sufficient detail to be considered
here.
Both Noble (1924) and Zweifel (1956) considered the Oligocene Macro-
pelobates to be more closely related to Pelobates and Scaphiopus than to the
Megophryinae. The free coccyx, form of the greatly expanded sacral
diapophyses, length and angle of the diapophyses of the more caudal presacral
vertebrae, and the shape and relative size of the ischium and the ilium are
characteristics that Macropelobates shares with many megophryine species of
Eopelobates and Megophrys. The presence of an apparently all bony pubis
and a large prehallux (with a digging tubercle) in Macropelobates could indi-
cate that the genus is near the point of divergence of the megophryine and
pelobatine evolutionary lines. However, the Oligocene time of origin for
Scaphiopus and Spea, as discussed later, rules against this thesis.
The distribution of the quadratojugal in the Pelobatinae (present in
Pelobates, absent in Scaphiopus ) clearly indicates an early point of evolu-
tionary divergence within the subfamily as it is now recognized. The length of
the sacrococcyx (less than six presacral vertebrae in Pelobates, greater than
seven in Scaphiopus ) and the width of the sacral diapophyses (equal to the
length of four presacral vertebrae in Pelobates, two in Scaphiopus ) are slightly
variable characters, but they too support the recognition of the point of
divergence based on the distribution of the quadratojugal. The size and form
of the palatine, the presence of a bony or cartilaginous sternal style, and the
angle of the diapophyses of vertebrae five through eight to the axis of the
vertebral column (see Zweifel, 1956; table 1) are considerably more variable
(see previous discussion, above), and while they also suggest the dichotomy,
they cannot be considered absolute indicators of it. The absence of the quad-
ratojugal, the probable cartilaginous sternal style, and the only slightly angu-
lated diapophyses of vertebrae five through nine in the fossil species neuter
strongly supports its placement in the genus Scaphiopus. Griffiths’ (1963 : 271 )
statement that all pelobatids have an ossified sternal apparatus is incorrect (see
Ramaswami, 1935: 67, and Zweifel, 1956: 24).
The presence of two natural groups of species within the genus Scaphi-
opus has been recognized for many years, and their taxonomic status has
received considerable debate. The utility of the generic category is without
question and yet the objective reality of the taxon has received ever increasing
discussion. In an attempt to form a more consistent definition of the generic
category, Inger (1954, 1958) has suggested that genera, particularly in
amphibians, should be defined on the basis of their adaptive features (“ge-
netics, developmental mechanics, behavior, ecology, etc.”) rather than, for
example, the more classical morphotype approach. From my own studies on
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No. 113
gekkonid lizards, and from similar investigations on plethodontid salamanders
(David B. Wake, pers. communication) it appears that an adaptive definition
by itself has no greater advantage over any other criterion that is founded on
evolutionary principles. From the gekko and salamander studies it has become
very obvious that adaptive parallelisms are much more common than formerly
realized and thereby rules against the adaptive criterion being the only
character used to define a genus. It is obvious that the more information we
accumulate on a group of related species, particularly from fossil forms, the
less clear the generic limits will be. It is my opinion that to continue to argue
whether the Scaphiopus and Spea groups are accorded generic or subgeneric
rank is without merit, as all higher taxonomic categories are entirely man-made
assemblages and therefore of a wholly arbitrary nature (Simpson, 1961). My
choice of considering Spea a subgenus, rather than a genus, is completely
arbitrary. Table 5 is a summary of the major characters, both morphological
and “adaptive” that distinguish the Scaphiopus and Spea evolutionary lines
(see previous discussion of osteological variation). The hybridization experi-
ments that have thus far been carried out by A. Blair (1947), Littlejohn
(1959), and Wasserman (1957; 1958) also support the recognition of this
dichotomy.
The lack of contact between the maxilla and squamosal, the completely
bone encircled prootic foramen, the presence of a frontoparietal fontanelle, the
absence of a pterygoid process of the maxilla, and the absence of a palatine
bone suggest that Scaphiopus neuter is closely related to the Spea evolutionary
line (see Table 5). In contrast to this inferred relationship, the presence of
encrusting dermal bone on the skull (although relatively limited) and the
presence of an operculum points to an affinity between S. neuter and the
Scaphiopus line. The head and body size of S. neuter may also be indicative of
a relationship with Scaphiopus (see Estes and Tihen, 1964). As has already
been discussed above), the number of vertebrae (including presacral-sacral
fusions), the degree of postsacral webbing, the size of the postsacral nerve
foramina, and the shape of the sacral cotyle are considerably more variable
than previously realized, and therefore do not appear to be useful in referring
S. neuter to either the Scaphiopus or Spea complexes. Without assigning
paleotelic values to the above noted characters, the relationships of S. neuter
appear to lie within the Spea line of divergence; however, those Scaphiopus
characteristics that it exhibits indicate that it is near the point of divergence
between the two subgenera. The Lower Miocene age of S. neuter points to at
least an Oligocene time of origin for Scaphiopus and Spea. Figure 8 pictorially
represents the proposed relationships.
The presence of a frontoparietal boss in both Scaphiopus intermontanus
and Scaphiopus bombifrons strongly suggests a closer degree of affinity than
either species has with Scaphiopus hammondii. The relationship between S.
intermontanus and S. bombifrons is also clearly exemplified by their similar
breeding calls (W. Blair, 1956; McAlister, 1959). The relatively smaller fronto-
1966
New Miocene Fossil Frog
19
Table 5
Summary of the Diagnostic Characters Distinguishing
SPEA Cope from SCAPHIOPUS Holbrook
Scaphiopus Holbrook
h. holbrookii Harlan — -Pleist. -Recent
(incl. albus Garman1)
h. hurterii Strecker2 — Pleist. -Recent
couchii (Baird) — Pleist.-Recent
(incl. rectifrenis Cope3)
war dor um Estes and Tihen — Lower
Plio. species from Florida —
Lower Mio.4
1. maxilla and squamosal in contact
2. prootic foramen widely emarginate
3. metatarsal tubercle sickle-shaped
4. frontoparietal fontanelle absent
5. dermal encrustation on skull
extensive
6. operculum usually present and
large
7. pterygoid process of maxilla
present
8. palatine present
9. adults large, up to 82 mm. snout to
vent length
10. paratoid gland present or indistinct
1 1 . vocal sac single or only slightly
divided into two compartments9
12. sound of call comes from vocal
cords proper and edges of vocal
cords9
13. call not trilled9.™
14. call slightly modulated9
15. eggs small, extensive pigmentation,
with single indistinct jelly
envelope11
16. tadpoles small, usually less than
30 mm., usually darkly colored3
17. usually occupy more mesic
environment12
Spea Cope
intermontanus (Cope)5 — Recent
bombifrons ( Cope ) — Pleist.-Recent
hammondii (Baird) — Recent
(incl. multiplicata Cope6)
diversa (Taylor) — Upper Plio.
(incl. noblei Taylor7)
pliobatracha (Taylor) — Mid. Plio.
(incl. antiqus Taylor8)
studeri (Taylor) — Mid. Plio.
alexander (Zweifel) — Lower Plio.
1 . maxilla and squamosal not in
contact
2. prootic foramen narrowly
emarginate
3. metatarsal tubercle cuneiform
4. frontoparietal fontanelle present
5. dermal encrustation on skull
absent
6. operculum absent
7. pterygoid process of maxilla
almost always absent
8. palatine almost always absent
9. adults of medium size, up to
65 mm. snout to vent length
10. paratoid gland absent
1 1 . vocal sac completely divided into
two compartments9
12. sound of call comes from edges of
vocal cords9
13. call trilled9*10
14. call considerably modulated9
15. eggs large, little pigmentation, two
or three jelly envelopes11
16. tadpoles large, up to 50 to 90 mm.,
seldom darkly colored3
17. usually occupy more xeric
environment
xfide Duellman and Schwartz (1958).
2as a subspecies of holbrookii : fide W. Blair (1958), Wasserman (1957, 1958).
3 fide Chrapliwy (1956).
4Auffenberg (1956). Presently being studied by J. Alan Holman.
5as a species distinct from hammondii : fide Chrapliwy (1956), W. Blair (1956),
McAlister (1959), McCoy (1962).
See page 20 for footnotes 6 through 12.
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No. 113
parietal fontanelle (with ragged margins) and boss in S. intermontanus may
indicate a more primitive stage of evolution than that shown by S. bombifrons.
The absence of a boss and the presence of a small frontoparietal fontanelle
(with ragged margins) in Scaphiopus neuter indicates that it may have been
phylogenetically near the point of separation between S. hammondii and the
closely related S. intermontanus and S. bombifrons.
Scaphiopus neuter was collected in association with numerous mammals:
(Macdonald, 1963) : a marsupial (Peratherium spindleri), insectivores (Ocajila
makpiyahe, Arctoryctes terrenus), rabbits (Palaeolagus hypsodus, Palaeolagus
philoi), an ischyromyid rodent (Prosciurus dawsonae), a heteromyid rodent
(Florentiamys agnewi), beavers (Palaeocastor nebrascensis, Capatanka cank-
peopi, Capacikala gradatus), canids (Nothocyon roii, Sunkahetanka geringen-
intermontanus bombifrons hammondii
neuter (Lower Miocene)
/
SCAPHIOPUS SPEA
\/
PELOBATES SCAPHIOPUS
MEGOPHRYINAE PELOBATINAE
PELOBATIDAE
Figure 8. Dendrogram showing the phylogenetic position of Scaphiopus neuter with-
in the Pelobatidae.
6the few diagnostic characters and the extreme width of the zone of integration does
not appear to warrant the further recognition of multiplicata (Chrapliwy, 1956;
Zweifel, 1956).
7my examination of the holotype of N eoscaphiopus nobiei (KUMVP 6367) con-
firms Tihen’s ( 1960) reference of that genus and species of the synonymy of diversa.
8my examination of the holotype of Scaphiopus antiqus (KUMVP 1469) confirms
Tihen’s ( 1960) reference of that species to the synonymy of pliobatracha.
Qfide McAlister (1959).
10 fide W. Blair (1955, 1956, 1958).
11 fide Hoyt ( 1960) ; not determined for intermontanus.
12some major exceptions in couchii.
1966
New Miocene Fossil Frog
21
sis, Enhydrocyon crassidens), horses (Miohippus near equinanus, Miohippus
equiceps), a rhinocerotid (Diceratherium cf. gregorii) and hypertragulids
(Leptomeryx sp., Nanotragulus intermedius). These faunal associates seem to
support Zweifel’s contention (1956: 41) that the evolution of the Spea group
may have been correlated with the establishment of a grassland.
Summary
A new species of pelobatid frog, Scaphiopus neuter, is described from the
Wounded Knee area of Shannon County, South Dakota. It was collected in the
Sharps Formation, Arikaree Group, of the Lower Miocene. Numerous skele-
tons of most of the Recent species of Pelobates and Scaphiopus were examined
to assess the intrageneric relationships of the fossil. The general form of the
pelvic girdle, the fused sacrococcyx, the reduced encrusting dermal bone on
the skull and the presence of a frontoparietal fontanelle, and the form, length
and direction of the diapophyses of the presacral vertebrae strongly indicate
that the fossil is a member of the subfamily Pelobatinae and more specifically
the genus Scaphiopus. The absence of a quadratojugal further supports its
placement in Scaphiopus. Scaphiopus neuter appears to be phylogenetically
near the point of divergence of the subgenera Scaphiopus and Spea. The Lower
Miocene age of the fossil indicates that the two subgenera probably originated
by the Oligocene.
Acknowledgments
I am greatly indebted to Dr. James R. Macdonald of the Section of
Vertebrate Paleontology, Los Angeles County Museum of Natural History
(LACM) for the opportunity to study the lower vertebrate material from the
Wounded Knee faunas of South Dakota. I am also indebted to Mr. Harley J.
Garbani of San Jacinto, California who collected the holotype of Scaphiopus
neuter, and the staff artists and photographers of the Los Angeles County
Museum of Natural History for preparing the illustrations for the present
paper. The following people have permitted me to examine their fossil, and
recent skeletal and preserved material of Pelobates and Scaphiopus, without
which the study could not have been completed: Dr. Theodore H. Eaton,
Museum of Natural History, University of Kansas (KUMVP); Dr. J. Alan
Holman, Illinois State University; Dr. Jay M. Savage (JMS) and Mr. Roy W.
McDiarmid, University of Southern California; Dr. Charles F. Walker,
Museum of Zoology, The University of Michigan (S); Dr. S. David Webb,
University of Florida Collections (UF). Dr. Richard G. Zweifel, American
Museum of Natural History, Dr. Joseph A. Tihen, University of Notre Dame,
Dr. Richard Estes, Boston University, and Dr. Peter S. Chrapliwy, Texas
Western College have contributed important information to the study. Drs.
Macdonald and Walker, and Mr. Kraig K. Adler of The University of Michi-
22
Contributions in Science
No. 113
gan have kindly read the manuscript and have offered many valuable
suggestions.
Skeletal material examined:
Pelobates
cultripes (4 specimens)— JMS 331, S 2629-31 (no locality data).
fuscus subsp. (1 specimen) — -S 1226 (Austria).
Scaphiopus
bombifrons (34 specimens) — United States: S 161 (Arizona); S 2265,
2284, 2592-9, 2600-19 (Kansas); S 123, 162-3 (Wyoming).
couchii (13 specimens) — Mexico: S 2415-6 (Coahuila); JMS 738, 740,
754, 776, 785 (Sinaloa); S 2011, 2263 (Sonora); an additional 76 preserved
specimens from the collections of the University of Southern California were
X-rayed (Baja California, Sinaloa and Sonora). United States: S 2410 (New
Mexico) ;S 156, 1593, 2358 (Texas).
hammondii (including multiplicatus) (20 specimens) — Mexico: S 2312-3
(Coahuila); S 964 (Distrito Federal); S 1674 (Durango); S 1171 (Michoa-
can) ; S 2218-9 (Oaxaca) ; S 157 (San Luis Potosi) . United States: S 159, 2409
(Arizona); JMS 415, 419, 451-2, S 2384-7 (California); S 2472 (Nevada);
S 2406 (New Mexico) .
h. holbrookii (20 specimens) — United States: S 158, 968, 1021, 1324,
2494 (Florida) ;S 974-5 (Georgia) ; S 1 174, 1260-70 (South Carolina) ; S 2493
(West Virginia) .
holbrookii hurterii (2 specimens)— United States: S 794 (no locality
data) ;S 1228 (Texas).
intermontanus (5 specimens) — -United States: S 155, 2353, 2868, 2924-5
(Utah).
Fossil material examined:
Scaphiopus antiqus Taylor (1941). Holotype KUMVP 1469. Middle Pliocene.
“Edson beds,” Ogallala Formation, Sherman County, Kansas.
Scaphiopus diversus Taylor ( 1942) . Holotype KUMVP 6368. Upper Pliocene.
Rexroad Formation, Meade County, Kansas.
Scaphiopus cf. holbrookii, Auffenberg (1956). Referred material UF 6502,
9896-9 (including material from Florida Geological Survey) and Museum
of Comparative Zoology, Harvard University (uncatalogued). Early
Miocene. Thomas Farm, Gilchrist County, Florida.
Neoscaphiopus noblei Taylor (1942). Holotype KUMVP 6367. Upper Plio-
cene. Rexroad Formation, Meade County, Kansas.
Scaphiopus pliobatrachus Taylor (1936). Holotype KUMVP 1430. Referred
material KUMVP 1431-6. Middle Pliocene. “Edson beds,” Ogallala
Formation, Sherman County, Kansas.
Scaphiopus studeri Taylor (1938). Holotype KUMVP 1478. Middle Pliocene.
“Rhino Hill Quarry,” Logan County, Kansas.
1966
New Miocene Fossil Frog
23
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New Miocene Fossil Frog
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LOS
ANGELES
COUNTY
MUSEUM
CONTRIBUTIONS
IN SCIENCE
Dumber 1 14
December 28, 1966
ADDITIONAL AVIAN RECORDS FROM THE MIOCENE OF
SHARKTOOTH HILL, CALIFORNIA
By Hildegarde Howard
Los Angeles County Museum of Natural History • Exposition Park
Los Angeles, California 90007
ADDITIONAL AVIAN RECORDS FROM THE MIOCENE OF
SHARKTOOTH HILL, CALIFORNIA
By Hildegarde Howard1
Abstract: Nineteen avian bones collected in the early
1960’s represent seven extinct species, one of which, a small
albatross, is described as new. Several specimens contribute
significant information concerning undescribed portions of the
skeleton of species previously recorded. The avifauna as a whole
is specifically distinct from those of other California marine
Miocene localities.
Introduction
Six species of birds were described from the Middle Miocene marine beds
of Sharktooth Hill, Kern County, California, by Wetmore (1930) and Miller
(1961, 1962), and three other taxa were recorded by genus only. For a com-
plete summary of previous research concerning the geology and paleontology
of the Sharktooth Hill deposits, including bibliographic references, the reader
is referred to Mitchell (1965).
In the early 1960’s collecting at Sharktooth Hill was carried on for the Los
Angeles County Museum of Natural History (LACM). Among the specimens
recovered were nineteen bird bones, the largest collection of avian material so
far obtained from the area. While these specimens add only one species to the
previous number of taxa recorded, they contribute significant additional infor-
mation concerning the described forms.
Table 1
Avian Record from Sharktooth Hill
Wetmore
1930
Compton
1936
Miller
1961 &
1962
Here
recorded
Total
Diomedea calif ornica
—
—
*1
1 ( + 1?)
3
Diomedea tnilleri n. sp.
—
—
—
*2
2
Puffinus inceptor
*1
—
—
1?
2
Puffinus prise us
—
—
*1
1 (+2?)
4
Puffinus mitchelli
1 ?a
—
*1
—
2
Morus vagabundus
*2
1
1
7 (+1?)
12
Morns sp.
— -
—
1
1
2
Presbychen abavus
*1
— .
—
1 ( + 1?)
3
Branta sp.
—
—
1
— -
1
Recurvirostra sp.
—
—
1
—
1
Totals
5
1
7
19
32
* Type description
aListed as Puffinus sp., size suggests P. mitchelli
Research Associate, Los Angeles County Museum of Natural History.
1
2
Contributions in Science
No. 114
Discussion of Species
Order Procellariiformes
Family Diomedeidae, Albatrosses
Diomedea calif ornica Miller (1962)
Figure 1, I
The type of Diomedea californica is a distal portion of tarsometatarsus
(breadth of distal end 20.6 mm.), slightly larger than that of D. albatrus (distal
breadth 19.3 mm.), but not as large as that of D. exulans (distal breadth 23.9
mm.). A distal end of humerus (LACM 16468) in the present collection is
assigned to D. californica on the basis of a similar intermediate size as com-
pared with humeri of the two Recent species noted (see Table 2) .
Other than size, distinctions from the Recent species lie in less proximal
extent of both the ectepicondylar prominence and the attachment of the
anterior articular ligament.
A badly eroded distal end of femur (LACM 7431) is so defaced that its
contours suggest either Diomedea or Pelecanus. It has the heavy proportion of
shaft found in the pelicans, but I would hesitate to introduce the genus
Pelecanus into the California Miocene record on the basis of this very ques-
tionable specimen. It is possible that it represents Diomedea californica.
Table 2
Measurements and Proportions of Albatross Humeri
(Measurements in millimeters, ratios in per cent)
D. californica
D. albatrus
D. exulans
A. Greatest breadth of distal end
B. Distance from distal notch between in-
ternal condyle and entepicondyle, to
proximal tip of attachment of anterior
27.5
26.0-26.7
31.6
articular ligament
C. Distance from distal contour of ex-
ternal condyle to proximal junction of
20.2
20.8-21.3
25.6
ectepicondylar process with shaft
Ratio of measurement B to measure-
18.2
18.8-20.7
23.0
ment A
Ratio of measurement C to measure-
73.5
79.7-81.4
81.2
ment A
66.2
71.7-77.5
72.7
Diomedea milleri, new species
Figure 1, C
Type: Proximal end of left ulna, LACM 7319, collected by J. Vlastnik,
March 28, 1964.
Locality and age: Sharktooth Hill, Kern County, Calif., LACM loc. 1655,
sect., 25, T 285, R 28E, Oil Center Quadrangle, U.S.G.S., 1954; Middle
Miocene.
Figure 1. A, B and J, Morus vagabundus Wetmore: A and J, referred humerus
LACM 7432, palmar view proximal section, and anconal view proximal and distal
sections placed together (contact missing); B, referred ulna LACM 16473, internal
view. C, Diomedea milleri, new species, type ulna LACM 7319, palmar view. D,
E, F, Presbychen abavus Wetmore, referred tarsometatarsus LACM 16466, ex-
ternal, anterior and internal views. G, Morus sp. (1), right tarsometatarsus LACM
16463, posterior view. H, Morus sp. (2) left tarsometatarsus LACM 16464, poste-
rior view. I, Diomedea californica Miller, referred humerus LACM 16467, palmar
view. All figures xl.
4
Contributions in Science
No. 114
Diagnosis: Proximal breadth of ulna across cotylae approximately 82
per cent of this dimension in D. nigripes; shaft below internal cotyla forming
distinct angle between palmar and internal surfaces; brachial impression well
depressed and distinctly delimited at its proximal tip by the prominent, broad
surface of the attachment of the anterior articular ligament; the latter attach-
ment relatively short, not extending distally beyond level of distal edge of
proximal radial depression.
Measurements: See Table 3.
Referred specimen: Proximal fragment of right tarsometatarsus, LACM
16474, collected by M. K. Hammer from LACM locality 1625, Sharktooth
Hill.
Compared with the tarsometatarsus of D. nigripes, the characters of
LACM 16474 are as follows: anterior face of shaft less depressed below
proximal cotylae; proximal foramina and attachment for tibialis anticus muscle
more proximally placed; posteriorly, two distinct converging calcaneal ridges
narrow the hypotarsal area below the proximal foramina. Broken contours
make impossible a comparison of measurements with D. nigripes; it is obvious,
however, that the fossil is smaller than in the Recent species.
Remarks: This is the smallest species of fossil albatross to be described.
It is not, however, the first fossil specimen of small albatross to be recorded.
Miller (1935) records an impression of carpometacarpus and wrist bones
from the Miocene shale of Lomita, in the Palos Verdes Hills, Los Angeles
County, California. Because of its poor preservation, the specimen was only
tentatively referred to Diomedea, with no specific assignment. Size was given
as between that of D. nigripes and D. immutabilis. Possibly the species repre-
sented is the same as here described as D. milleri.
The species is named in honor of Dr. Loye Miller, pioneer in California
paleornithology.
Table 3
Measurements (in millimeters) of Albatross Ulnae
D. milleri D. nigripes
Type
LACM Bi 1268
Breadth across proximal cotylae
14.4
17.6
Depth from internal cotyla to acromion
12.6
14.7
Breadth (anconally) below cotylae at point of greatest
prominence of attachment of anterior articular ligament
12.6
14.3
Family Procellariidae, Shearwaters
Puffinus priscus Miller (1961)
A distal fragment of right humerus (LACM 16469) is assigned to Puffinus
priscus based on its resemblance to the type in shape of ectepicondylar process
and height of the process above the distal end (10.1 mm. in type, 10.3 mm. in
LACM 16469). Two humeral shafts, one near the distal end (LACM 16476),
1966
Miocene Fossil Birds
5
the other more central, but including the tip of the deltoid crest (LACM
16475), are tentatively assigned to P. priscus on the basis of similarity of
proportions of the shaft.
Puffinus ? inceptor Wetmore (1930)
A fragment of humerus near the proximal end (LACM 6972) represents
a shearwater. The specimen lacks the head, both tuberosities, median crest
and bicipital crest and most of the deltoid crest. The contour of the shaft
anconally below the pneumatic fossa is preserved, and suggests that the median
crest terminated closer to the angular ridge of the shaft than in Recent species
of Puffinus. Also the attachment of the supraspinatus muscle is more medially
placed and more deeply depressed. This portion of the humerus is not known
in the three shearwaters recorded from Sharktooth Hill, or in the other Cali-
fornia Miocene species, P. diatomicus. However, Miller (1961:400) remarks
that the distal end of the humerus (the type) of P. inceptor differs so notably
from Recent Puffinus that “Were it a bird in full flesh and plumage today, it
would doubtless be assigned to a separate subgenus at least.” The divergence
of the proximal fragment now at hand from typical Puffinus suggests the possi-
bility that it, too, may represent P. inceptor. The possibility is strengthened
by the relatively stout shaft: depth 6.5 mm., breadth 3.8 mm.; the dimensions
toward the distal end in the type, as given by Miller (1961:400) are 6.0 x
3.5 mm.
Order Pelecaniformes
Family Sulidae, Boobies and Gannets
Morus vagabundus Wetmore (1930)
Figure 1 , A, B and J
At least seven of the nine sulid specimens in the collection can be assigned
to Morus vagabundus. Previous records of the species include proximal and
distal ends of humerus, incomplete ulna, and cranium. The following speci-
mens are represented in the collection now at hand: proximal and distal ends
of two right humeri (LACM 7432 and 16467), distal end of left humerus
(LACM 13980), proximal end of left humerus (LACM 16471); two prox-
imal ends of left ulnae (LACM 16472 and 16473), proximal end of right
ulna (LACM 16470).
The distal ends of humeri conform well with the description of the type
(Wetmore, 1930:89). Of two measurable specimens, one equals the type in
breadth, the other is 1.1 mm. smaller. The ulnae are assigned on the basis
of size, in keeping with proportions noted in Recent species; all (like the
humeri) fall within the size range of Sula leucogaster. On the basis of these
specimens, the following additional skeletal characters may be recorded for
the species.
Humerus: the proximal end has more flared deltoid crest (seen in palmar
view) than in Recent sulid species, and the bicipital surface is deeply grooved
6
Contributions in Science
No. 114
at its distal edge. Anconally, the contour from the external tuberosity to the
shaft is more abrupt than in Recent specimens of Sulci or Morus; the shaft has
greater angularity than in Morus, but greater breadth below the head than in
Sula. The line of the latissimus dorsi muscle on the external side of the shaft
below the deltoid crest terminates near the anconal edge of the shaft as in
Morus (distinguished from the condition in Sula in which the line is more
centrally placed on the shaft).
These specimens of humeri allow more definite comparison with Sula
pohli from the Miocene of the San Fernando Valley, California, than was
previously possible (Howard, 1958:10). A measurement of length can be
estimated from the proximal and distal segments of specimen no. 7432, the
two totalling at least 170 mm. (See Fig. 1,1). This dimension exceeds by 20
mm. or more the humeral length of S. pohli, although breadth of distal end is
overlapping in range. Furthermore, the internal condyle in M. vagabundus is
more d 1st ally extended.
It was previously noted (Howard, 1958:10) that size range in Recent
species of sulids suggested the advisability of reviewing the relationship of
M. vagabundus to M. loxostylus of the Miocene of Maryland. The additional
specimens of humeri of M. vagabundus now at hand indicate a range down-
ward in size from the type, and place the minimum specimen (with a distal
breadth of 17.2 mm.) below the probable size range of M. loxostylus. In three
Recent specimens of Sula leucogaster, measured at the Los Angeles County
Museum of Natural History, the maximum is 17 per cent larger than the
minimum specimen. The recorded (Wetmore, 1930:90) humerus of M.
loxostylus, with a distal breadth of 21.1 mm., is 22 per cent larger than M.
vagabundus humerus LACM 13980.
Ulna: The proximal ends of ulna resemble those of Morus rather than
Sula in narrow, more depressed brachial impression, with upper end of impres-
sion deeply indented (there is some tendency to pneumaticity at this point,
although less than in Sula) ; heavier, less pointed acromion; a depression at
base of acromion; and absence of pneumatic foramen on palmar side of shaft
below external cotyla.
Morus, sp.
Figure 1, G, H
Proximal ends of right and left tarsometatarsi (LACM 16463 and 16464)
represent the family Sulidae, and, presumably, the genus Morus. Although
there is considerable specific variation in this element among Recent sulids,
certain characters of the hypotarsus appear to be reliable in distinguishing
Morus from Sula. In Morus the middle calcaneal ridge does not project as
far posteriorly as the internal and is bridged on a level with the external ridge;
the two tendinal canals are nearly equal in size. In Sula, the middle and internal
ridges are of approximately equal posterior extent and are bridged together
by a large posterior capping; the internal canal is notably larger than the
1966
Miocene Fossil Birds
7
external. Both tarsometatarsi at hand are closer to Morus in these characters,
but differ from Recent M. bassanus in having a sturdier connection between
the middle and internal ridges. Also, on the proximal surface of both tarso-
metatarsi, the intercotylar area is raised as a tuberosity as in some species
of Sula.
In breadth of proximal end, LACM 16463 is only one millimeter broader
than LACM 16464, but the posterior surfaces of the calcaneal ridges are
longer and the bone is markedly deeper anteroposteriorly through the hypo-
tarsus, even exceeding M. bassanus in this dimension (see Table 4). The
following qualitative characters of LACM 16463 also distinguish it from
LACM 16464: a diagonal line (not present in 16464) extends distally from
the external calcaneal ridge to merge medianly with the internal ridge; an-
teriorly the external border of the shaft is narrow (slightly inflated in 16464),
and a narrow flange (lacking in 16464) extends downward from the external
cotyla; the outer contour of the external cotyla is straight (rounded in 16464) .
Table 4
Measurements and Proportions of Tarsometatarsi of Morus
(Measurements in millimeters, ratios in per cent)
Fossils Recent
Breadth of proximal end
Morus sp. ( 1 )
LACM 16463
13.7
Morus sp. (2)
LACM 16464
12.7
M. bassanus
LACM 18173
14.6
Depth of proximal end from an-
terior edge of internal cotyla to
posterior tip of hypotarsus
14.2
12.6
13.6
Length of posterior surface of in-
ternal calcaneal ridge
7.2
5.9
8.6
Length of posterior surface of ex-
ternal calcaneal ridge
5.6
4.5
5.0
Ratio of depth of proximal end to
breadth of proximal end
103.1
99.1
93.3
Ratio of length of posterior surface
of internal calcaneal ridge to
breadth of proximal end
52.5
46.5
58.9
Ratio of length of posterior surface
of external calcaneal ridge to
breadth of proximal end
40.9
35.4
34.2
It is obvious that two species are represented by these tarsometatarsi, and
presumably one should be referred to M. vagabundus. The lesser antero-
posterior depth of LACM 16464 suggests that this specimen is the more likely
representative of that species. The other bone is possibly assignable to the
same species represented by the large ulna recorded by Miller (1961:401) as
Morus sp. Whether or not this species can be correlated with one of the large
sulids recorded from other California Miocene localities (as, for example,
M. lompocanus ) must await additional evidence to determine.
8
Contributions in Science
No. 114
Order Anseriformes
Family Anatidae
Subfamily Anserinae, geese and swans
Presbychen abavus Wetmore (1930)
Figure 1, D, E, F
A proximal end of tarsometatarsus (LACM 16466) is assigned to Pres-
bychen abavus. Wetmore (1930:92-93) described this species from the Shark-
tooth Hill deposits on the basis of a single distal end of tibiotarsus said to be
intermediate in size between “the largest of the Canada geese and the whistling
swan” ( Branta canadensis and Olor columbianus ) ; qualitative characters were
also noted, in comparison with Branta. Compared with proportions in Recent
skeletons of O. columbianus, the tarsometatarsus, LACM 16466, possibly rep-
resents a smaller individual than does the type tibiotarsus, but the discrepancy
would be within the range of variability to be expected in anserine birds (in
both of these leg elements, a difference of 19-20 per cent between maximum
and minimum individuals is noted in O. columbianus (see table 5). Further-
more, the fossil tarsometatarsus is less flared proximally than in living geese
or swans, and would, therefore, be expected to be narrower in proportion to
the tibiotarsus than is the case in either Branta or Olor.
Table 5
Comparison of Measurements (in millimeters) of Leg Bones of
Presbychen abavus and Olor columbianus
Presbychen abavus Olor columbianus
no. in parentheses
LACM
is number of
speci-
Type
16466
mens measured
Breadth of distal end of tibiotarsus
18.6
20.7-24.6
(4)
Breadth of proximal end of tarsometatarsus
Depth of internal side of shaft of
18.3
21.5-25.5
(4)
tarsometatarsus
9.0
8.9- 9.8a
(2)
a The specimens on which maximum and minimum breadth of proximal end were
taken do not provide a reliable measurement of depth of shaft.
While superficially resembling the comparable element in Branta, the fossil
tarsometatarsus exhibits notable differences, some more closely resembling
characters of Cygnus olor, but all sufficiently significant to indicate generic
distinction: (1) proximal end lacking marked internal flare of articular area;
(2) anterior face of shaft deeply depressed in region of proximal foramina; (3)
external border of shaft anteriorly very distinctly marked by a straight ridge
running distally from anterior edge of external cotyla; (4) internal ridge of
anterior face of shaft short, becoming indistinct toward level of middle of
attachment of tibialis anticus muscle (somewhat as in Cygnus olor); (5)
outer surface of internal calcaneal ridge depressed above foramen (closest
to Cygnus olor ; this area flat in Branta and O. columbianus) ; (6) external side
1966
Miocene Fossil Birds
9
of shaft at level of distal terminus of median calcaneal ridge, deep antero-
posteriorly as in swans (narrowing proximally in Branta) . See Table 5.
A proximal fragment of femur (LACM 16465) is tentatively assigned to
P. abavus. It is too badly eroded to show characters in detail, or to permit
precise measurements, but general appearance and size place it with the
Anserinae. The head is large, and upturned, the neck well defined; the anterior
edge of the trochanter is blunt, lacking the upcurved, pointed tip found in
Branta (more closely resembling the condition found in swans); the external
contour of the articular end has its greatest lateral extent approximately
centered with respect to anteroposterior dimensions. Like the tarsometatarsus,
the femur appears to represent an individual of smaller size than does the type
of Presbychen abavus. Miller (1961:401) records an imperfect fragment of
ulna as Branta sp., stating “it is far too small to represent Wetmore’s species”
( Presbychen abavus ). Unfortunately no reliable measurements could be pro-
vided by the specimen. Now that a tarsometatarsus and femur, distinct from
living geese, yet of smaller individuals than the type of P. abavus, can, with
reasonable certainty, be assigned to Presbychen, the assignment of the ulnar
fragment to Branta is open to question.
The fact that certain characters of the tarsometatarsus here discussed
resemble those of the Mute Swan, Cygnus olor, in contrast to the condition
found in Branta or Olor, suggests a possible North American ancestral back-
ground for this group of birds. We are reminded that Cygnus paloregonus, a
larger and closer relative of Cygnus olor, is a dominant species in the Pleisto-
cene avifauna of Fossil Lake, Oregon.
Comparison of California’s Marine Miocene Avifaunas
Avian fossils have been found in more than a dozen southern California
marine Miocene localities, not all of which have been recorded. The avifaunas
of nine of the recorded localities are presented in comparison with that of
Sharktooth Hill in Table 6. Admittedly, it is difficult to compare the dis-
associated fragments from Sharktooth Hill with the partial skeletons impressed
in shale found in most of the other deposits. The latter specimens, although
more spectacular and more revealing of proportions of one element to another,
provide scant information as to the detailed characters of the articular ends,
which form the chief means of identification of the Sharktooth Hill material.
Sharktooth Hill is presumably older than any of the other deposits, with
the exception of the locality in Tepusquet Canyon, Santa Barbara County
(Middle Miocene) . All of the others are now considered to be of Late Miocene
age. The distinction in age seems to be reflected in the avifaunas. In spite of
the fact that Sharktooth Hill now boasts the greatest number of recorded
avian taxa, the possibility of specific identity with any of the forms from the
later avifaunas is entirely tentative and inconclusive (i.e., small albatross,
Diomedea milled, and the gannet, Morus sp.) . On the other hand, such species
as Puffinus diatomicus and Sula willetti, which predominate at the Lompoc
10
Contributions in Science
No. 114
locality and have been recorded, also, from other Late Miocene localities, are
clearly lacking at Sharktooth Hill, replaced by other, ecologically similar forms.
Possibly the shearwater specimen found at the Middle Miocene Tepusquet
Canyon locality (Santa Barbara Museum of Natural History specimen 319)
noted by Howard (1957: 1 ) can be correlated with Puffinus mitchelli of Shark-
tooth Hill. The wing impressions (with some fragments of bone present)
clearly indicate a bird of larger size than P. diatomicus, and the humerus is
of the same general proportions as in the type of P. mitchelli, with heavy ectepi-
condylar process. A definite assignment must await additional discoveries.
Table 6
Avifaunas of California Marine Miocene Localities
X >,
£ §
Ja
u a
2 &
£ ^
U
u
+*> ”P
§ 3
a"®
S B
o* a
r4> Cd
H co
o
O
cd
in
Cd
-O
u
o «
Cd
S C
O cd
t-J on
o
U
„ <D <L>
52 d -ti u
- o
iS u
cd co
O 22
<0
S w>
2 c
P <
a c
.2 CO ^
Cd O 5 w w — ww
Ol-J C/j l— 1 C/Dl-J W H J t— 1 1-2
O
U
CO
- s,
d
c oo
<u c
CO 00
W °
o
U
o
« OjO
cd c
f «
C co
o o
Diomedea calif ornica
X
Diomedea miller i
X
?a
Puffinus inceptor
X
Puffinus priscus
X
Puffinus mitchelli
X
?a
Puffinus diatomicus
X
X
X
Oceanodroma hubbsi
Osteodontornis orri
X
X
Phalacrocorax femoralis
X
Sula willetti
X
X
?
Sula pohli
X
Morus vagabundus
X
Morus lompocanus
X
Morus sp.
X
Miosula media
X
Paleosula stocktoni
X X
Presbychen abavus
X
Branta sp.
X
Limosa vanrossemi
X
Recurvirostra sp.
X
Cerorhinca dubia
X
o
U
CO
^ CL)
2 «
£
Ph <
c £
03 O
cd 3
1 8>
2 .2
.2 S
g-c
U co
aTentative referral of specimens recorded but heretofore specifically unassigned.
^Tentative referral of a specimen heretofore unrecorded.
Omitted from the table are two recorded localities: (1) one at Pt. Fermin,
Los Angeles County, where the only specimen was a single water-worn float
pebble containing questionably identified foot bones; and (2) one at Laguna
Hills, Orange County, from which the genus Praemancalla was recently
described (Howard, 1966); this latter locality has a large avian assemblage
that is yet to be identified in detail.
1966
Miocene Fossil Birds
11
Summary
The present study has more than doubled previous records of avian
specimens known from the Sharktooth Hill deposits. Significant information
gained includes the description of an extinct species of small albatross,
Diomedea milleri, and increased knowledge of the skeletons of previously
described species, Diomedea calif ornica, Morns vagabundus, and Presbychen
abavus.
Although ecologically the Sharktooth Hill environment presents no
marked distinction from that of the Late Miocene marine localities of Cali-
fornia, the dominant species at Sharktooth Hill are not found in the later
localities; likewise the dominant forms of the Late Miocene are absent from
Sharktooth Hill.
Acknowledgments
Without the initiative of Mr. Edward Mitchell, the excavations that
yielded the bird bones here recorded would not have been made. My thanks
are extended to him and to the several others who worked with him in the field.
Photography is by Mr. Mike Hatchimonji, through funds supplied under
a John Simon Guggenheim Memorial fellowship for study of fossil birds of
western United States.
Literature Cited
Compton, Lawrence
1936. The cranium of the Miocene gannet, Moris vagabundus Wetmore.
Proc. California Acad. Sci., ser. 4, 23(5) : 83-84.
Howard, Hildegarde
1957. A gigantic “toothed” marine bird from the Miocene of California.
Santa Barbara Mus. Nat. Hist., Bull. 1, Dept. Geol., pp. 1-23.
1958. Miocene sulids of southern California. Los Angeles County Mus.,
Cont. in Sci., 25:1-15.
1966. A possible ancestor of the Lucas Auk (Family Mancallidae) from the
Tertiary of Orange County, California. Los Angeles County Mus.,
Cont. in Sci., 101 : 1-8.
Miller, Loye
1961. Birds from the Miocene of Sharktooth Hill, California. Condor, 63:
399-402.
1962. A new albatross from the Miocene of California. Condor, 64:471-472.
Mitchell, Edward
1965. History of research at Sharktooth Hill, Kern County, California. Spec.
Publ. Kern County Hist. Soc. and County of Kern through its Museum,
vi -{- 45 pp.
Wetmore, Alexander
1930. Fossil bird remains from the Temblor Formation near Bakersfield,
California. Proc. California Acad. Sci., ser. 4, 19:85-93.
LOS
ANGELES
COUNTY
MUSEUM
CONTRIBUTIONS
IN SCIENCE
UMBER 115
December 28, 1966
LATE TERTIARY RADIATION OF VIPERFISHES
( CH AULIODONTID AE ) BASED ON A COMPARISON OF
RECENT AND MIOCENE SPECIES.
By Jules M. Crane, Jr.
Los Angeles County Museum of Natural History • Exposition Park
Los Angeles, California 90007
CONTRIBUTIONS IN SCIENCE is a series of miscellaneous technical papers
in the fields of Biology, Geology and Anthropology, published at irregular intervals
by the Los Angeles County Museum of Natural History. Issues are numbered sepa-
rately, and numbers run consecutively regardless of subject matter. Number 1 was
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change basis. Copies may also be purchased at a nominal price.
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Orders for additional copies should be sent to the Editor at the time corrected galley
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David K. Caldwell
Editor
LATE TERTIARY RADIATION OF VIPERFISHES
( CH AULIODONTID AE ) BASED ON A COMPARISON OF
RECENT AND MIOCENE SPECIES.
By Jules M. Crane, Jr.1
Abstract: The Miocene species Chauliodus eximius from
southern California is redescribed and compared with all extant
species of the genus. Postponement of the time of ossification
of the anterior vertebrae in some recent species is considered as
the major speciating mechanism in the late Tertiary. Chauliodus
sloani is proposed as the first modern species to possess this char-
acteristic. Patterns of radiation from the eastern Pacific are sug-
gested to explain the present geographical distribution of the
family.
Introduction
The family Chauliodontidae is represented by the single genus Chauliodus.
Berg (1947) places this family in the order Clupeiformes (Isospondyii) ,
suborder Stomiatoidei, superfamily Stomiatoidae (Lepidophotodermi) . Gos-
line (1960), in his revision of Clupeiformes, concurred but added the division
Clupei. Greenwood, et al. (1966) suggests that the suborder Stomiatoidei be
placed in the order Salmoniformes, superorder Protacanthopterygii, Division
III. Six species are recognized in the latest revision of the genus by Morrow
(1961) with representatives found in the temperate and tropical regions of all
oceans. These species occur at depths ranging from 20 to 2,800 meters with
the greater depths characterized by larger specimens. There appears to be a
segregation of distinct populations in different water masses (Haffner, 1952).
This has resulted in the description of a number of species and subspecies.
The first description of the genus was by Catesby (1771) who assigned
the name Vipera marina. His designation was invalidated by opinion no. 89
of the Commission for International Rules of Zoological Nomenclature which
eliminated all the systematic names devised by Catesby. The first recognized
use of the name Chauliodus was by Bloch and Schneider ( 1801 ) in describing
C. sloani.
The remaining living nominal species and subspecies were described in
the following order: Chauliodus macouni Bean (1890), C. pammelas Alcock
(1892), C. barbatus Garman (1899), C. dannevegi McCulloch (1916), C.
danae Regan and Trewavas (1929), C. sloani secundus Ege (1948), C. sloani
schmidti Ege (1948). In 1961, Morrow placed Chauliodus dannevegi and C.
sloani secundus in synonomy with C. sloani and elevated Chauliodus sloani
schmidti to specific rank.
Jordan and Gilbert (in Jordan, 1925) described a fossil specimen from
Lompoc, California as Eostomias eximius. In 1943, David recognized the
specimen as a viperfish and changed the name to Chauliodus eximius. The type
department of Biology, Cerritos College, Norwalk, California.
1
2
Contributions in Science
No. 115
specimen has been declared lost, according to Dr. George S. Myers, Division
of Systematic Biology, Stanford University, after a thorough search by the
author.
Chauliodus barbatus frazeri David (1943) was described on the basis of
one almost complete specimen and several fragments from the upper Modelo
formation (upper Miocene) of the Santa Monica Mountains, Los Angeles
County, California. A search of the literature and personal communication
with Prof. Camille Arambourg, Musee Nationale d’Histoire Naturelle, Paris,
France, indicates that no other fossil specimens of the genus were known prior
to 1959 despite the fact that fossil faunal assemblages similar in their deep sea
aspects to those in southern California have been reported from the late
Miocene of North Africa and Italy, and the Oligocene of the Carpathian
Mountains and of the Caucasus (Arambourg, 1927; Jerzmanska, 1960;
Danilchenko, 1960). Since then, 43 additional fossil specimens of Chauliodus
have been collected in southern California. This new and more complete fossil
record forms the comparative basis for this paper.
Methods and Materials
I. Recent forms
Alcohol-preserved specimens of Chauliodus barbatus (17 specimens), C.
macouni (35 specimens), C. sloani (23 specimens), C. danae (14 specimens),
C. pammelas (12 specimens), and C. schmidti (3 specimens) were X-rayed or
cleared and stained using the Hollister (1934) technique.
Satisfactory radiographs were obtained by using Kodak Industrial type
M film with two types of Soft X-ray units, G.E. Mobile 90-1 1, and Softex “B”,
KXK, Tokyo. A hospital unit, Picker “300’’ was unsatisfactory using Dupont
SL-313 and Kodak Blue Brand films because even the lowest voltage over-
exposed the negative and the film was too grainy for good definition. Since the
outlines of alcohol drops show on the film, the specimens were first blotted
dry, then positioned on a thin sheet of clear plastic laid over the loaded cassette.
Varying degrees of exposure on a single specimen were achieved either by
blocking out an area by building a bridge with a lead sheet or laying several
thicknesses of paper toweling over the tail or other portions likely to be burned
out by overexposure. The latter method was very effective and time-saving since
it eliminated the necessity for two separate exposures. Test film was exposed
to determine the optimum exposure for each size range. The results are
summarized in Table 1.
The fossil specimens were X-rayed, but the results were unsatisfactory.
Some barely visible outlines were obtained with the Softex “B” machine at
6 MA, 15 KV, 10 inches, with a two minute exposure. Further experimentation
in this area should prove fruitful.
Measurements and counts were made from the X-rayed or stained
material of those characters which may be discerned in these fossils. Such
1966
Viperfish Evolution
3
Table 1
X-ray data for the genus Chauliodus using
Kodak Industrial Type M film
Standard
Length
Milli-
Kilo-
Exposure
Time
Distance
X-ray Model
(mm.)
amperes
volts
(sec.)
(inches)
G.E. Mobile 90-11
50
5
40
20
40
175
5
40
50
40
KXK Softex-B
50
6
15
3.5
10
80
6
15
5
10
125
6
15
15
10
125
6
15
35
16
190
6
15
30
10
characters include the teeth, vertebral column, fins, and general body pro-
portions but exclude otoliths, scales, and most of the bones of the head. Photo-
phores of the ventral series appear in only two fossil specimens. In addition
to the generally accepted meristic and morphometric characters used in fishes,
the following counts and measurements were used:
1 . Number of cervical vertebrae: Counted from the second vertebra and
specifically excluding the most anterior vertebra (which possesses an enlarged
neural arch, no centrum, and an enlarged haemal arch) to that vertebra on a
vertical line drawn from the base of the first dorsal ray. (Note: All vertical
and perpendicular lines are drawn in reference to the axis of the vertebral
column.)
2. Number of acentrous vertebrae: Counted from the second cervical
vertebra, specifically excluding the first cervical vertebra, which is not discern-
ible in the fossil because it is obscured by the supracleithrum and is therefore
omitted from all counts. Only those vertebrae without any trace of ossification
of the centrum were included in this category.
3. Number of thoraco-abdominal vertebrae: Counted from, but not in-
cluding, the last cervical vertebra to that vertebra on a perpendicular line from
the first anal ray.
4. Number of caudal vertebrae: Counted from the last thoraco-abdominal
vertebra.
5. Number of vertebrae under dorsal fin: Counted between vertical lines
drawn from the base of the first dorsal ray and the base of the last dorsal ray.
6. Number of vertebrae over pelvic bone: Counted between perpendicu-
lars drawn from the anterior-most margin of the pelvic bone and from its
articulating border.
7. Number of vertebrae over anal fin: Number of vertebrae between
perpendiculars drawn from the base of the first anal ray and from the base
of the last anal ray.
8. Length of head: Distance between vertical lines from the anterior tip
of the premaxillary and to the most posterior margin of the cleithrum.
4
Contributions in Science
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9. Length of caudal peduncle: Measured from a vertical drawn from the
base of the last anal ray to the end of the hypural.
10. Length of teeth: Measured as the distance in a straight line between
the tip of the tooth and its point of emergence from the bone. No attempt was
made to follow the curvature of the tooth.
11. Standard length: Measured from the anterior margin of the pre-
maxillary to the posterior margin of the hypural.
A few of the fishes examined were so damaged by capture, preservation,
or preparation methods that data from them are incomplete. However, usable
data from these specimens were retained.
II. Fossil forms
The holotype of Chauliodus barbatus frazeri David, from California
Institute of Technology (CIT) Location No. 326 (Fig. 1) was cleaned and
examined, as were two fragments from CIT 385. Forty-three additional
specimens from Los Angeles County Museum of Natural History (LACM)
Locations Nos. 1035, 1267, 1806, 1925, 6589 and CIT 332 were collected
and measured in the manner previously described. Six of these specimens are
complete. Parts of the remaining specimens were either lost in collecting,
broken in preparation, or had been lost prior to fossilization. In some, the
bones of the head were disarticulated. It was necessary in preparing the teeth
to destroy some of the bones in the area of the mouth. If bones were scattered
or fragmented, only those components which could be measured accurately
were used. The supracleithrum generally obscured the area of the centrum
of the anterior-most vertebrae. No attempt was made to remove this bone.
Thirty of the specimens collected came from LACM No. 1267 (Fig. 1)
at the northeast end of the Santa Monica Mountains, Los Angeles County,
California, in the most easterly portion of what is now an inaccessible San
Diego Freeway (Interstate Highway 405) road cut. The remaining specimens
were unearthed at LACM No. 1806: Milbrook Road, Beverly Glen Canyon,
Santa Monica Mountains (Fig. 1); LACM No. 6589: Brush Ridge Quarry,
near Lompoc, California; LACM No. 1925: Cabrillo Beach, San Pedro, Cali-
fornia; LACM No. 1035: behind Mulholland Fire Station, Santa Monica
Mountains, Los Angeles County, California; CIT 332: Sulfur Canyon, Santa
Susana Mountains, Ventura County, California; CIT No. 385: near LACM
6589, Lompoc, California.
The fact that Chauliodus is a relatively small fish made it possible to
collect good materials using small hand tools. All of the fossils were found by
splitting the diatomaceous shale in the field and then finishing the cleaning in
the laboratory. Rocks were trimmed with a handsaw to facilitate transporta-
tion and storage. In no instance was any portion of a fossil lost or damaged
because of this technique. The greatest damage was done as the rock was split
initially, when a pick or chisel inadvertently went through part of an unseen
specimen.
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Figure 1. Map of a portion of the Santa Monica Mountains, Los Angeles County,
California, showing collecting localities of Chauliodus eximius, LACM Nos. 1267,
1806, CIT No. 326.
Location of map in Van Nuys Quadrangle U.S.G.S. 1953.
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Figure 2. Photograph of Chauliodus eximius, LACM Specimen No. 5244, showing
photophores and typical body form. Anterior vertebrae are displaced laterally.
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The fossils are thin, brittle, compressed remains, appearing brown when
laminated between the sheets of punky white diatomaceous rock in which they
are found. Frequently portions of the skin are present as an ultra-thin film
readily destroyed by contact. Two specimens show carbonized spots repre-
senting photophores (Fig. 2). Because of the delicate nature of this material,
cleaning was done under a dissecting microscope using needles with flattened
points and modified dental cleaning tools. The cleaned surface was sprayed
lightly with clear Krylon plastic spray to protect it from abrasion. Saturating
the rock with the plastic resulted in a contraction of the saturated area as the
plastic hardened, causing that portion of the rock to lift off the plane beneath
it. This was a desirable cleaning technique for removing overlying matrix.
When too much plastic was applied to fossils lying in thinly bedded diatomite,
it caused the fossil to lift off in a thin, brittle sheet.
In order to establish a basis for comparison between extant forms which
have been extensively described (Morrow, 1961; Ege, 1948) and the fossil
species whose description was based on a single, now lost, specimen (Jordan,
1925) or on an incomplete specimen (David, 1943), it is necessary to re-
Figure 3. X-ray photograph of adult specimen of Chauliodus macouni, showing the
single acentrous cervical vertebra at the anterior most end of the vertebral column.
Figure 4. X-ray photograph of adult specimen of C. barbatus showing single acen-
trous cervical vertebra and partial ossification in vertebra immediately posterior.
Figure 5. X-ray photograph of adult specimen of C. danae showing five acentrous
cervical vertebrae.
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Figure 6. Chauliodus eximius, juvenile, 68 mm. SL, LACM specimen No. 11,440, showing greatest ossification in
caudal region.
Figure 7. Enlargement of cervical region of Figure 6.
Figure 8. Chauliodus eximius, adult, approx. 160 mm. SL, LACM specimen No. 11,441, showing apparent com-
plete anterior ossification.
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describe the fossil in terms comparable with modem data. For this purpose,
the David specimen is inadequate and the following description is therefore
based on the additional fossil specimens collected since 1959.
Results
Acentrous vertebrae: Examination of X-rays of up to 67-mm. standard
length juvenile specimens of all extant species of Chauliodus reveals that ossi-
fication of the vertebral column begins in the caudal region and procedes
anteriorly as the size of the organism increases. In C. barbatus and C. macouni
the number of acentrous vertebrae continues to decrease until a minimum of
one is reached at a standard length of about 145 mm. in C. macouni (Fig. 3)
and about 178 mm. in C. barbatus (Fig. 4). In the remaining extant species,
ossification of vertebral centra appears to cease at the fifth, sixth, or seventh
vertebra. One unusually large specimen (272 mm.) of C. sloani showed 2,
possibly 3 (specimen badly distorted in this area), acentrous vertebrae.
Chauliodus danae is used in Figure 5 as an example of a species with five
acentrous vertebrae.
In an exceptionally well preserved fossil specimen of a juvenile Chauliodus
eximus 68 mm. in standard length (LACM specimen No. 11,440), ossification
is shown to be greatest in the caudal region and progressively less as it procedes
anteriorly (Fig. 6). The area of acentrous vertebrae (Fig. 7) is the greatest
of any of the fossil specimens. In larger specimens, 86 to 161 mm. in standard
length, ossification has proceded to the point where there are 0-2 acentrous
vertebrae evident (Fig. 8). A comparison of the fossil and Recent species
with regard to acentrous vertebrae is shown graphically in Fig. 9.
Vertebral counts: A summary of cervical, thoracic and caudal counts for
all known species of Chauliodus appears in Table 2. The proportions of verte-
brae over the pelvic bone, over the anal base, and under the dorsal base for
adult C. macouni, C. bartabus, and the fossil C. eximius within the same size
range are shown in Figure 10.
Dentition: The general arrangement and size of the teeth in the fossil
specimens are typical of the family Chauliodontidae. Measurements of teeth
made of intact premaxillary and mandibular teeth in C. eximius (Specimen
Nos. LACM 5244, 5247, 5248, 5250, 5253, 5254, 5256, 5258, 5260, 5261;
LACM: CITNo. 10163; 2 uncatalogued) showed that in twelve out of thirteen
specimens in which comparison of the third and fourth premaxillary tooth
was possible, the third was longer than the fourth. In specimen No. LACM
5253, the two were equal in length. No measurements were made on the
modern species because the data are reported in the literature by Morrow
(1961 : 253). Chauliodus macouni and C. barbatus are the only living species
having the third premaxillary tooth longer or rarely equal to the fourth.
In C. eximius the first mandibular, and the second and third premaxillary
teeth show no terminal modification into a triangular expansion generally
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typical of C. sloatii, C. danae, C. pammelas and C. schmidti. Both C. barbatus
and C. macouni show only a slight terminal modification of these teeth. There
is no recurving of the teeth in C. eximius as is found moderately developed in
C. macouni and C. barbatus and distinctly in all other species of this genus.
Fin rays: The dorsal fin in all living species has a minute first fin ray
followed by an elongated filamentous second ray plus five, rarely four or six,
smaller rays. Morrow (1961: 262) describes the first ray as much produced
into a long filament, thereby omitting mention of the minute first true dorsal
ray. Chauliodus eximius seems to possess the minute first dorsal ray. It is
difficult to determine the presence of this first ray with certainty because of
its small size, proximity to the base of the filamentous second ray, and the
general state of preservation. The total dorsal ray count for C. eximius
including the presumed minute first ray is seven to eight.
Counts of the rays in the paired fins of the fossil specimens were difficult
to make accurately because the rays had split during the process of fossiliza-
tion. Estimates of the pectoral and ventral fin rays place C. eximius within the
range of the genus. The anal fin in C. eximius has 10 to 13 rays which is the
same as the range for C. macouni, C. barbatus, and C. sloani. Chauliodus
danae and C. schmidti have 10 to 12 anal rays and C. pammelas has 12 to 13
rays. No counts of caudal fin rays were made.
Table 2
Comparison of vertebral counts in Miocene and Recent species of Chauliodus.
Brackets surround data from Morrow (1961). TV = number of
specimens used for each count.
Vertebrae (without first cervical)
Species
Total
Cervical
Thoracic
Caudal
eximius
(fossil)
487/50-54
N = 16
9.2(8.5-10)
N = 22
32.2(30-35)
N = 12
9.8(9-11)
N - 13
barbatus
50-54
N = 15
12.6(10.5-14.0)
N - 14
29.7(28-32)
N = 15
10.3(9.0-11.5)
N — 15
macouni
56-60[55-61]
N = 29
10.5(8.5-12.5)
N = 27
36.8(35-39)
N = 29
10.3(10.0-12.0)
N = 27
sloani
53-58[61]
N = 20
9.2(8-11)
N = 19
36.7(33-39)
N - 19
9.4(9-10)
N = 20
danae (Gulf of Mexico)
(Peru-Chile Trench)
53-56[50-56]
N = 8
58-60
12.6(12-13)
N = 8
12.1(12-13)
32.0(30-34)
N = 8
35.2(34-36)
9.6(9-10)
N = 8
11.6(11-12)
pammelas
N = 6
51-53 [49-52]
N = 6
9.5(9-10)
N — 6
32.1(30-33)
N = 6
10.2(9-11)
N - 10
N = 11
N = 9
N = 9
5 3 [5 1-54] 8 34 11
N = 1 N = 1 N = 1 N = 1
schmidti
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3Vd83±d3A SnoaiN30V
Figure 9. Graph showing relationship between standard length and number of acentrous vertebrae in three Recent and
one fossil species of Chauliodus. Arrows indicate that the number of acentrous vertebrae exceeds eight but an exact
count is not possible. Chauliodus danae, C. pammelas, and C. schmidti of comparable size follow a pattern similar
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Measurements of length: Measurements from the snout to dorsal origin
are shown as an average percentage of the standard length in Figure 1 1 . Snout
to anal origin, length of anal base, and the length of the caudal peduncle are
similarly represented in Table 3. The snout to dorsal origin shows a steady
progression of the dorsal fin posteriorly from the fossil to Chauliodus barbatus
and C. danae. Beginning with C. macouni the trend is reversed as the dorsal
fin is advanced.
In C. danae the anal fin is more posteriorly placed than in any other, and
the fin itself is the shortest. In C. barbatus the anal fin has advanced somewhat
while the size of the fin has remained constant.
In all species the caudal peduncle has increased in length over the fossil
form. The greatest increase is in C. barbatus.
• C. macouni ■ C.barbatus A C. eximius (fossil)
Figure 10. Triangular graph comparing the proportional distribution of vertebrae
within individuals of two Recent and the Miocene species of Chauliodus. (Each
point represents a single fish whose vertebral counts in the areas represented have
been summed and each count plotted as a percentage of this sum. The coordinates
of any point on this graph, therefore, add up to 100%.)
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Table 3
Comparison of caudal region of Miocene and modern species of Chauliodus.
All figures expressed as average
percentage of standard length.
Species
Snout-anal origin
(data, except fossil,
from Morrow, 1960)
Length of
anal base
Length of caudal
peduncle
danae
86.0
6.6
7.4
barbatus
81.8
8.8
9.4
eximius (fossil)
84.6
8.9
6.5
macouni
83.8
8.1
8.1
pammelas
83.0
9.0
8.0
sloani
84.6
7.5
7.9
schmidti
84.8
8.2
7.0
Discussion
Acentrous vertebrae: Tchernavin (1953: 23) states, “In Chauliodus (as
in many Stomiatoids) the anterior part of the notochord, corresponding
probably to seven vertebrae, persists through life . . Counts of acentrous
vertebrae from X-rayed and cleared specimens of extant species of Chauliodus
indicate that this statement is not applicable to C. barbatus or C. macouni,
since the extent of ossification is a function of size in these species (Fig. 9).
Examination of juvenile stages of these two species shows that ossification
occurs first in the anal region and proceeds anteriorly to the first cervical
vertebra which remains unossified throughout the adult life. (For a resume
of larval development, see Morrow, 1964.)
In Chauliodus sloani, C. pammelas, C. schmidti and C. danae the anterior
4 to 5 vertebrae remain unossified up to 200 mm. standard length. Specimens
of all these species in excess of 200 mm. were not available for examination,
but the one C. sloani at 272 mm. with only two vertebrae remaining unossified
suggests that this may be the ossification pattern for the others as well. Juvenile
stages of all species likewise were not available. Seemingly they would show
a growth pattern of ossification, up to the fourth or fifth cervical vertebra,
similar to that of C. barbatus and C. macouni.
The low (1 to 2) acentrous counts in Chauliodus eximius may not be
accurate because there may be no acentrous vertebrae at all. No specimen
clearly shows the complete anterior-most portion of the vertebral column
because in all specimens it is obscured by the fragile supracleithrum and dis-
located small bones. Since the count of acentrous vertebrae in X-rayed indi-
viduals of modern species is based on observable paired neural and epineural
spines, and since the position of these elements and the spacing of centra in
this material indicates that some evidence of the anterior vertebrae should be
visible in fossil specimens, it is believed that the estimates of the vertebral
number in C. eximius are fairly reliable. It is logical to assume that the Miocene
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fishes displaying ossification of the vertebrae up to at least the second cervical
vertebra, and possibly with all vertebrae ossified, represent a primitive condi-
tion and that the increase in the number of anterior acentrous vertebrae
typical of many living forms of this genus is a subsequent evolutionary
modification.
The spacing between the cervical vertebrae in all species of Chauliodus
is greater than in other portions of the vertebral column. This increases the
flexibility in the anterior portion of the body. Increasing the number and
decreasing the length of the anterior vertebrae increases flexibility even further.
The unossified notochord provides still greater flexibility than either increased
spacing or increased numbers. The feeding habits of Chauliodus involve very
great flexion of the cervical region (Tchernavin, 1953). Seemingly the reten-
tion of the acentrous condition in the adult has great survival value to
Chauliodus sloani. This species has a greater geographic distribution than any
other species of the family Chauliodontidae, and this is probably due to its
more efficient food-catching abilities — the result of the retention of the larval
acentrous condition in the cervical region.
Similarly, the acentrous cervical vertebrae in Chauliodus danae, C.
pammelas, and C. schmidti confer a feeding advantage upon these species.
Vertebral counts: Since the method used here of counting cervical,
thoracic, and caudal vertebrae is dependent upon the positions of the dorsal
Figure 11. Graph relating the number of cervical vertebrae and the position of the
dorsal fin in Chauliodus spp. The curve (y=:0.59X-3.94) represents the “best fit”
line derived by the method of least squares.
1966
Viperfish Evolution
15
fin origin and the anal fin origin, it is necessary to compare the various counts
with the positions of the referred elements. Thus, in Figure 1 1, it may be seen
that the number of cervical vertebrae has increased by at least one from the
fossil to Chauliodus macouni and by at least three from the fossil to C. barbatus.
The remaining apparent differences are the result of the movement of the dorsal
fin. The movement of the dorsal fin in C. macouni and C. barbatus plus the
increase in number of cervical vertebrae increased the flexibility of the cervical
region giving these two species a feeding advantage over C. eximius. In C.
danae flexibility reaches a maximum when acentrous vertebrae are combined
with the features of high cervical vertebral count and posterior placement of
the dorsal fin.
Chauliodus sloani, C. pammelas and C. schmidti tend to maintain or
reduce the number of cervical vertebrae but achieve flexion by reducing
cervical ossification.
The thoracic vertebrae count depends on both the position of the origin
of the dorsal fin and the position of the origin of the anal fin. In Chauliodus
barbatus, the anal fin has moved forward slightly as the dorsal has moved
posteriorly resulting in a decrease in thoracic vertebrae. Chauliodus danae,
whose dorsal fin position relative to C. barbatus, is unchanged, and whose
anal fin is most posterior, has gained thoracic vertebrae. Chauliodus macouni
and C. sloani whose dorsal fin has moved forward have relatively fixed anal fin
origins. Each appears to have gained at least four thoracic vertebrae, but
proportionately the gain is twice as great in C. macouni as in C. sloani. This
gain is reflected in their generally higher total vertebral counts.
Chauliodus pammelas and C. schmidti have a more anteriorly placed
dorsal fin than the fossil, but their anal fin placement is the same. Chauliodus
pammelas, however, shows no increase in thoracic vertebrae and C. schmidti
only a slight increase.
The number of caudal vertebrae remains essentially the same in all species
of Chauliodus. This may relate to the fact that ossification is completed first
in this region.
Since the number of cervical vertebrae varies directly with the position
of the dorsal fin and since the number of caudal vertebrae remains constant,
it must be the vertebrae between the dorsal and anal fin that account for the
variation in vertebral counts among the species of Chauliodus.
Dentition: The significance of the fact that the third premaxillary tooth
is longer than the fourth in C. eximius, C. barbatus, and C. macouni may be
related to their feeding abilities. It is assumed that those species of Chauliodus
which retain the acentrous cervical vertebrae are able to open their mouths
wider because of the increased flexibility of the anterior portion of the vertebral
column. Chauliodus eximius, C. barbatus, and C. macouni all have fewer
acentrous vertebrae as adults than do the other species in the genus. This fact,
together with the correlated proportions of the third and fourth teeth, makes
it plausible to assume that they cannot open their mouths wide enough for
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the fourth tooth to be effective. Chauliodus sloani, C. danae, C. pammelas and
C. schmidti retain the acentrous condition as an adult and the fourth pre-
maxillary tooth is longer than the third, presumably because it is able to play
a greater role in food-getting since the mouth can be opened wider. The fourth
premaxillary tooth is shorter in C. eximius, C. macouni, and C. barbatus
because an excessively long tooth at that position in the jaw would be ineffective
as a food-getting instrument in a fish that cannot open its mouth very wide.
Measurements of length: The position of the dorsal fin suggests that two
trends exist in this genus. In one, as exemplified by Chauliodus barbatus and
C. danae, the dorsal fin is moving posteriorly, and in the other, from C.
macouni to C. pammelas to C. sloani and C. schmidti, this fin is moving
anteriorly. Part of the significance of the dorsal fin position is undoubtedly
related to the function of the elongated second dorsal ray. If, as Brandes
(1898; cited by Tchernavin, 1953) postulates, this ray bends far forward
to dangle a luminous lure to attract prey, then the position of the dorsal fin
plus the length of the second ray would play an important role in food getting.
Unfortunately, no one has ever seen a living specimen of Chauliodus feed. Still
another difficulty lies in the fact that this long filamentous ray is quite fragile
and is most often broken off during capture. Until sufficient material can be
collected to correlate the measurements of the second dorsal ray, this aspect
of the problem must lie in abeyance.
Taxonomy: David (1943: 124) suggested that Jordan’s specimen of
Chauliodus eximius from Lompoc, California, might be synonomous with the
Southern California subspecies which she named C. barbatus frazeri. With
more specimens on hand, it became apparent that all the known fossil mem-
bers of the family Chauliodontidae are indeed the same species. While David’s
name appropriately connotes a strong resemblance to C. barbatus, Jordan’s
specific name, eximius, holds priority and, therefore, must be used to designate
the fossil species. Furthermore, since the type specimen of Chauliodus eximius
has been lost, a specimen from the same formation at Lompoc, California,
must be chosen as a neotype. Because the Lompoc specimens are not as clear
as those from the Santa Monica Mountains, Los Angeles County, California,
a plesiotypic series composed of specimens from this latter locality needs to
be established (see below).
Evolution: Morrow (1961) suggests that a Chauliodus sloani-Yike form
gave rise to C. macouni, C. danae, C. barbatus, and C. sloani. Chauliodus sloani
then branched off to form C. pammelas and C. schmidti. Morrow supports this
position by pointing out that a trend exists in the increased number of pig-
mented SM organs (clusters of minute ventral photophores), in C. sloani
from the Indo-Pacific area to the Pacific Equatorial water mass with C.
barbatus as “the end result of whatever forces are at work . . .” (1961 : 284).
He also notes that the length of the barbel, which is shortest in North Atlantic
species, increases in length circum-globally until the greatest length is attained
in the mid-Pacific forms.
1966
Viperfish Evolution
17
Until a function is determined for the barbel and the SM organs, it is
difficult to assess the significance of their reduction or increase. Presumably
structures which confer upon the organism some selective advantage are
those which are retained in succeeding generations. Evolutionary trends in
this genus cannot be reconstructed reliably on the basis of structures whose
functional significance in Recent fish is unknown. Fossil evidence, however,
has in this instance provided a structure whose function can be reasonably
construed and which is observable in modern members of the genus; that is,
the nature of me vertebrae in the cervical region of Chauliodus.
It is assumed that any adaptation which would confer greater feeding
capability on an organism would give that organism a strong selective ad-
vantage over its congeners. Such an adaptation in Chauliodus was the indefinite
postponement of the ossification of the anterior vertebrae. By being more
flexible in the cervical region, those species with this adaptation should be
more successful than those retaining the primitive condition. This conjecture
is borne out by the observation that those species enjoying the widest distribu-
tion are the ones with the acentrous anterior vertebrae; i.e., Chauliodus sloani
and C. danae, while those retaining the anterior ossification characteristic of
the fossil species remain relatively restricted; i.e., C. barbatus and C. macouni
(Fig. 12). Chauliodus pammelas and C. schmidti, which have very restricted
ranges and have acentrous cervical vertebrae, are regarded as recently-evolved
species, closely allied to C. sloani, whose speciating mechanism probably
involved a physical-chemical isolation (see Haffner, 1952).
It does not seem reasonable to argue that an organism would evolve in a
direction that would cause it to lose feeding capabilities. This is especially
true in a deep sea environment where, presumably, carnivorous fishes compete
strongly for food. Within the family Chauliodontidae, therefore, evolution
could only have progressed in the direction of greater feeding capability,
marked by the acquisition of greater flexibility of the neck region.
Evidence supporting the choice of the southern California fossil species
as at least part of the ancestral stock for the family is found in two observations.
One, the Recent species most closely resembling the fossil form are found in
the Eastern Pacific. Two, in other fossil deposits in which the fauna is similar
to the southern California fossil fauna, there have been no reports of Chaulio-
dus. This is despite the fact that in some instances C. sloani is now quite
abundant in the waters near these deposits. There is no question regarding
recognition of the family in these different deposits since the other bathypelagic
fishes therein are beautifully preserved and any such preservation of Chaulio-
dus would be instantly identifiable.
If Chauliodus eximius formed the primitive stock for the family Chauli-
dontidae, then the Recent forms morphologically and geographically closest
to it should represent the chronologically oldest extant species. These forms
are C. barbatus and C. macouni which still retain the anterior ossification char-
acteristic of C. eximius and are found in the Eastern Pacific. Each exhibits the
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1966
Viperfish Evolution
19
beginning of two opposite trends in the movement of the dorsal fin relative to
its position in C. eximius. In C. barbatus the dorsal fin is more posterior while
in C. macouni it is more anterior. Both show a slight increase in the number of
cervical vertebrae. As the water temperature in the Eastern Pacific during the
late Miocene continued to decrease (Durham, 1960), that portion of the
primitive population which was able to adapt to the colder water may have
radiated into the North Eastern Pacific waters, increased the number of
vertebrae and became C. macouni. The remaining part of the population could
have contracted southward into the present tropical range of C. barbatus.
Following the trend of anterior movement of the dorsal fin, accompanied
by a favorable genetic change postponing anterior ossification, Chauliodus
macouni could have evolved to C. sloani in the Western Pacific. Once this
highly successful species was established, it probably then invaded the Indo-
Pacific area and the Atlantic Ocean. The Peru-Chile Trench population may
represent either radiation from the South Atlantic or an eastward migration
from the Western Pacific. There appear to be few, if any, barriers to the
radiation of this species.
The more posterior position of the dorsal fin in Chauliodus barbatus
suggests a close relationship to C. danae. If the trend to a more posterior
placement of the dorsal continued, accompanied by the postponement of
cervical ossification, then C. danae could quite conceivably have evolved from
C. barbatus. In this case, however, a geological isolating mechanism is postu-
lated in the form of the closure of the Bolivar Trough sometime in the late
Miocene or early Pliocene (Olssen, cited in Whitmore, 1965). If the popula-
tion of C. barbatus once occupied part of the present western Carribean area
and was continuous with the modern tropical Pacific population, it is possible
to conjecture that the mutant forms with greater cervical flexion arose in the
Carribean some time near the end of the Miocene. These were cut off from the
parent population by the development of the land bridge between North and
South America. Their feeding advantage would be great because of lack of
cervical ossification in addition to the more posterior placement of the dorsal
fin. This species could then radiate throughout the Atlantic and even around
to the Eastern Pacific and the Peru-Chile trench.
The apparent overlap in the ranges of Chauliodus sloani and C. danae
could plausibly be explained in terms of varying oxygen concentrations isolating
the populations vertically (Haffner, 1952).
An alternative radiation pattern could have been from the Eastern Pacific
around South America to the Atlantic. In this case the isolating mechanisms
might involve currents and deep trench environments.
The possibility that the fossil, Chauliodus eximius, gave rise to C. sloani
directly by the isolating mechanism postulated for C. barbatus to C. danae,
thus requiring a genetic change for lack of anterior ossification to occur only
once, cannot be overlooked. Chauliodus danae, C. schmidti and C. pammelas
would then have evolved from C. sloani. The difficulty with this interpretation
20
Contributions in Science
No. 115
is that it requires an anterior movement of the dorsal fin from C. eximius to
C. sloani then back again some 7 per cent of the standard length in order to
evolve C. danae from C. sloani. Chauliodus danae is closer to C. barbatus in
total number of vertebrae, the position of the dorsal fin and the structure of the
barbel. It is closer to C. sloani in lacking anterior ossification, in having the
third premaxillary tooth shorter than the fourth and in general morphometric
characters.
Whether Chauliodus danae evolved from C. sloani or from C. barbatus is
a problem which, at this point in time, does not seem capable of resolution.
Feeding habits: Eleven specimens of C. eximius showed evidence of fossil-
ized fish remains in the area of the stomach (Fig. 13). All such stomach con-
tents showed that C. eximius swallowed its prey headfirst. This is consistent
with reports of the feeding behavior in the modern members of the genus
(Morrow, 1961: 265).
Paleoecology: The punky, white diatomite in which Chauliodus eximius
occurs forms the major part of the upper strata of the upper Modelo formation
interpreted as upper Mohnian and lower Delmontian in age (Hoots, 1931).
This late Miocene formation, approximately 15 million years old, is exposed
on the northern slopes of the Santa Monica Mountains, Los Angeles County,
California. The upper Modelo lies conformably on the lower Modelo which
differs in composition because it is composed mainly of hard platy siliceous
shales and thinly interbedded sandstone. Megafossils of the upper member are
epipelagic and bathypelagic fishes, algal remains, and terrestrial spermatophyte
leaves. Delectopecten pedroanus Trask is the only mollusk represented. Two
crab carapaces and one bryozoan colony have been collected by the author
but no report on these has been published. Microfossils other than diatoms
include Foraminifera, Radiolaria, sponge spicules, and silicoflagellates.
Hoots (1931: 112) suggested a shallow (95 meters) water origin for at
least some of these diatomite deposits. He based his conclusion on the presence
of fossil-barren standstone intercalated with finely laminated diatomaceous
shale which, he held, could only have been formed by wave base action. How-
ever, it is now thought (Ladd, 1959) that turbidity currents are capable of
carrying sand and off-shore terrestrial remains to great depths. David (1943:
79) suggested that the upper Modelo “was laid down in the deep sea but not
at a great distance from shore” at depths of 200 to 500 meters in deep basins
occurring on the continental shelf or just off the continental shelf. Her con-
clusions were based on analyses of the fishes present in the faunal assemblage
of the upper Modelo of the Santa Monica Mountains. The deep basin interpre-
tation is supported by the physiography of the Recent seascape off the southern
California coast (Emery, 1960: 50). Sullwold (1960: 436) suggested, from
analysis of the Foraminifera, a depth of 920 meters. Interpretations suggesting
depths of 1 ,000 meters or more are consistent with the depths at which modern
species of Chauliodus occur.
Determinations of paleotemperature (Emiliani, 1954) indicate that the
1966
Viperfish Evolution
21
22
Contributions in Science
No. 115
climate during the Miocene was warmer than at present. What effect this would
have on the bathypelagic environment is uncertain. Since part of the geographic
distribution of Chauliodus may be related to water temperature (Haffner,
1952), it seems likely that even a small change in the extremes of the tempera-
ture range would exert some evolutionary pressure on a bathypelagic genus
such as Chauliodus which may have influenced its evolutionary pattern.
The preservation of articulated skeletons, the absence of benthonic
scavengers, and the evenly layered diatomite suggests a calm sterile bottom
similar to present off-shore areas described by Emery (1960: 168) : “A parallel
effect of low content of oxygen in the water is that much of the organic debris
produced near the surface reaches the bottom without having undergone much
oxidation during settling; therefore considerable oxidation continues within
the bottom sediments . . . the dissolved oxygen . . . becomes depleted” resulting
in an inhospitable benthonic environment. This statement also contains a
suggestion concerning circumstances that may have originally killed the fishes
preserved as fossils. Much work is presently under way on the study of marine
sediments which should provide data making interpretation of diatomite
deposits more accurate.
DESCRIPTION OF Chauliodus eximius (JORDAN AND GILBERT)
Synonymy: Eostomias eximius Jordan and Gilbert 1925; Stanford Univ.
Press, Univ. Ser: Biol. Sciences, 4:1, p. 13. Chauliodus barbatus frazeri David
1943; Geol. Soc. Amer., Spec. Paper 43, p. 61.
Study material: Forty-four specimens, 68 to 161 (200?) mm. standard
length. Thirty-three of these are from the upper Modelo formation of the NE
slope of the Santa Monica Mountains, Los Angeles County, California (LACM
1267, 1806, 1035, CIT 326). Seven are from Cabrillo Beach, San Pedro,
California (LACM 1925). Two are from Sulphur Canyon, Santa Suzanna
Mountains, Ventura County, California (CIT 332), and three are from the
area near Lompoc, California (LACM 6589, CIT 385).
Type designations: The type of Chauliodus eximius (Jordan and Gilbert) ,
originally Eostomias eximius Jordan and Gilbert, has been lost. Los Angeles
County Museum of Natural History (LACM) specimen No. 11044 from
Lompoc, California is hereby designated the neotype. Specimen No. 5253,
11048, 11439, and 11440 from LACM locality No. 1267, specimen No. 5242
from LACM locality No. 1806, and specimen No. 10163 from LACM locality
No. 326 (David’s type for C. barbatus frazeri) are designated plesiotypes.
Diagnosis: General characteristics are those of the family Chaulio-
dontidae. Large fang-like teeth in premaxillary and mandible; elongate body;
dorsal fin in anterior one-third with first ray minute, second ray much produced
into long filament.
Chauliodus eximius is similar to C. barbatus and C. macouni in that the
third premaxillary tooth is longer than the fourth and ossification procedes
anteriorly up to the first cervical vertebra. Chauliodus eximius differs from
1966
Viperfish Evolution
23
these species in proportional distribution of vertebrae, in having fewer cervical
vertebrae, in the position of the dorsal fin and in possessing non-recurving
teeth. It differs from all other extant species of Chauliodus in having ossifica-
tion of the anterior centra and in not having terminally modified teeth or teeth
which recurve.
Description: Proportional measurements given as average percentages of
standard length. Range of variation is shown in parenthesis.
Head; 16.8 (15 to 21), N = 14.
Distance from snout; to origin of dorsal fin 26.2 (23.0 to 29.8) ; to origin
of anal fin 85.7 (82.0 to 88.0); to origin of ventral fin 41.4 (37.1 to 45.0),
N = 12.
Pre-anal length without head; 68.3 (66 to 72), N = 13.
Origin of dorsal to origin of anal; 59.6 (56.0 to 64.0), N = 13.
Origin of ventral to origin of anal; 44.2 (41.0 to 47.0), N = 14.
Length of caudal peduncle; 6.5 (5.6 to 7.7), N = 14.
Length of mandible; 13.9 (12.6 to 16.3), N = 13.
Dorsal fin; rays 7 to 8 (includes minute first ray) , N = 23.
Anal fin; rays 10 to 13, N = 14.
Pectoral fin; length 10.9 (7 to 13.7), N = 12; rays 10 to 13,N = 20.
Ventral fin; length 16.6 (12 to 23.7), N = 10; rays 7 to 8, N = 22.
Vertebrae; (counts exclude first cervical vertebra; see Methods and Ma-
terials, above) Total 48?, 50 to 53, N = 15; acentrous 1 to 2; most often 1,
N = 29; cervical (including acentrous vertebrae) 9.2 (8 to 10), N = 22;
thoracic 32.1 (30 to 35), N= 12; caudal 9.8 (9 to 11), N = 13.
Serial photophores; Ventral row: VAV 20 or 22, N = 1; PV 17, N = 1.
Lateral row present.
Body elongate, slender, depth about 10 per cent of standard length. Barbel
not detectable. Head slightly less than 20 per cent of standard length. Orbit not
well enough defined to measure.
Mouth large, mandible length about 80 per cent of head. Premaxilla
with four teeth, second tooth longest; third longer than fourth (one specimen,
third equal to fourth) , N = 15. Many small oblique teeth on maxilla. Mandible
with up to 7 teeth, generally 6, first tooth about half the length of mandible,
longest of any in jaws. Teeth typical of the family Chauliodontidae except that
they taper to a point uninterrupted by secondary curving or terminal modifica-
tions. Distally the teeth appear translucent and the tip is black. No denticles
detectable near corner of mouth.
Dorsal fin with 7 to 8 rays, including minute first ray visible on at least
one specimen. Second dorsal ray long, filamentous, at least 33 per cent of
standard length. Ventral fin abdominal, length about 15 per cent of standard
length with 7 to 8 rays. Pelvic bones elongate, possibly fused anteriorly. Length
of pelvic bone 8 (6 to 10) per cent of standard length, N = 8. Anal fin near
caudal with 10 to 13 rays. Length of base about equal to pectoral length.
Length of caudal peduncle about 7 per cent of standard length.
24
Contributions in Science
No. 115
Size: The largest complete specimen is 161 mm. (LACM 5242) in
standard length. The size of the head of an incomplete specimen indicates that
C. eximius probably grew to at least 200 mm.
Relationships: Chauliodus eximius is more closely allied to C. harbatus
and C. macouni than any other extant species. Only these three species share
the characteristic of ossified centra in the anterior vertebrae. The third pre-
maxillary tooth is longer than the fourth in all three and in no other members
of the family Chauliodontidae. Chauliodus macouni and C. harbatus are the
only Recent species found in the eastern Pacific, which was the habitat of C.
eximius. It is likely that C. eximius was the ancestral stock for both the north
temperate C. macouni and the tropical C. barbatus. Total vertebral count is
similar in both C. eximius and C. barbatus but the distribution of cervical and
thoracic vertebrae is different. Chauliodus macouni is closer to C. eximius in
number of cervical vertebrae. Both C. barbatus and C. macouni differ from
C. eximius in number of thoracic vertebrae. Chauliodus eximius is more distinct
from C. barbatus and C. macouni in number of vertebrae over the pelvic bone
than they are from each other. The Miocene species Chauliodus eximius is
therefore assumed to be the ancestral species which gave rise to C. barbatus
and C. macouni.
Conclusions
General: The members of the genus Chauliodus have not changed
in over-all body form since the middle Miocene. Adult specimens of
Chauliodus barbatus and C. macouni have retained the primitive, as defined
by the condition in the fossil C. eximius, almost completely ossified condition
of the anterior vertebrae and also possess the primitive characteristic of having
the third premaxillary tooth longer than the fourth. The remaining extant
species have developed a tendency to postpone the ossification of the anterior-
most vertebrae and have the fourth premaxillary tooth longer than the third.
Vertebral ontogeny proceeded from the caudal region anteriorly in the
fossil, Chauliodus eximius, as it does in all recent species. Differences in
vertebral counts among the various species of Chauliodus appear to be attribut-
able to variation in the number of vertebrae in the area between the dorsal fin
and the anal fin. The caudal vertebral count is constant for all species, while
the cervical vertebral count varies directly with the position of the dorsal in
all species except the fossil which has a proportionately lower number of
cervical vertebrae.
There is not significant difference in fin ray counts among the various
species except for the anal fin in Chauliodus pammelas which tends to have
two more rays but which is still within the range for the genus.
Evolution: The acquisition of the ability to postpone the ossification of
the anterior vertebrae, thus increasing the flexibility of the cervical region and
conferring a great feeding advantage, is considered to be the major factor
influencing speciation within the genus Chauliodus since the middle Miocene.
1966
Viperfish Evolution
25
Because C. barbatus and C. macouni most closely resemble the fossil C. eximius
in lacking this ability, they are considered the more primitive recent species.
Accepting this, and using as criteria the two trends in the movement of the
dorsal fin, i.e., anteriorly versus posteriorly when compared with the fossil,
several patterns of radiation are possible. There is not, at this time, sufficient
evidence to choose one among them as the pattern and therefore, all three are
presented. In each case, C. pammelas and C. schmidti, because of their great
resemblance to C. sloani are regarded as species most recently evolved from
C. sloani. Note that evolution from C. barbatus or C. macouni requires the
selection for postponing time of ossification of anterior vertebrae. Further-
more, it is conceived that C. sloani was well established before C. danae evolved
because of the more wide spread range of C. sloani.
1 . Chauliodus eximius — > C. barbatus — » C. danae, which was isolated by
the closure of the Bolivar Trough. Chauliodus danae then carried throughout
the North and South Atlantic, and moved around South America into the
Peru-Chile Trench.
Chauliodus eximius C. macouni C. sloani at the extreme Western
Pacific range of C. macouni. Chauliodus sloani then would have radiated west-
ward and into the South Pacific. The westerly movement continued circum-
globally until C. sloani occupied all oceans.
2. Chauliodus eximius — > C. barbatus C. danae in the southernmost
part of the C. barbatus range, C. danae then radiating from the Peru-Chile
Trench into the Atlantic Ocean. The evolution of C. sloani would have pro-
ceded as in ( 1 ) above.
3. Chauliodus eximius — > C. sloani via either the Bolivar Trough route
or around South America. Chauliodus sloani would then have evolved C.
danae, isolation in this case probably being of a physico-chemical nature.
A trend is evident in the modification of the distal portion of the teeth in
that the fossil displays a simple curving to the tip, C. barbatus and C. macouni
show a slight recurving, and in the remaining species the teeth tend to expand
into a triangular modification as well as being recurved.
Paleoecology: Chauliodus eximius probably occupied much the same
niche in a Miocene deep basin environment as its related species do today.
However, further data on Recent marine sediments are needed to properly
estimate the depositional environment of the Modelo diatomite. Records of
bottom depth as well as net depth for captures of Recent bathypelagic fishes
are needed if an accurate interpretation of the depth of the Modelo basin is to
be based on magafossil evidence. The exact vertical position for any species of
Chauliodus in a specific water mass is presently not determinable from
published data.
Acknowledgments
Materials available for this study were from the collections of the Los
Angeles County Museum of Natural History; Leland Stanford University, Palo
Alto; Scripps Institution of Oceanography, United States Fish and Wildlife
26
Contributions in Science
No. 115
Service Laboratories, La Jolla and Washington, D.C.; University of Southern
California; British Museum (Natural History); Marinbiologisk Laboratory,
Charlottenlund Slot; Museum of Comparative Zoology, Harvard University;
and the University of California, Los Angeles. Thanks for the use of equipment
and specimens and for their thoughtful suggestions are tendered to: Dr. Rolf
Bolin, Stanford University; Dr. Carl Hubbs, Scripps Institution of Ocean-
ography; Dr. Vladimir Walters, Mr. Wayne Baldwin, University of California,
Los Angeles; Dr. Daniel Cohen, United States Fish and Wildlife Service
Ichthyological Laboratory, United States National Museum, Washington,
D.C.; Mr. Frederick H. Berry, Tropical Atlantic Biological Laboratory, Miami,
Florida; Dr. Theodore Downs, Dr. Shelton P. Applegate, Los Angeles County
Museum of Natural History; Dr. Henry Childs, Mr. William Keim, Cerritos
College, Norwalk, California; and Drs. George Jakway, Robert Meade, and
James Welsh, California State College, Los Angeles. A special acknowl-
edgment is due Mr. Michael Bell who collected most of the fossil specimens.
List of Specimens Studied
Chauliodus eximius : Los Angeles County Museum of Natural History,
CIT No. 10163, LACM Nos. 5242 to 5261, 10560, 11038-11044, 11047,
11048, 11228, 11239-11241, 11251, 11440, 11441, 12239, 12410, and three
uncatalogued.
Chauliodus barbatus: Scripps Inst. Oceangr., Nos. H52-363 (2), 52-390,
H52-404, H52-409, 55-229, 55-244, 55-246, 55-258 (3), 55-265, 60-215,
60-216, 60-232, 60-247, 61-215. University of Southern California, Allan
Hancock Foundation, Eltanin Sta. 53, 58 (Peru-Chile Trench).
Chauliodus macouni: Scripps Inst. Oceangr., Nos. 51-363 (9), H5508,
H6204 (2), H6204-60.60 (2), H6204-80.55; U.S. Fish Wildl. Serv. Lab.,
La Jolla, No. 224-1; Univ. So. Calif., Nos. 2943 (4), 7344 (2), 7394, 8020,
8027 (5 postlarval) , 8028, 8122, 8236; Univ. Calif. Los Angeles, No. 1036
(cleared specimen); U.S. Fish Wildl. Serv. Lab., Washington, D.C.,
uncatalogued (3).
Chauliodus sloani: U.S. Fish Wildl. Serv. Lab., Washington, D.C., un-
catalogued (20). Univ. of So. Calif., Allan Hancock Foundation, Eltanin 14
(Greenland Cruise), Eltanin 743 (2) (Peru-Chile Trench). Scripps Inst.
Oceanography, La Jolla, Calif.: SIO 65-621 (1).
Chauliodus danae: Mus. Comp. Zook, Harvard Univ., Nos. 42247 (2),
42249 (8); Univ. So. Calif., Allan Hancock Foundation, Eltanin 52, 61, 165,
741,742 (2) (Peru-Chile Trench).
Chauliodus pammelas: British Museum (Nat. Hist.) 1939.5.26.412;
1939.5.26.417; 1939.5.26.418-419; 1939.5.26.410-411 (2); 1939.5.26.413-
415 (2). Marinbiol. Lab., Charlottenlund, Dana Coll., Nos. 39091, 3909III,
3912 1 (2), 3912II, 3912III.
Chauliodus schmidti: Mus. Comp. Zool., Harvard Univ. Nos. 42082,
42097 (2).
1966
Viperfish Evolution
27
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Durham, J. W.
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1961. History of the earth. San Francisco: Freeman Co., 610 p.
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1961. Taxonomy of the deep sea fishes of the genus Chauliodus. Bull. Mus.
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Addendum
After this paper was in galley proof, a juvenile Chauliodus eximius, 50 mm. in
standard length, was collected by John E. Fitch at Lompoc, California. The speci-
men, LACM 17143, shows weak anterior ossification. The number of vertebrae in
this region is not determinable.
LOS
ANGELES
COUNTY
MUSEUM
CONTRIBUTIONS
IN SCIENCE
Nmber 116
December 28, 1966
RECOGNITION OF THE CANCELLARIID GENUS NEADMETE
HABE, 1961, IN THE WEST AMERICAN FAUNA, WITH
DESCRIPTION OF A NEW SPECIES FROM THE LOMITA
MARL OF LOS ANGELES COUNTY, CALIFORNIA
By George P. Kanakoff and James H. McLean
||
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Los Angeles County Museum of Natural History
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RECOGNITION OF THE CANCELL ARIID GENUS NEADMETE
HABE, 1961, IN THE WEST AMERICAN FAUNA, WITH
DESCRIPTION OF A NEW SPECIES FROM THE LOMITA
MARL OF LOS ANGELES COUNTY, CALIFORNIA
By George P. Kanakoff1 and James H. McLean2
Abstract: The genus Neadmete Habe, 1961, is suggested
for the cool-water cancellariid species previously treated in the
genus Massy la H. and A. Adams, 1854, and a new species, Nead-
mete sutherlandi, is described from the Late Pliocene Lomita
Marl formation of San Pedro, California.
In February of 1965, Mr. Roger Reimer brought to the Los Angeles
County Museum of Natural History samples of fossiliferous marl and some
shells and otolith material from the San Pedro area. Excavations by Mr. John
E. Fitch, Director of the California Fish and Game Laboratories, followed and
yielded an unusually large number of minute otoliths and molluscan species
resembling those of the hitherto single rich source of megafossils at the forma-
tion from the locality known and recorded in the literature as “Hilltop Quarry,”
a source that has been unavailable since 1940. Extensive excavations of these
outcrops by Mr. Fitch and the Museum staff yielded a large and unique fauna
with new fossil records of mollusks. The formation suggests Late Pliocene,
rather than Early Pleistocene age, as has previously been accepted. Further
analysis of molluscan and foraminiferal material will be required to document
this conclusion. The description of a striking new species of cancellariid is
presented at this time.
Acknowledgments
We are indebted to Dr. A. M. Keen of Stanford University for helpful
suggestions and critically reviewing the manuscript. Photographs are by Mr.
Armando Solis, Museum photographer.
Genus Neadmete Habe, 1961
Neadmete Habe, 1961a, App., p. 28. Type species, by original desig-
nation, Neadmete japonica (Smith, 1879). Recent, Japan. Habe, 1961a, p. 73,
pi. 36, fig. 2; 1961b, p. 435, pi. 24, fig. 3; 1964, p. 113, fig. 2.
1Research Associate, Los Angeles County Museum of Natural History.
2Curator of Invertebrate Zoology, Los Angeles County Museum of Natural History.
1
2
Contributions in Science
No. 116
Figure 1. Neadmete sutherlandi, new species. Holotype, LACM 1150. Long. 39.6,
lat. 16.1 mm.
1966
New Fossil Mollusk
3
Figure 2. Neadmete sutherlandi, new species. Holotype, LACM 1150. Long. 39.6,
lat. 16.1 mm.
4
Contributions in Science
No. 116
A translation from Japanese of the original generic diagnosis of Habe
follows:
Shell small, whitish; surface covered with periostracum; spire high,
length of body whorl more than half of shell length; spiral sculpture fine and
distinct; axial sculpture forming cancellations, which sometimes become
obsolete on body whorl.
Through a misinterpretation of the type species of Massyla H. and A.
Adams, 1854, Massyla has erroneously been applied to related west American
species, dating from its first application by Strong in Burch (1945: 10). The
type species, Cancellaria corrugata Hinds, 1843, has been little known until
syntype material was recently figured by Keen (1966: 27, pi. 46, fig. 7). It is
clear that this relatively smooth species from the Panamic province having
strong columellar folds has nothing in common with the cool-water group
under consideration.
Dali (1921) applied the generic name Sveltia Jousseaume, 1888, type
species, Voluta varicosa Brocchi, 1814, Pliocene, Italy. The type species, as
figured by Jousseaume (1888: 24, fig. 3) has prominent raised axial ribbing,
with spiral incisions, two heavy columellar plaits, and strong spiral ribbing on
the inner surface of the outer lip. It clearly represents a tropical group not
related to the boreal forms under consideration.
Neadmete Habe thus appears to be the genus most applicable to the
relatively large cool-water species in the North Pacific differing from Admete
Moller, 1842, ex Kroyer MS, type species, Admete crispa Moller, 1842 (— A.
couthouyi Jay, 1839) in having a higher spire relative to the length of the body
whorl, lacking the tendency toward development of a broad columellar shield
of Admete, having a straighter canal than Admete, with continuous spiral
sculpture in the columellar area, that increases the number of columellar plaits.
Only two or possibly three Recent species of Neadmete are known from
the northeastern Pacific: N. modesta (Carpenter, 1864) (Palmer, 1958: 224,
pi. 24, figs. 17-18, holotype) and TV. circumcincta (Dali, 1873) (p. 59, pi. 2,
fig. 2). Neadmete unalashkensis (Dali, 1873) (p. 58, pi. 2, fig. 1) appears to
be a sculptural variant of N. modesta having more prominent axial ribs. Both
N. modesta and TV. circumcincta are highly variable in shell proportions,
heaviness of shell, and strength of sculpture.
Neadmete Sutherland!, new species
Figures 1 and 2
Description: Shell relatively large, slender, whorls 10, spire elevated,
turreted. Nucleus simple, whorls two. Approximately 16 axial ribs on early
whorls, becoming obsolete on body whorl. Spiral sculpture of prominent,
widely spaced ribs, superimposed on the axial ribs; 3 on each postnuclear whorl
and 8 on body whorl, with single intercalary ribs between major ribs. Whorls
1966
New Fossil Mollusk
5
subcylindrical, tabulate; sutures deep; outer lip thick. Columellar area with
two main plaits and 5 plicae formed by extension of primary spiral ribbing.
Aperture oval; anterior canal short and deep. Dimensions (in mm.): Long.
39.6, lat. 16.1, long, of aperture 15.4 (holotype).
Type Locality: Los Angeles County Museum of Natural History, Inverte-
brate Paleontology locality no. 435: a fossiliferous outcrop on the east side of
the canyon below the convergence of Park Western Drive, West Coralmount
Drive, and Host Place, San Pedro, California. This locality probably cor-
responds to USGS locality no. 12222 (Woodring, Bramlette and Kew, 1946).
Age and Formation: Late Pliocene (previously reported in the literature
as Early Pleistocene), Lomita marl.
Type Material: Holotype, Los Angeles County Museum of Natural
History, Invertebrate Paleontology, cat. no. 1150, 3 paratypes, cat. no. 1151.
Two of the paratypes are smaller specimens, having 8 to 9 whorls, and measure
(in mm.), long. 19.0, lat. 18.6; long. 17.9, lat. 7.8 mm. One specimen is lacking
the spire but has an aperture 20 mm. long, which enables an estimate for the
total length of the shell to be approximately 50 mm., larger than the holotype.
Discussion: Neadmete sutherlandi attains nearly twice the size of other
known species of Neadmete . It has sculptural features in common with both
N. modesta and N. circumcincta. The whorls are not as tabulate as those of N.
modesta, which generally has 5 major ribs exposed on the early whorls. The
tabulation of N. sutherlandi is considerably more pronounced than that of
N. circumcincta which has a larger number of spiral ribs of uneven strength.
Neadmete sutherlandi resembles N. japonica in details of sculpture, but com-
parison with the illustrations of the latter given byHabe (1961a, 1961b, 1964),
shows it is a much larger and more slender shell than that of N. japonica.
The species is named for Mr. J. Alden Sutherland, Museum Field Assistant
in Paleontology, who collected the type material for the Museum.
Literature Cited
Dali, W. H.
1873. Preliminary descriptions of new species of Mollusca from the coast
of Alaska, with notes on some rare forms. Proc. Calif. Acad. Sci.,
5: 57-62, pi. 2.
1921. Summary of the marine shell-bearing mollusks of the northwest coast
of America, from San Diego, California, to the Polar Sea, mostly
contained in the collection of the United States National Museum,
with illustrations of hitherto unfigured species. Bull. U.S. Natl. Mus.,
112: 1-127, 22 pis.
Habe, Tadashige
1961a. Coloured illustrations of the shells of Japan (II). Osaka, Japan:
Hoikusha, xii -f 183 pp., 66 pis. [in Japanese].
1961b. Description of four new cancellariid species, with a list of the Japa-
nese species of the family Cancellariidae. Venus: Jap. J. Malacology,
21(4): 431-441, pis. 23-24.
6
Contributions in Science
No. 116
1964. Shells of the Western Pacific in color, vol. II. Osaka, Japan: Hoikusha,
vii + 233 pp., 66 pis. [English ed. of Habe, 1961a.]
Jousseaume, M. le D’
1888. La famille des Cancellariidae. Le Naturaliste, Revue Illustree des
Sciences Naturelles. Ann. 9, Ser. 2, pp. 1-31, illust.
Keen, A. M.
1966. West American mollusk types in the British Museum (Natural His-
tory), II. Species described by R. B. Hinds. Veliger, 8(4) : 265-275, pis.
46-47, 6 text figs.
Palmer, K. V. W.
1958. Type specimens of marine Mollusca described by P. P. Carpenter from
the west coast (San Diego to British Columbia), vi + 376 pp., 35 pis.
Strong, A. M.
1945. Family Cancellariidae. In J. Q. Burch, Minutes of the Conchological
Club of So. Calif., no. 49, pp. 2-14.
Woodring, W. P., M. N. Bramlette, and W. S. W. Kew
1946. Geology and paleontology of Palos Verdes Hills, California. U.S.
Geol. Survey Prof. Paper 207, v + 145 pp., 37 pis.
LOS
ANGELES
1;
COUNTY
MUSEUM
CONTRIBUTIONS
IN SCIENCE
IJMBER 117
December 28, 1966
A NEW SPECIES OF ARCHITECTONICA FROM THE SANTA
SUSANA MOUNTAINS, VENTURA COUNTY, CALIFORNIA
By J. Alden Sutherland
ii
j
i
Los Angeles County Museum of Natural History • Exposition Park
Los Angeles, California 90007
CONTRIBUTIONS IN SCIENCE is a series of miscellaneous technical papers
in the fields of Biology, Geology and Anthropology, published at irregular intervals
by the Los Angeles County Museum of Natural History. Issues are numbered sepa-
rately, and numbers run consecutively regardless of subject matter. Number 1 was
issued January 23, 1957. The series is available to scientific institutions on an ex-
change basis. Copies may also be purchased at a nominal price.
INSTRUCTIONS FOR AUTHORS
Manuscripts for the LOS ANGELES COUNTY MUSEUM CONTRIBU-
TIONS IN SCIENCE may be in any field of Life or Earth Sciences. Acceptance of
papers will be determined by the amount and character of new information and the
form in which it is presented. Priority will be given to manuscripts by staff members,
or to papers dealing largely with specimens in the Museum’s collections. Manuscripts
must conform to CONTRIBUTIONS style and will be examined for suitability by
an Editorial Committee. They may also be subject to critical review by competent
specialists.
MANUSCRIPT FORM. — ( 1 ) The 1960 AIBS Style Manual for Biological
Journals is highly recommended as a guide. (2) Typewrite material, using double
spacing throughout and leaving ample margins, on only one side of 8V2 x 1 1 inch
standard weight paper. (3) Place tables on separate pages. (4) Footnotes should be
avoided if possible. (5) Legends for figures and unavoidable footnotes should be
typed on separate sheets. Several of one kind may be placed on a sheet. (6) Method
of literature citation must conform to CONTRIBUTIONS style— see number 90 and
later issues. Spell out in full the title of non-English serials and places of publication.
(7) A factual summary is recommended for longer papers. (8) A brief abstract must
be included for all papers. This will be published at the head of each paper.
ILLUSTRATIONS/ — All illustrations, including maps and photographs, should
be referred to as “figures!’ All illustrations should be of sufficient clarity and in the
proper proportions for reduction to CONTRIBUTIONS page size. Permanent ink
should be used in making line drawings and in lettering (do not type on drawings);
photographs should be glossy prints of good contrast. Original illustrations will not
be returned unless specifically requested when the manuscript is first submitted.
PROOF. — Author will be sent galley proof which should be corrected and re-
turned promptly. Changes after the paper is in galley will be billed to the author. Un-
less specially requested, page proof will not be sent to the author. 100 copies of each
paper will be given free to a single author or divided equally among multiple authors.
Orders for additional copies should be sent to the Editor at the time corrected galley
proof is returned; appropriate forms for this will be included when galley is sent.
David K. Caldwell
Editor
A NEW SPECIES OF ARCHITECTONICA FROM THE SANTA
SUSANA MOUNTAINS, VENTURA COUNTY, CALIFORNIA
By J. Alden Sutherland1
Abstract: Architectonica llajasensis; a new species of gas-
tropod from the Llajas formation, Middle Eocene, Santa Susana
Mtns., Ventura Co., California.
A new species of Architectonica has been recognized during the study of
material from Las Llajas Canyon, Santa Susana Mountains, Ventura County,
California. Two locations have been of primary interest: LACMIP 461 -A, in
the lowest exposed layer of Las Llajas Canyon, and LACMIP 461-B, on the
northern slope of a small canyon intersecting Las Llajas Canyon from the
east. LACMIP 461-B is stratigraphically 70 feet higher than 461 -A, and is
200 feet from the top of the Llajas formation, (see McMasters, 1933; Stipp
& Tolman, 1934) The new species of Architectonica is from LACMIP 461-B.
The faunal differences of the two locations are quite distinct and constant.
LACMIP 461 -A is characterized by Cylichnina tantilla Anderson & Hanna,
and Lyria andersoni Waring; while 461-B is characterized by V enericardia
hornii (Gabb) forma calafia Stewart, and Eucrassatella semidentata (Cooper) .
Other species common to both locations are Cernina hannibali (Dickerson),
and Amaurellina clarki Stewart.
Architectonici llajasensis, new species
Figures 1 and 2
Diagnosis: Architectonica llajasensis differs from any other described
species in that it has eight tuberculate spiral ribs with one intercalary thread
on the body whorl, in the extreme acute peripheral carina, and also in the order
of sculpture of the undersurface, and umbilical whorls.
Description of holotype: Shell medium size for genus; carinate; low conic;
walls thin; suture impressed; aperture oblique, quadrate; umbilicus broad;
whorls six. First three whorls not sculptured, fourth whorl sculptured by six
tuberculate spiral ribs, penultimate whorl sculptured by eight tuberculate
spiral ribs, body whorl sculptured by eight tuberculate spiral ribs, with one
tuberculate intercalary thread. Growth lines appear on body whorl behind
aperture; whorls convex below suture, concave above; peripheral carina acute,
tuberculate; undersurface concave at peripheral margin, convex from middle
to basal keel; undersurface sculptured by nine spiral ribs, one or two inter-
calaries, two ribs nearest basal keel tuberculate; umbilical whorls sculptured
by five tuberculate spiral ribs, with one tuberculate intercalary thread.
Diameter 26.5, altitude 14, aperture height 7, width 7 mm.
1Field Assistant in Invertebrate Paleontology, Los Angeles County Museum of
Natural History.
1
2
Contributions in Science
No. 117
Type locality: LACMIP 461-B, Middle Eocene, Llajas formation, Las
Llajas Canyon, Santa Susana Mountains, Ventura County, California.
Holotype: LACMIP 1 140. Collected by author, Lebruary 5, 1966.
Architectonica llajasensis is named for the geologic formation at the type
locality. A single specimen is known.
Figure 1. Architectonica llajasensis, new species. Holotype, LACMIP 1140.
Diameter 26.5 mm.-.
1966
New Fossil Gastropod Mollusk
3
Discussion: This species occurs in the same sediments with A. cognata
Gabb, which it resembles only in general configuration. It differs chiefly in
spiral sculpture. It resembles A. elaboratum Conrad (see Conrad, 1833), of
the Claiborne, Alabama, Eocene, to a greater degree than any western Eocene
species. Both species are low conic, and sculptured by tuberculate spiral ribs.
A. elaboratum differs in being uniformly convex; the peripheral carina is not
as acute, and the sculpture of the basal keel is more ornate.
Acknowledgments
I wish to thank Mr. James Runkle of Simi, California, for his kind per-
mission to collect paleontological material from his property. I also wish to
Figure 2. Architectonica llajasensis, new species. Holotype, LACMIP 1140.
Diameter 26.5 mm.
4
Contributions in Science
No. 117
thank George P. Kanakoff, formerly Curator of Invertebrate Paleontology of
the Los Angeles County Museum of Natural History, for his encouragement
and guidance in preparing this description.
Literature Cited
Conrad, T. A.
1833. Fossil shells of the Tertiary formations of North America. Phila-
delphia Acad. Bull, 1(3): 29-38.
McMasters, J. H.
1933. Eocene Llajas Formation, Ventura County, California. Bull. Geological
Soc. Amer, 44(1): 217-218.
Stipp, T. F, Tolman, F. D.
1934. Eocene stratigraphy of the north side of Simi Valley. Pan-American
Geol, 62(1): 79.
LOS
ANGELES
CONTRIBUTIONS
COUNTY
MUSEUM
IN SCIENCE
Timber 118
December 28, 1966
1
l
j
A NEW TOE BITER FROM MEXICO
(BELOSTOMATIDAE, HEMIPTERA)
By A. S. Menke
Los Angeles County Museum of Natural History
Los Angeles, California 90007
Exposition Park
CONTRIBUTIONS IN SCIENCE is a series of miscellaneous technical papers
in the fields of Biology, Geology and Anthropology, published at irregular intervals
by the Los Angeles County Museum of Natural History. Issues are numbered sepa-
rately, and numbers run consecutively regardless of subject matter. Number 1 was
issued January 23, 1957. The series is available to scientific institutions on an ex-
change basis. Copies may also be purchased at a nominal price.
INSTRUCTIONS FOR AUTHORS
Manuscripts for the LOS ANGELES COUNTY MUSEUM CONTRIBU-
TIONS IN SCIENCE may be in any field of Life or Earth Sciences. Acceptance of
papers will be determined by the amount and character of new information and the
form in which it is presented. Priority will be given to manuscripts by staff members,
or to papers dealing largely with specimens in the Museum’s collections. Manuscripts
must conform to CONTRIBUTIONS style and will be examined for suitability by
an Editorial Committee. They may also be subject to critical review by competent
specialists.
MANUSCRIPT FORM.— (1) The 1960 AIBS Style Manual for Biological
Journals is highly recommended as a guide. (2) Typewrite material, using double
spacing throughout and leaving ample margins, on only one side of 8V2 x 11 inch
standard weight paper. (3) Place tables on separate pages. (4) Footnotes should be
avoided if possible. (5) Legends for figures and unavoidable footnotes should be
typed on separate sheets. Several of one kind may be placed on a sheet. (6) Method
of literature citation must conform to CONTRIBUTIONS style— see number 90 and
later issues. Spell out in full the title of non-English serials and places of publication.
(7) A factual summary is recommended for longer papers. (8) A brief abstract must
be included for all papers. This will be published at the head of each paper.
ILLUSTRATIONS. — All illustrations, including maps and photographs, should
be referred to as “figures!’ All illustrations should be of sufficient clarity and in the
proper proportions for reduction to CONTRIBUTIONS page size. Permanent ink
should be used in making line drawings and in lettering (do not type on drawings);
photographs should be glossy prints of good contrast. Original illustrations will not
be returned unless specifically requested when the manuscript is first submitted.
PROOF. — Author will be sent galley proof which should be corrected and re-
turned promptly. Changes after the paper is in galley will be billed to the author. Un-
less specially requested, page proof will not be sent to the author. 100 copies of each
paper will be given free to a single author or divided equally among multiple authors.
Orders for additional copies should be sent to the Editor at the time corrected galley
proof is returned; appropriate forms for this will be included when galley is sent.
David K. Caldwell
Editor
A NEW TOE BITER FROM MEXICO
(BELOSTOMATIDAE, HEMIPTERA)
By A. S. Menke1
Abstract: A new species of Abedus, subgenus Abedus,
from the Mexican states of Sonora and Jalisco is described and
named A. parkeri. The relationships with A. breviceps and A.
signoreti sonorensis are discussed.
Recent collecting trips in northwestern Mexico have produced a new
species of the genus Abedus, subgenus Abedus. The material studied is in the
Los Angeles County Museum of Natural History (LACM) which includes the
A. S. Menke Collection (ASM). Terminology used follows Menke (1960).
Abedus parkeri Menke, new species
Figures 2a, b; 3a, b
2a. parkeri 2 b.
Figure 1. Phallus of Abedus breviceps Stal, lateral view.
Figure 2. Phallus of Abedus parkeri Menke, new species. A, Lateral view; B, Apical
view.
department of Entomology, University of California, Davis.
1
2
Contributions in Science
No. 118
Holotype 8 : 25.5 mm. long, 14 mm. wide.
Head: Eye slightly flattened externally, slightly higher than interocular
space; eye width about one-half interocular distance; length of tylus before eye
slightly more than one half eye length; antenna with four segments, II and III
with long finger-like processes.
Hemelytron: Embolial fracture present; membrane broad with seven
elongate cells, (fig. 3a), width (including translucent margin) slightly greater
(one third greater in females) than median length of posterior lobe of pro-
notum; apex of hemelytron broadly rounded.
Abdominal venter: Hair pattern as in Abedus breviceps (see Menke,
1960, fig. 39).
Air strap : As in breviceps (see Menke, 1960, fig. 11) except somewhat
narrower.
Phallus: Opening of ejaculatory duct heart shaped (Fig. 2b); roof over
duct, a broadly rounded triangular ridge; lateral wings strong posteriorly
(Fig. 2a) , disappearing before apex and not visible in apical view (Fig. 2b) .
Types: Holotype 8, Alamos, Sonora, Mexico, IX-1-1960, R. L. West-
cott (LACM). Nine paratypes from the following Mexican localities:
JALISCO: 3 mi. S.E. Plan de Barrancas, VII-8-1963, 1 8, F. D. Parker and
L.A. Stange (ASM). SONORA: Alamos, IX-1-1960, 1 8,1 $ , R. L. Westcott,
(ASM, LACM); Alamos, VII-16-17-1963, 1 9, R. L. Westcott (LACM); La
Aduana, VI- 12- 1961, 3 8,2 9 , A. S. Menke, F. D. Parker, L. A. Stange
(ASM).
Discussion: Abedus parked is most similar to A. breviceps Stal but the
broad hemelytral membrane with its long cells, and the smaller eyes are diag-
nostic (compare Figs. 3a and 4a). Abedus signoreti sonorensis Menke is also
similar to parked but sonorensis does not have ventral laterotergite I com-
pletely covered by appressed pubescence. Photographs of all three species are
given for comparison (Figs. 3 to 5).
The phallus of A. parked is quite different from that of breviceps but is
similar to that of signoreti. It differs from the phallus of A. breviceps in the
short, basal lateral wings which are not evident in apical view (Figs. 2a, b).
The phallus of breviceps is shown in lateral view for comparison (Fig. 1 ) .
The Alamos and Plan de Barrancas specimens were collected at lights
according to their collectors. This is the second species of the genus in which
this behavior has been noted (for notes on the first, A. signoreti, see Menke
and Truxal, 1966) .
The La Aduana specimens were collected in small muddy pools of a dried
up stream bed. Abedus signoreti sonorensis were found with parked.
This species is named in honor of Frank D. Parker, one of its collectors.
Based on my 1960 key to the species of Abedus, A. parked will run to
couplet 4. The following changes in the key should be made (figures refer to
those found in the 1960 revision) :
1966
New Mexican Toe Biter
3
Figure 3. Abedus parked Menke, new species, holotype. A, Dorsal view; B, Ventral
view.
4
Contributions in Science
No. 118
Figure 4. Abedus breviceps Stal. A, Dorsal view; B, Ventral view.
1966
New Mexican Toe Biter
5
Figure 5. Abedus signor eti sonorensis Menke. A, Dorsal view; B, Ventral view.
6
Contributions in Science
No. 118
4. Air strap with a subapical dorsal sac (fig. 10, a, b) ; mesal glabrous area of
abdominal plates narrowed towards base of abdomen, usually extending
from base of plate IV to apex of plate V (occasional specimens have a
mesal glabrous area on plate III) (fig. 38) Abedus ovatus Stal
Air strap without a dorsal sac (fig. 11, a, b); mesal glabrous area of ab-
dominal plates broad, usually extending from base of plate III to apex
of V (occasional specimens have a mesal bare area on plate II
(fig. 39) 4a
4a. Hemelytral membrane narrow, cells narrow or poorly defined, largest
wider than high, or at most square (some females), combined width of
membrane and translucent margin about equal to median length of poste-
rior lobe of pronotum2; eye much higher than interocular space (fig. 33)
Abedus breviceps Stal
Hemelytral membrane broad, largest cells longer than wide, combined
width of membrane and translucent margin slightly greater than (males)
to one third greater than (females) median length of posterior lobe of pro-
notum; eye only slightly higher than interocular space (similar to fig. 29)
Abedus parkeri Menke
Literature Cited
Menke, A. S.
1960. A taxonomic study of the genus Abedus Stal. Univ. Calif. Publ. Ento-
mol., 16(8): 393-440.
Menke, A. S. and F. S. Truxal
1966. New distribution data for Martarega, Buenoa, and Abedus, including
the first record of the genus Martarega in the United States. Los An-
geles Co. Mus., Contr. in Sci., 106: 1-6.
2In some female breviceps the combined width of membrane and translucent
margin may be greater than the width of the posterior pronotal lobe, but in these
specimens the translucent margin is broader than the row of cells in the membrane.
In parkeri females the translucent margin is narrower than the row of cells in the
membrane.
angeles CONTRIBUTIONS
ZTZ IN SCIENCE
juMBER 119 December 31, 1966
lb c / ^ / <
\l 0 / Q/q
\sU ^ 0 b o -
ADDITIONAL FISH REMAINS, MOSTLY OTOLITHS, FROM A
PLEISTOCENE DEPOSIT AT PLAYA DEL REY, CALIFORNIA
I
By John E. Fitch
.
Ii
:[
ij
jl
Los Angeles County Museum of Natural History • Exposition Park
Los Angeles, California 90007
CONTRIBUTIONS IN SCIENCE is a series of miscellaneous technical papers
in the fields of Biology, Geology and Anthropology, published at irregular intervals
by the Los Angeles County Museum of Natural History. Issues are numbered sepa-
rately, and numbers run consecutively regardless of subject matter. Number 1 was
issued January 23, 1957. The series is available to scientific institutions on an ex-
change basis. Copies may also be purchased at a nominal price.
INSTRUCTIONS FOR AUTHORS
Manuscripts for the LOS ANGELES COUNTY MUSEUM CONTRIBU-
TIONS IN SCIENCE may be in any field of Life or Earth Sciences. Acceptance of
papers will be determined by the amount and character of new information and the
form in which it is presented. Priority will be given to manuscripts by staff members,
or to papers dealing largely with specimens in the Museum’s collections. Manuscripts
must conform to CONTRIBUTIONS style and will be examined for suitability by
an Editorial Committee. They may also be subject to critical review by competent
specialists.
MANUSCRIPT FORM.— (1) The 1960 AIBS Style Manual for Biological
Journals is highly recommended as a guide. (2) Typewrite material, using double
spacing throughout and leaving ample margins, on only one side of 8V2 x 11 inch
standard weight paper. (3) Place tables on separate pages. (4) Footnotes should be
avoided if possible. (5) Legends for figures and unavoidable footnotes should be
typed on separate sheets. Several of one kind may be placed on a sheet. (6) Method
of literature citation must conform to CONTRIBUTIONS style — see number 90 and
later issues. Spell out in full the title of non-English serials and places of publication.
(7) A factual summary is recommended for longer papers. (8) A brief abstract must
be included for all papers. This will be published at the head of each paper.
ILLUSTRATIONS. — All illustrations, including maps and photographs, should
be referred to as “figures!’ All illustrations should be of sufficient clarity and in the
proper proportions for reduction to CONTRIBUTIONS page size. Permanent ink
should be used in making line drawings and in lettering (do not type on drawings);
photographs should be glossy prints of good contrast. Original illustrations will not
be returned unless specifically requested when the manuscript is first submitted.
PROOF. — Author will be sent galley proof which should be corrected and re-
turned promptly. Changes after the paper is in galley will be billed to the author. Un-
less specially requested, page proof will not be sent to the author. 100 copies of each
paper will be given free to a single author or divided equally among multiple authors.
Orders for additional copies should be sent to the Editor at the time corrected galley
proof is returned; appropriate forms for this will be included when galley is sent.
David K. Caldwell
Editor
ADDITIONAL FISH REMAINS, MOSTLY OTOLITHS, FROM A
PLEISTOCENE DEPOSIT AT PLAYA DEL REY, CALIFORNIA
By John E. Fitch1
Abstract: When the washed screenings from a 200-pound
field sample of fossiliferous “dirt” from the Playa del Rey
locality (LACMIP 59) were examined a spoonful at a time
with the aid of a microscope, 1,174 identifiable otoliths (rep-
resenting 34 “species”) were recovered. Twelve of these species
were new to the deposit, as were two of the elasmobranchs from
which teeth were found. The yield of fish remains from this “re-
sampling experiment” emphasizes the necessity for using a micro-
scope or similar viewing aid for obtaining accurate information
on the fauna contained in a fossil deposit.
Fish otoliths, teeth, and other remains routinely are picked from washed
fossiliferous screenings by numerous volunteers (unpaid) and a few salaried
employees at the Los Angeles County Museum of Natural History. Most of
the sorting has been done by volunteer personnel working in the Invertebrate
Paleontology section of the Museum, and their primary concern has been
fossil mollusks. Fortunately, most of these student workers were trained by
George P. Kanakoff, formerly Curator of Invertebrate Paleontology, who
subsequently supervised their activities. Thus, recognizable vertebrate remains,
as well as mollusks and other invertebrates, have been saved whenever
encountered.
The search-and-removal technique employed at the Museum (Inverte-
brate Paleontology) involves spreading a thin layer of washed screenings in
front of the searcher, and picking from this residue all identifiable vertebrate
and invertebrate remains that are observed. Most of the time this work is
accomplished without the help of any device that will aid in detection ( e.g .,
magnifying glass, hand lens, microscope, etc.).
Over the years I have found that when I am searching through fossil-
iferous material for otoliths, I often fail to “see” the numerous mollusks and
other invertebrate remains that pass before my vision. Conversely, when I
have been looking for and removing mollusks, I often passed up otoliths that
I did not see. Because of this, and because many fish otoliths are minute, or
odd-shaped, or both, I decided to test the efficiency of the Museum’s “perusal-
by-eye” technique by examining under a microscope some of the fine material
they were discarding after having finished sorting through it.
For finding fossil otoliths, I spread a tablespoonful of washed fossil-
iferous screenings evenly in a flat plastic dish with slightly raised margins, and
systematically search through this material with a pair of forceps while using
1Research Associate, Los Angeles County Museum of Natural History; Marine
Biologist, California Department of Fish and Game, Terminal Island, California.
1
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Contributions in Science
No. 119
a binocular microscope at six magnifications. Almost every spoonful of the
Museum’s discard yielded one or more otoliths when this technique was used.
Since it appeared that many otoliths were escaping detection, I decided
to resample a number of the Museum’s fossil localities while they were still
accessible. (Each month in southern California fossil deposits of long standing
are lost because of freeway construction, housing developments, cut-and-fill
projects, and a multitude of other “progressive” activities of modern man.)
Since the fish fauna of the Playa del Rey (Lincoln Avenue) locality
already had been reported upon (Fitch, 1964), it seemed an appropriate place
to start resampling. The 1,376 identified otoliths from this site (Table 1)
had been gleaned by Museum personnel from several tons (at least) of fossil-
Table 1. Percentages of otoliths sorted from Playa del Rey screenings by eye
(Fitch, 1964) and with the aid of a microscope
(Relative size of otolith: large, medium, small, indicated to left of name)*
How sorted How sorted
Species
By With
eye scope
Species
By With
eye scope
s. Anchoa compressa
0.3
1. Micropogon ectenes
0.1
1. Anisotremus davidsoni
0.1
s. Occa verrucosa
0.1
s. Argentina sialis
0.1
1. Otophidium spp.
16.3
9.7
s. Atherinops affinis
0.1
0.9
m. Paralabrax sp.
0.3
m. Atherinopsis calif orniensis 0.2
0.1
m. Paralichthys calif ornicus
0.1
0.4
s. Chitonotus pugetensis
0.1
0.6
m. Parophrys vetulus
0.1
0.3
m. Citharichthys spp.
21.2
19.0
m. Phanerodon furcatus
0.6
1. Coelorhynchus scaphopsis
0.1
s. Pleuronichthys ritteri
0.1
m. Cymatogaster aggregata
0.4
0.8
1. Porichthys spp.
12.8
24.0
1. Cynoscion nobilis
0.4
m. Prionotus stephanophry s
0.2
1. Cynoscion reticulatus
0.3
0.1
1. Roncador stearnsi
0.7
s. Engraulis mordax
4.9
16.5
1. Sebastodes spp.
1.8
0.7
1. Genyonemus lineatus
15.8
9.9
1. Seriphus politus
20.1
9.9
s. Icelinus quadriseriatus
0.8
1. Sphyraena argentea
0.1
s. Lepidogobius lepidus
0.1
0.7
s. Stenobrachius leucopsarus
0.1
1. Lepophidium negropinna
0.1
s. Symbolophorus californiense
0.1
s. Leuresthes tenuis
0.4
0.3
s. Symphurus atricauda
1.1
m. Lyopsetta jordani
0.1
m. Trachurus symmetricus
0.3
m. Menticirrhus undulatus
0.1
0.4
1. Umbrina roncador
1.0
0.3
1. Merluccius productus
1.8
0.9
s. Zaniolepis latipinnis
0.1
Total otoliths 1,376 1,174
Percent 99.9 99.1
*large = > Vz inch; medium = 14 to Vz inch; small = <14 inch
1966
Pleistocene Fish Remains
3
iferous material gathered during a period of three decades or more. In fact,
this lens-shaped deposit has been so thoroughly sampled in the past that it took
some rather diligent searching to obtain a 200-pound field sample.
I routinely screen field samples through three sieves (2, 1, and 0.5 mm,
U.S. Standard Sieve Series) that fit one into the other “piggy-back” style.
After soaking my field sample in a tub of water, I place several handfuls of the
saturated dirt into the top (largest mesh) sieve, submerge all three in a second
tub of water to within one-half inch of the top of the upper sieve, and filter
the mixture by gently rotating and shaking the submerged screens. When the
residue in each screen is clean (a running hose played over the top of each one
as it is removed will guarantee best results) I “dump” the contents onto
several thicknesses of newspaper and allow it to dry in the sun.
When the sample is dry, I screen the coarsest material through Va -inch
mesh to remove “large” shells, rocks, bone fragments, and similar items. The
residue retained by the Va -inch mesh can be checked by eye for the rare shark
tooth or very large otolith it might contain. All remaining material is exam-
ined through a binocular microscope at six magnifications. By having washed
the samples through three screens, the particles are graded by size and the
task of examining the material under the microscope is greatly simplified.
Using these techniques of washing, screening, and sorting, I gleaned
1,174 identifiable otoliths from the 200-pound Playa del Rey field sample
(Table 1). Interestingly, while 19 of the 40 “species” were encountered in both
samplings, 9 kinds were found only in the earlier diggings, and 12 only in the
resampling. The same six “species” ( Citharichthys spp., Engraulis mordax,
Genyonemus lineatus, Otophidium spp., Porichthys spp., and Seriphus politus )
were important in both samplings, but the proportions generally were markedly
different. Genyonemus lineatus, Otophidium spp., and Seriphus politus com-
prised 15.8, 16.3, and 20.1 percent of the 1,376 otoliths sorted by eye, but
only 9.9, 9.7, and 9.9 percent of the 1,174 obtained with the aid of the micro-
scope. Obviously, because of their large maximum sizes, otoliths of these
three species were “found” with fair frequency by eye. On the other hand, the
small otoliths of Engraulis mordax comprised only 4.9 percent of the earlier
sample, but made up 16.5 percent of the yield from resampling— their small
maximum size unquestionably contributed to the low numbers found with the
naked eye. The 24.0 percent recovery rate for Porichthys spp. otoliths with
the microscope, compared to 12.8 percent by eye, was a result of two factors:
failure of earlier sorters to recognize them because of their odd shape, and
the difficulty of discerning with the naked eye the great numbers of small
midshipman otoliths present in this deposit (many of the Porichthys otoliths
recovered by use of the microscope were 2 mm or less in greatest dimension).
Otoliths of six of the nine species found by Museum personnel, but not
encountered in my resampling, were large, two were medium sized, and one
was small. For only three of these nine species, were more than two otoliths
found. Thus, it may be assumed that the otoliths of most of these species,
Table 2. Fish remains found during resampling
of the Playa del Rey Pleistocene
Type and number of remains
Scientific name Common name otoliths teeth vertebrae other
ELASMOBRANCHS
Alopias vulpinus
thresher
1
Carcharhinus sp.
requiem shark
7
Dasyatis dipterurus
diamond stingray
2
Galeorhinus zyopterus
soupfin shark
1
Heterodontus francisci
horn shark
1
Isurus oxyrinchus
mako
2
Myliobatis calif ornicus
bat stingray
32
Raja spp.
skates
4*
Squatina calif ornica
Pacific angel shark
6
Sphyrna sp.
hammerhead shark
6
Urolophus halleri
round stingray
5**
unidentified elasmobranchs
15
TELEOSTS
Anchoa compressa
deepbody anchovy
4
Argentina sialis
Pacific argentine
1
Atherinopsis calif orniensis
jacksmelt
1
atherinids
atherinids
10
Chitonotus pugetensis
roughback sculpin
7
Citharichthys sordidus
Pacific sanddab
7
Citharichthys stigmaeus
speckled sanddab
181
Citharichthys spp.
sanddabs
47
Coelorhynchus scaphopsis
Gulf rattail
1
Cymatogaster aggregata
shiner perch
10
Cynoscion reticulatus
striped corvina
1
Engraulis mordax
northern anchovy
194
Genyonemus lineatus
white croaker
116
Icelinus quadriseriatus
yellowchin sculpin
10
Lepidogobius lepidus
bay goby
8
Leuresthes tenuis
grunion
4
Menticirrhus undulatus
California corbina
5
Merluccius productus
Pacific hake
11
Occa verrucosa
warty poacher
1
Otophidium scrippsae
basketweave cusk-eel
49
Otophidium taylori
spotted cusk-eel
65
Paralabrax sp.
bass
3
Paralichthys calif ornicus
California halibut
5
Parophrys vetulus
English sole
3
Porichthys myriaster
specklefin midshipman
3
Porichthys no tat us
plainfin midshipman
278
Prionotus stephanophrys
lumptail searobin
2
Sebastodes spp.
rockfish
8
Seriphus politus
queenfish
116
Stenobrachius leucopsarus
northern lampfish
1
Symbolophorus calif orniense
California lanternfish
1
Symphurus atricauda
California tonguefish
13
Trachurus symmetricus
Pacific jackmackerel
3
Umbrina roncador
yellowfin croaker
4
Zaniolepis latipinnis
longspine combfish
1
unidentified teleosts
8
18
32 3t
1,182
♦skate “wing” spines
♦♦caudal “stings”
t2 fin spines, 1 jaw fragment
1966
Pleistocene Fish Remains
5
although rare in the deposit, were recovered efficiently because of their large
size.
On the other hand, otoliths of 8 of the 12 species found in the resampling,
but not found previously, were small, three were medium sized, and only one
was large. Six of the 12 ( Argentina sialis, Coelorhynchus scaphopsis, Occa
verrucosa, Stenobrachius leucopsarus, Symbolophorus calif orniense, and
Zaniolepis latipinnis ) were represented by only one otolith each, but one
species ( Icelinus quadriseriatus ) yielded 10 (Table 2). Thus, although sorting
by eye seems to be productive of large otoliths, it is not an efficient method
for recovering the small ones.
Elasmobranch remains in the resampling consisted primarily of teeth,
but some vertebrae, caudal stings, and dermal denticles also were found
(Table 2). Only two of the 11 species ( Dasyatis dipterurus and Heterodontus
francisci ) were not previously reported for this site (Fitch, 1964). The teeth
of both of these are relatively small compared with the other shark and ray
teeth found in this deposit, and the flattened laterals of Heterodontus could be
easily overlooked by anyone not familiar with them. (The single Heterodontus
tooth found with the aid of the microscope was a lateral.) Although finding
the teeth of two additional elasmobranch species in my relatively small field
sample is not conclusive proof that sorting for these remains without a micro-
scope is inefficient, recent experiments have shown that a microscope (or some
similar magnifying device) is an absolute necessity for finding the tiny teeth
of seven of California’s commonest inshore elasmobranchs. In these experi-
ments, I found that all of the teeth of the swell shark, Cephaloscy Ilium uter,
shovelnose guitarfish, Rhinobatos productus, banded guitarfish, Zapteryx exas-
perata, thornback, Platyrhinoidis triseriata, electric ray, Torpedo californica,
butterfly ray, Gymnura marmorata, and round stingray, Urolophus halleri,
will pass through an 18-mesh screen (1 mm, U.S. Standard Sieve Series), as
will most of the teeth of the gray smoothhound, Mustelus calif ornicus. These
small teeth would be impossible to find with the naked eye, particularly when
one considers they would comprise an infinitesimal part of the sample being
searched.
Systematic Account
Heterodontidae— horn sharks
Heterodontus francisci— horn shark
Horn sharks are abundant in rocky subtidal areas between about Morro
Bay, California, and Magdalena Bay, Baja California. They seldom move
about during daylight hours, but at night they can be found foraging the
bottom for food, primarily crustaceans. During their nocturnal feeding, they
occasionally stray short distances away from their preferred rocky habitat,
into areas of sandy or sandy-mud substrate.
Material : 1 lateral tooth.
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Alopidae— thresher sharks
Alopias vulpinus — thresher shark
The thresher shark previously was reported from the Playa del Rey site
based upon two teeth (Fitch, 1964).
Additional material: 1 tooth.
Isuridae— mako sharks
lsurus oxyrinchus— mako
The mako was previously reported from this site as lsurus glaucus
based upon two teeth (Fitch, 1964).
Additional material : 2 teeth.
Carcharhinidae— requiem sharks
Carcharhinus sp.— carcharhinid shark, species undetermined
Unidentified requiem sharks were previously reported from this site
based upon 26 teeth (Fitch, 1964). The additional teeth found in the re-
sampling probably came from one of the same unidentified species.
Additional material : 7 teeth.
Galeorhinus zyopterus—sowpfin shark
Soupfin shark teeth (15) previously were reported from this site (Fitqh,
1964).
Additional material : 1 tooth.
Sphyrnidae— hammerhead sharks
Sphyrna sp.— hammerhead shark, species undetermined
The 3 hammerhead shark teeth and 17 vertebrae previously reported
from this site were not identified to species (Fitch, 1964), nor are the teeth
from my resampling.
Additional material : 6 teeth.
Squatinidae— angel sharks
Squatina calif ornica— Pacific angel shark
The Pacific angel shark previously was reported from this deposit based
upon 10 teeth and 8 vertebrae (Fitch, 1964).
Additional material : 6 teeth.
Rajidae— skates
Raja spp.— skates, species undetermined
Unidentified skate remains (8 vertebrae and 3 “wing” spines) have been
reported from this site (Fitch, 1964).
Additional material : 4 “wing” spines.
1966
Pleistocene Fish Remains
7
Dasyatidae— stingrays
Dasyatis dipterurus— diamond stingray
Diamond stingrays have been reported from as far north as British
Columbia, but their occurrence north of Point Conception can be considered
unusual. They range south at least to Central America and possibly to Peru.
A large female caught in Los Angeles Harbor in 1963 was about 5 feet 2
inches long (34 inches across the disk) and weighed 113 Vi pounds.
Material : 2 teeth.
Urolophus halleri— round stingray
Although it is difficult, if not impossible, to distinguish broken and worn
fragments of the caudal “sting” of round stingrays from those of bat stingrays
or juvenile diamond stingrays, the 164 stings previously reported from this
deposit were assigned to U. halleri (Fitch, 1964). The teeth of the round
stingray are extremely small (they will pass through an 18-mesh screen) so
even if they occur in a deposit, they would not be found unless residue retained
by a 30- or 32-mesh screen was carefully examined under a microscope.
Additional material: 5 stings.
Myliobatidae— eagle rays
Myliobatis calif ornicus— bat stingray
The bat stingray previously has been reported from this deposit based
upon 205 teeth (Fitch, 1964).
Additional material: 32 teeth.
Unidentified elasmobranchs
Material: No attempt was made to assign to species the 15 elasmobranch
vertebrae found during resampling.
Engraulidae— anchovies
Engraulis mordax— northern anchovy
Sixty-eight northern anchovy otoliths were reported for this deposit by
Fitch (1964). Use of the microscope in searching the residue greatly in-
creased the yield of Engraulis otoliths.
Additional material: 194 otoliths.
Anchoa compressa— deepbody anchovy
The range of the deepbody anchovy appears to be quite restricted (Morro
Bay, California, to Todos Santos Bay, Baja California). The species seldom
attains 5 inches in length, and 40 or 50 large individuals are required for a
pound of weight. They usually inhabit quiet waters of back bays and sloughs,
but have been recorded from relatively sheltered areas of the open coast, in-
cluding Santa Monica Bay.
Material: 4 otoliths (Fig. 8).
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Contributions in Science
No. 119
1966
Pleistocene Fish Remains
9
Argentinidae— argentines
Argentina sialis— Pacific argentine
The Pacific argentine ranges from off San Francisco south into the Gulf
of California. Although they have been considered a bathypelagic species,
they probably are most abundant just above the bottom in water shallower
than 1,000 feet. Small schools of Argentina sialis have been photographed in
these depths from a diving saucer, and the stomachs of occasional large in-
dividuals trawled from 600 to 800 feet of water contain bottom-living or-
ganisms. An 814 -inch female weighed slightly less than 2 ounces (53 grams).
Material : 1 otolith (Fig. 1 1 ) showing signs of having been in the stomach
of some predatory species (i.e., general erosion of all surfaces including
concave).
Myctophidae— lanternfishes
Symbolophorus californiense—C alifornia lanternfish
S. calif orniense is a bathypelagic species that ranges throughout the north
Pacific Ocean— from British Columbia to below Ensenada in the eastern
Pacific. A large individual might be 5 inches long and weigh about one-half
ounce.
Material: 1 otolith (Fig. 3) in good condition.
Stenobrachius leucopsarus— northern lampfish
S. leucopsarus is one of the most abundant bathypelagic fishes in the
eastern Pacific, where it ranges from the Bering Sea to about Cedros Island,
Baja California. A large individual might be 5 inches long and weigh about
one-half ounce.
Material: 1 otolith (Fig. 5), somewhat eroded, as if from the stomach of
a predator.
Figure 1. Inner face, left sagitta (badly worn, rostrum missing) of Trachurus sym-
metries 6.7 mm long.
Figure 2. Inner face, left sagitta (badly worn) of Zaniolepis latipinnis 3.3 mm long.
Figure 3. Inner face, right sagitta of Symbolophorus californiense 4.4 mm long.
Figure 4. Inner face, right sagitta of Prionotus stephanophry s 6.8 mm long.
Figure 5. Inner face, left sagitta (badly worn) of Stenobrachius leucopsarus 2.0 mm
long.
Figure 6. Inner face, left sagitta of Icelinus quadriseriatus 2.8 mm long.
Figure 7. Inner face, left sagitta of Coelorhynchus scaphopsis 7.9 mm long.
Figure 8. Inner face, right sagitta of Anchoa compressa 1.9 mm long.
Figure 9. Inner face, right sagitta of Symphurus atricauda 2.5 mm long.
Figure 10. Inner face, right sagitta of Occa verrucosa 3.8 mm long.
Figure 11. Inner face, left sagitta (badly worn) of Argentina sialis 3.0 mm long.
Figure 12. Inner face, right sagitta (posterior portion) of Paralabrax sp. 7.8 mm
long.
Photographs by Jack W. Schott.
10
Contributions in Science
No. 119
Macrouridae ( =Coryphaenoididae ) — rattails
Coelorhynchus scaphopsis— Gulf rattail
This species is one of the commonest rattails in the northern Gulf of
California, where it can be trawled on the bottom in depths greater than 100
fathoms. During recent years, a single individual was caught off Santa Barbara,
California, in an otter trawl. No measurements are available for the Gulf
rattail, but none of several dozen observed during the past 10 years exceeded
15 inches total length.
Material : 1 otolith (Fig. 7) in good condition.
Merlucciidae— hakes
Merluccius productus— Pacific hake
Pacific hake otoliths (24) previously were reported from the Playa del
Rey deposit (Fitch, 1964).
Additional material'. 11 otoliths.
Bothidae— lefteyed flounders
Paralichthys californicus—C alifornia halibut
A single broken California halibut otolith was reported from this site by
Fitch (1964).
Additional material : 5 otoliths.
Citharichthys spp.— sanddabs, species as listed below.
The otoliths of all three sanddabs known to Californian waters are easily
distinguished if they are in good condition. Those of C. stigmaeus have
straight margins and never attain large sizes. C. sordidus and C. xanthostigma
otoliths have rounded margins, and at maximum size are two to three times
larger than C. stigmaeus otoliths. The antero-dorsal margin of the otolith of
C. sordidus is sharply notched, distinguishing it from C. xanthostigma. Otoliths
(290) of C. sordidus and C. stigmaeus were reported from this deposit by
Fitch (1964).
Additional material'. 235 otoliths— 7 from C. sordidus, 181 from C. stig-
maeus, and 47 undeterminable.
Pleuronectidae— righteyed flounders
Parophrys vetulus— English sole
A single English sole otolith previously was reported from this deposit
(Fitch, 1964).
Additional material'. 3 otoliths.
1966
Pleistocene Fish Remains
11
Cynoglossidae— tonguefishes
Symphurus California tonguefish
The California tonguefish lives on sandy or sandy-mud bottoms at moder-
ate depths between Big Lagoon, northern California and about Magdalena
Bay, Baja California. Greatest concentrations seem to be in water shallower
than 150 feet. A very large individual was 7 Vi s inches long; its weight was not
recorded but probably was not in excess of 3 ounces. Because of their small
size and odd shape, Symphurus otoliths probably would not be found without
using a microscope.
Material: 13 otoliths (Fig. 9).
Serranidae— basses
Paralabrax spp.— kelp and sand basses
Three species of Paralabrax are abundant in shallow waters along the
southern California coast. The kelp bass, P. clathratus, prefers rocky habitat,
especially where kelp beds are prolific. Of the other two, P. nebulifer (sand
bass) and P. maculatofasciatus (spotted sand bass), P. nebulifer is the most
likely to be found in quiet offshore waters where the bottom is sandy or sandy-
mud. P. maculatofasciatus seems to prefer the habitats found in bays and
estuaries. Paralabrax otoliths are difficult, if not impossible, to distinguish to
species even when they are in perfect condition. None of the Paralabrax
otoliths found in resampling was complete.
Material'. 3 otoliths, all with anterior ends missing (Fig. 12).
Atherinidae— silversides
Leuresthes tenuis— grunion
Grunion otoliths (6) previously were reported from the Playa del Rey
deposit by Fitch (1964).
Additional material: 4 otoliths.
Atherinopsis calif or niensis— jacksmelt
Three jacksmelt otoliths previously were reported from this locality
(Fitch, 1964).
Additional material’. 1 otolith.
Atherinids— species undetermined
The only other silverside known to California is the topsmelt, Atherinops
affinis. A single topsmelt otolith previously was reported from this deposit
(Fitch, 1964). Several broken atherinid otoliths found during resampling
probably were from A. affinis, but were not sufficiently entire to make a posi-
tive identification.
Additional material: 10 broken otoliths.
12
Contributions in Science
No. 119
Carangidae— jacks
Trachurus symmetricus— Pacific jackmackerel
The Pacific jackmackerel is a schooling species that ranges from British
Columbia to Cape San Lucas and offshore for several hundred miles. A large
individual might be 30 inches long and weigh 5 or 6 pounds, but the com-
mercial catch, comprising thousands of tons each year, consists mostly of
15-inch and smaller fish.
Material: 3 otoliths (Fig. 1) in poor condition, possibly from having
been partially digested in the stomachs of predators.
Sciaenidae—croakers
Cynoscion reticulatus— striped corvina
The otoliths (4) of this southern species previously were reported for
this site (Fitch, 1964).
Additional material : 1 otolith.
Genyonemus lineatus— white croaker
White croaker otoliths were abundant in the original sampling of the
Playa del Rey deposit, 217 having been reported (Fitch, 1964).
Additional material : 116 otoliths.
Menticirrhus undulatus— California corbina
The California corbina previously was reported from this locality based
upon the broken posterior half of a single otolith (Fitch, 1964).
Additional material : 5 otoliths.
Seriphus politus— queenfish
The 275 queenfish otoliths reported from this site (Fitch, 1964) repre-
sented over 20 percent of the total otoliths on hand at that time.
Additional material : 116 otoliths.
Umbrina roncador—yeMowfin croaker
Yellowfin croaker otoliths (13) previously were reported from this
deposit (Fitch, 1964).
Additional material : 4 otoliths.
Embiotocidae— surfperches
Cymatogaster aggregata— shiner perch
Shiner perch otoliths (6) previously were reported from the Playa del
Rey locality (Fitch, 1964).
Additional material: 10 otoliths.
1966
Pleistocene Fish Remains
13
Scorpaenidae— rockfishes
Sebastodes spp.— rockfishes, species undetermined
The otoliths of most of the 52 species of Sebastodes inhabiting the waters
of California can be distinguished from each other if they are in perfect or
near-perfect condition. Such characters as length and shape of rostrum, con-
figuration of posterior end, angle of posterior taper, depth of sulcus, and
number of growth zones (annuli) for otolith size are helpful for determining
species or species-complex. Although five species were identified from among
the 25 Sebastodes otoliths previously reported upon (Fitch, 1964), the rock-
fish otoliths obtained from resampling the deposit were in such poor condition
it was impossible to determine the species involved.
Additional material : 8 otoliths.
Zaniolepidae— combfishes
Zaniolepis latipinnis— longspine combfish
The longspine combfish is fairly abundant in moderate depths (50 to 400
feet) on sandy-mud bottoms between Puget Sound and San Martin Island,
Baja California (at least). A large individual may exceed 10 inches in length,
but weights for a combfish that size are unavailable. An 8 Vi -inch specimen
weighed nearly 3 ounces (72 grams).
Material : 1 otolith (Fig. 2).
Cottidae— sculpins
Chitonotus pugetensis— roughback sculpin
One roughback sculpin otolith previously was reported from this site
(Fitch, 1964).
Additional material : 7 otoliths.
Icelinus quadriseriatus— yellowchin sculpin
The yellowchin sculpin is one of the most abundant members of the
family in moderate depths (50 to 250 feet) between about Pt. Reyes, Califor-
nia, and Cape San Lucas, Baja California. They never attain very large sizes,
about 3 Vi inches being maximum, and their otoliths are small enough that a
microscope is needed to recover them. Three other species of Icelinus are fairly
common in the same general areas as /. quadriseriatus, but only the otoliths
of I. tenuis are difficult to distinguish from those of the yellowchin sculpin.
Some of the Icelinus otoliths found during resampling could have been from
/. tenuis, but I. tenuis is less abundant in our area and typically inhabits some-
what deeper water, so I assigned all of the present material to I. quadriseriatus.
Material : 10 otoliths (Fig. 6), some in rather poor condition.
14
Contributions in Science
No. 119
Triglidae— gurnards
Prionotus stephanophrys— lumptail searobin
Although the lumptail searobin has been reported from as far north as
San Francisco, its occurrence in California is sporadic and usually during years
when water temperatures are higher than normal. It is relatively abundant in
Mexican waters, and occurs in the Gulf of California with several other mem-
bers of the genus. A very large individual from southern California was 15Vi
inches long and weighed just over 2 pounds.
Material : 2 otoliths (Fig. 4).
Agonidae— poachers
Occa verrucosa— warty poacher
The warty poacher ranges from about Vancouver Island to Point Con-
ception, where it is found on sandy-mud bottoms in moderate depths (30 to
150 feet or so). A large individual might be 8 inches long and weigh 2 ounces,
but most are much smaller. This appears to be the only species from the Playa
del Rey deposit that has not been reported south to the same latitude during
recent times. The lack of recent records from south of Point Conception may
be a reflection of inadequate sampling with small-mesh trawl nets in the right
depths, but several hundred tows with small-mesh otter trawls in Santa Monica
Bay during the past decade failed to yield a single warty poacher.
Material : 1 otolith (Fig. 10).
Gobiidae— gobies
Lepidogobius lepidus— bay goby
One otolith from a bay goby previously was found in this deposit (Fitch,
1964).
Additional material : 8 otoliths.
Batrachoididae— toadfishes
Porichthys myriaster— specklefin midshipman
Specklefin midshipman otoliths (15) previously have been reported from
this locality (Fitch, 1964).
Additional material : 3 otoliths.
Porichthys notatus— plainfin midshipman
Plainfin midshipman otoliths (161) previously were reported from this
deposit (Fitch, 1964).
Additional material : 278 otoliths.
1966
Pleistocene Fish Remains
15
Ophidiidae— cusk-eels
Otophidium scrippsae— basketweave cusk-eel
O. scrippsae otoliths were nearly twice as abundant as the otoliths of
O. taylori in the earlier samples from this site, 140 having been reported as
O. scrippsae by Fitch (1964). This ratio did not hold up during resampling.
Additional material : 49 otoliths.
Otophidium taylori— spotted cusk-eel
Eighty-three otoliths of O. taylori were reported from this site by Fitch
(1964).
Additional material: 65 otoliths.
Discussion
The fish remains found during resampling of the Playa del Rey deposit
represent 11 species (at least) of sharks, skates, and rays belonging to nine
families, and 34 or more species of bony fishes belonging to 21 families. Two
of the elasmobranchs and 12 of the bony fishes had not previously been found
at this locality, and brought the known number of species from this deposit to
62 (at least). The elasmobranch remains consisted of 58 teeth, 15 vertebrae,
5 caudal “stings’’ and 4 “wing” spines, whereas the teleost remains consisted
of 1,182 otoliths, 32 vertebrae, 18 teeth, 2 fin spines, and 1 jaw fragment.
These remains do not detract from the contention that the habitat (at the
time and place of deposition) was an area of fine grained silty sand, typical
of a quiet-water embayment 10 to 12 fathoms deep (Valentine, 1961), even
though two of the species ( Stenobrachius leucopsarus and Symbolophorus
calif orniense) are bathypelagic forms that almost never are captured where
the water is shallower than 1,000 feet. Two other species ( Coelorhynchus
scaphopsis and Argentina sialis) usually are taken at or near the bottom in
600 feet or more of water. The otoliths of these four species are quite rare
in the deposit (only one of each was found), and possibly all were carried
there in the stomachs of predators or scavengers. Dead fish, including deep-sea
species, that float to the surface usually are picked up by gulls and other
scavenging birds which are capable of traveling great distances in a short time
after eating such a meal. Otoliths that pass through gulls, terns, and other
fish-eating birds, often are scarcely altered by digestive action (Martini, 1964).
The finding of Coelorhynchus scaphopsis and Prionotus stephanophrys
otoliths during the resampling brought to six the number of southern fish
species in this deposit that seldom, if ever, are taken north of Mexico. Previ-
ously, a tooth from Rhizoprionodon (formerly Scoliodon ) longurio, and oto-
liths of Micropogon ectenes, Cynoscion reticulatus, and Lepophidium negro-
pinna had been reported from this site (Fitch, 1964). An otolith of Cynoscion
reticulatus also was found during resampling. These southern species lend
additional support to the contention that this deposit was laid down at a time
16
Contributions in Science
No. 119
when local ocean temperatures were considerably higher than is normal for
our area at present.
Finally, the excellent yield of otoliths, large as well as small, from the
relatively small field sample obtained, points up the need for using a micro-
scope to search out fish remains in fossil deposits.
Acknowledgments
This study was supported in part by a research grant (GB-1244) from the
National Science Foundation. In addition, Richard A. Fitch resampled the
Playa del Rey deposit for me, and washed and screened the field sample he
brought back. Jack W. Schott took the excellent photographs used to illustrate
this paper, and Mrs. Loretta Morris typed the final draft of the manuscript.
Literature Cited
Fitch, John E.
1964. The fish fauna of the Playa del Rey locality, a southern California
marine Pleistocene deposit. Los Angeles County Mus., Cont. in Sci.,
82: 1-35.
Martini, Erlend
1964. Otolithen in Gewollen der Raubseeschwalbe ( Hydroprogne caspia ).
Bonner Zoologische Beitrage, 15(1 & 2) : 59-71.
Valentine, James W.
1961. Paleoecologic molluscan geography of the Californian Pleistocene.
Univ. Calif., Publ. Geol. Sci., 34(7): 309-442.
LOS
ANGELES
COUNTY
MUSEUM
CONTRIBUTIONS
IN SCIENCE
number 120 December 31, 1966
A NEW SPECIES OF DIOPTOPSIS FROM CALIFORNIA
(DIPTERA: BLEPH AROCERIDAE )
1
i
By Charles L. FIogue
■I
i
Los Angeles County Museum of Natural History • Exposition Park
Los Angeles, California 90007
CONTRIBUTIONS IN SCIENCE is a series of miscellaneous technical papers
in the fields of Biology, Geology and Anthropology, published at irregular intervals
by the Los Angeles County Museum of Natural History. Issues are numbered sepa-
rately, and numbers run consecutively regardless of subject matter. Number 1 was
issued January 23, 1957. The series is available to scientific institutions on an ex-
change basis. Copies may also be purchased at a nominal price.
INSTRUCTIONS FOR AUTHORS
Manuscripts for the LOS ANGELES COUNTY MUSEUM CONTRIBU
TIONS IN SCIENCE may be in any field of Life or Earth Sciences. Acceptance of
papers will be determined by the amount and character of new information and the
form in which it is presented. Priority will be given to manuscripts by staff members,
or to papers dealing largely with specimens in the Museum’s collections. Manuscripts
must conform to CONTRIBUTIONS style and will be examined for suitability by
an Editorial Committee. They may also be subject to critical review by competent
specialists.
MANUSCRIPT FORM.— (1) The 1960 AIBS Style Manual for Biological
Journals is highly recommended as a guide. (2) Typewrite material, using double
spacing throughout and leaving ample margins, on only one side of 8V2 x 11 inch
standard weight paper. (3) Place tables on separate pages. (4) Footnotes should be
avoided if possible. (5) Legends for figures and unavoidable footnotes should be
typed on separate sheets. Several of one kind may be placed on a sheet. (6) Method
of literature citation must conform to CONTRIBUTIONS style— see number 90 and
later issues. Spell out in full the title of non-English serials and places of publication.
(7) A factual summary is recommended for longer papers. (8) A brief abstract must
be included for all papers. This will be published at the head of each paper.
ILLUSTRATIONS. — All illustrations, including maps and photographs, should
be referred to as “figures!’ All illustrations should be of sufficient clarity and in the
proper proportions for reduction to CONTRIBUTIONS page size. Permanent ink
should be used in making line drawings and in lettering (do not type on drawings);
photographs should be glossy prints of good contrast. Original illustrations will not
be returned unless specifically requested when the manuscript is first submitted.
PROOF.- — Author will be sent galley proof which should be corrected and re-
turned promptly. Changes after the paper is in galley will be billed to the author. Un-
less specially requested, page proof will not be sent to the author. 100 copies of each
paper will be given free to a single author or divided equally among multiple authors.
Orders for additional copies should be sent to the Editor at the time corrected galley
proof is returned; appropriate forms for this will be included when galley is sent.
David K. Caldwell
Editor
A NEW SPECIES OF DIOPTOPSIS FROM CALIFORNIA
(DIPTERA: BLEPHAROCERIDAE )
By Charles L. Hogue1
Abstract: Described in the genus Dioptopsis is a new
species of net-winged midge, D. alpina. Adult males, females
and pupae are recorded only from northern California. The
species is most closely related to D. arizonico Alexander, from
Arizona.
In order to make its name available for a forthcoming review of the
California Blepharoceridae planned for the Bulletin of the California Insect
Survey, I am describing the following new species in the genus Dioptopsis.
I am grateful to Dr. J. Powell, University of California, Berkeley-Cali-
fornia Insect Survey [CIS] and to Dr. Alan Stone, Entomology Research
Division, A.R.S., United States Department of Agriculture (United States
National Museum) [USNM] for making available to me the material upon
which this species is based. Thanks also are due Mrs. L. McTernan for assist-
ance in the preparation of the illustrations.
Dioptopsis alpina, new species
Figures 1 through 9
Adult female :
Size. Small: measurements as
follows (lengths
in mm
from allotype) :
Overall body: 5
Wing: 8.5
Legs:
fore
mid
hind
femur
4.6
5.2
6.1
tibia
4.3
4.3
5.6
tarsis 1
2.51
1.86
2.16
2
.96
.80
.81
3
.53
.47
.43
4
.30
.31
.29
5
.48
.49
.41
Coloration. General integument evenly testaceous, lightly sclerotized.
Mesoscutum gray, finely pruinose; scute! lum and pleuron clear yellow, like-
wise finely pruinose. Abdominal sternites pale yellow, tergites gray like
mesoscutum with pale yellow caudal borders. Legs testaceous proximad,
Curator of Entomology, Los Angeles County Museum of Natural History.
1
2
Contributions in Science
No. 120
1966
New Species of California Diptera
3
Dioptopsis alpina
becoming moderately darker distad. No conspicuous markings of any kind
on any part of body or appendages.
Head. As figured (Fig. 3). Eyes widely separated; upper division of eye
very small, comprising only a poorly differentiated hemi-lenticular area of
4 to 5 rows of ommatidia; ommatidia of both eye divisions equal in size. An-
tennae short; 15 segmented; shape and size of flagellar segments about equal
except basal which is 1.4 length of adjoining distal segment, ultimate segment
slightly smaller than penultimate. Mouthparts with normal elements, mandi-
bles present; hypopharynx with well-developed marginal teeth, those on the
tip about half the size of lateral.
Thorax. Scutellum with small, dense setal patches at outer corners;
scattered, fine setae mesad across posterior border. A few short, fine setae on
lateral wings of anterior pronotum. Other sclerites nude. Legs with tibial
spurs 0-1-2, spur of mid tibia very small; fore basitarsus somewhat bowed
upward in allotype (artifact?). Wing shape, venation, and trichiation as
figured (Fig. 1); membrane entirely hyaline.
Genitalia. As figured (Fig. 6). Spermathecae elongate, slightly swollen
cephalad, one slightly smaller than other two.
4
Contributions in Science
No. 120
Adult male :
Size. Small: slightly smaller than female; measurements as follows
(lengths in mm from holotype) :
Overall body: 4.5
Wing: 6.4
fore
mid
hind
femur
3.4
4.0
5.0
tibia
3.4
3.5
2.7
tarsis
1
2.1
1.72
1.78
2
.84
.77
.64
3
.58
.54
.39
4
.36
.36
.28
5
.36
.36
.30
Coloration. Essentially as in female.
Head. As figured (Fig. 2). Eyes widely separated; essentially undivided,
upper division even less well-differentiated than in female. Antennae short;
15 segmented; shape and size of flagellar segments about equal except basal
which is narrowed proximad and 1.4 length of adjoining distal segment; ulti-
mate segment slightly smaller than penultimate. Mouthparts with mandibles
absent. Tip of labrum slightly emarginate; hypopharynx evenly tapering to a
narrowly truncate tip, lateral margins with obsolescent teeth.
Thorax. Thoracic chaetotaxy as in female. Legs with tibial spurs 0-1-2,
mid small as in female; fore basitarsus straight. Wing as in female.
Genitalia. As figured ( Figs. 4 and 5 ) .
Pupa :
Size. Medium, length typical specimen: 6.2 mm. Interbranchial index
(see Fig. 9 for definition) = .86.
Structure. As figured (Figs. 7 to 9). General shape subfusiform. Trans-
verse dorsal mesoscutal rugae zigzagging. Branchial sclerite beaded and
strongly swollen cephalad, projecting laterally over the frontal sclerite (Fig. 8).
Material :
Holotype male (genitalia, head and wing on slides Nos. CLH 641 105-3,
6607-1, and 6608-2 resp.): Alpine Co., Lake Alpine, 15 July 1950 (L. W.
Quate) [USNM].
Allotype female (genitalia and head on slides Nos. CLH 650504-6 and
6607-2 resp.) : same data as holotype.
Additional specimens: 11 pupae and pupal skins: same data as types.
16 pupal skins: El Dorado Co., Fred’s Place, 5 July 1950 (L. W. Quate) [CIS].
1966
New Species of California Diptera
5
Diagnosis :
Dioptopsis alpina is most similar to D. arizonica Alexander, 1958, de-
scribed from the Sierra Ancha Mountains of Arizona. The two species form
a distinctive division of the genus characterized by nearly undivided eyes in
both sexes, IX tergite lobes of the male genitalia apically truncate and very
short (nearly as broad as long), spermathecae elongate and the mid tibia with
only one, tiny spur.
Unfortunately, arizonica is known only from a single male, which is a
poor specimen, having been in a teneral state when collected. It is therefore
not possible to complete the diagnosis with alpina. Essential differences are
apparent nevertheless in the male genitalia (Figs. 4 and 5) especially in regard
to the shape of the tip of the inner lobe of the dististyle (Figs. 4a and 6).
There is a cup-shaped depression or apodeme here in arizonica ; this lobe is
simply recurved like a button-hook in alpina.
CONTRIBUTIONS
IN SCIENCE
LOS
ANGELES
COUNTY
MUSEUM
UMBER 121
December 31, 1966
0 7
By James R. Dixon and Philip A, Medica
SUMMER FOOD OF FOUR SPECIES OF LIZARDS
FROM THE VICINITY OF WHITE SANDS, NEW MEXICO
Los Angeles County Museum of Natural History • Exposition Park
Los Angeles, California 90007
CONTRIBUTIONS IN SCIENCE is a series of miscellaneous technical papers
in the fields of Biology, Geology and Anthropology, published at irregular intervals
by the Los Angeles County Museum of Natural History. Issues are numbered sepa-
rately, and numbers run consecutively regardless of subject matter. Number 1 was
issued January 23, 1957. The series is available to scientific institutions on an ex-
change basis. Copies may also be purchased at a nominal price.
INSTRUCTIONS FOR AUTHORS
Manuscripts for the LOS ANGELES COUNTY MUSEUM CONTRIBU-
TIONS IN SCIENCE may be in any field of Life or Earth Sciences. Acceptance of
papers will be determined by the amount and character of new information and the
form in which it is presented. Priority will be given to manuscripts by staff members,
or to papers dealing largely with specimens in the Museum’s collections. Manuscripts
must conform to CONTRIBUTIONS style and will be examined for suitability by
an Editorial Committee. They may also be subject to critical review by competent
specialists.
MANUSCRIPT FORM.— (1) The 1960 AIBS Style Manual for Biological
Journals is highly recommended as a guide. (2) Typewrite material, using double
spacing throughout and leaving ample margins, on only one side of 8 Vi x 1 1 inch
standard weight paper. (3) Place tables on separate pages. (4) Footnotes should be
avoided if possible. (5) Legends for figures and unavoidable footnotes should be
typed on separate sheets. Several of one kind may be placed on a sheet. (6) Method
of literature citation must conform to CONTRIBUTIONS style— see number 90 and
later issues. Spell out in full the title of non-English serials and places of publication.
(7) A factual summary is recommended for longer papers. (8) A brief abstract must
be included for all papers. This will be published at the head of each paper.
ILLUSTRATIONS. — All illustrations, including maps and photographs, should
be referred to as “figures!’ All illustrations should be of sufficient clarity and in the
proper proportions for reduction to CONTRIBUTIONS page size. Permanent ink
should be used in making line drawings and in lettering (do not type on drawings);
photographs should be glossy prints of good contrast. Original illustrations will not
be returned unless specifically requested when the manuscript is first submitted.
PROOF. — Author will be sent galley proof which should be corrected and re-
turned promptly. Changes after the paper is in galley will be billed to the author. Un-
less specially requested, page proof will not be sent to the author. 100 copies of each
paper will be given free to a single author or divided equally among multiple authors.
Orders for additional copies should be sent to the Editor at the time corrected galley
proof is returned; appropriate forms for this will be included when galley is sent.
David K. Caldwell
Editor
SUMMER FOOD OF FOUR SPECIES OF LIZARDS
FROM THE VICINITY OF WHITE SANDS, NEW MEXICO
By James R. Dixon1 and Philip A. Medica2
Abstract: A total of 3 1 1 stomachs of four species of lizards
from the vicinity of White Sands, New Mexico, were examined
for their food items. A graphic representation of the summer
food of the four species is given along with a description of the
foraging activities for each species. An explanation is presented
to account for the absence of Uta in the major portion of the
dunes.
In conjunction with a study of the biology of the lizards of White Sands,
New Mexico, 311 stomachs of four species of lizards collected during the summer
months of 1963 were examined for their food items.
Uta stansburiana is the only lizard examined that is not found in the
major portion of the gypsum soils. Uta is apparently restricted to the more
stabilized gypsum dunes along the periphery of the active dunes. This species
seldom occupies the gypsum dunes for more than 30 meters from the edge of
the adobe soil. The lizards that occupy the major portion of the active dune
area are Holbrookia maculata, Sceloporus undulatus, and Cnemidophorus
inornatus.
Of 31 1 stomachs only 275 contained food. The species, with their respec-
tive number of empty stomachs are Holbrookia , 9; Sceloporus , 26, Cnemido-
phorus, 3.
Methods'. The total volume of each stomach was taken by placing it in a
volumetric flask and measuring the millimeters of water displaced. The con-
tents were then removed and the volume of water displaced by the stomach
lining was recorded. The volume of food items was determined by subtracting
the latter figure from the former. The analysis of the various food items in the
stomach was obtained by using a binocular dissecting microscope. The vol-
umes for each food item were then given an ocular estimate as they were
separated. All of the arthropods were keyed to family wherever possible. In
some instances where only parts of arthropods were found it was possible only
to key to order. No attempt was made to separate the stomachs into the
monthly periods when the lizards were taken. The percentages of the orders
of arthropods eaten by each species of lizard were totaled for a three-month
summer period and graphically diagrammed (Figs. 1 through 4).
Food and Foraging Activity : The food items of the three sympatric
species of lizards inhabiting the gypsum soils are similar but of varying per-
centages. Holbrookia (Fig. 1) eat essentially the same food items as Scelo-
porus (Fig. 2), but the methods employed in catching food and the micro-
habitat occupied belies the factor of direct competition.
Curator of Herpetology, Los Angeles County Museum of Natural History,
department of Zoology, Brigham Young University, Provo, Utah.
1
2
Contributions in Science
No. 121
Holbrookia are often seen foraging in open spaces between vegetation in
the dune depressions and on the open dunes. They have been observed taking
Coleoptera, Lepidoptera and Hymenoptera from the small flowers of scorpion-
weed, a low prostrate plant commonly found in the dune depressions. Hol-
brookia were observed pursuing slow flying insects across open dunes, and
feeding upon ants near the ant colony site.
Sceloporus were seldom seen in open spaces about vegetation. This lizard
was observed on several occasions feeding on the ground immediately beneath
rabbit bush, yucca, mormon tea, and in clumps of alkali sacaton. Sceloporus
were often seen climbing about on the branches of shrubs, indicating an ability
HOLBROOKIA MACULATA
Figure 1 . Per cent by volume of food items of 82 stomachs of Holbrookia maculata
taken from the White Sands of New Mexico, during the summer of 1963.
1966
Summer Food of Lizards
3
SGELOPORUS UNDULATUS
Figure 2. Per cent by volume of food items of 1 17 stomachs of Sceloporus undulatus
taken from the White Sands of New Mexico, during the summer of 1963.
to take insects that may be on the shrubs some distance above the ground.
Holbrookia were never seen climbing in shrubs or herbs.
The major food items of Holbrookia and Sceloporus consisted of Hy-
menoptera and Coleoptera. However, Holbrookia consumed larger quantities
of other orders of insects available to them than did Sceloporus. This is
probably accounted for by their foraging in a greater variety of microhabitats
than Sceloporus. Holbrookia are lighter in color and color pattern than Scelo-
porus, a factor that allows them a distinct advantage in food gathering in the
open dunes without being seen by a predator, and thus a wider variety of food
is available to them.
4
Contributions in Science
No. 121
A few of the stomachs of Sceloporus contained the tails of young Cnemi-
dophorus, and two Cnemidophorus stomachs contained hatchlings of Scelo-
porus, indicating that the two genera may feed on the young of one another.
In addition, some Holbrookia stomachs contained partly digested remains of
both Sceloporus and Cnemidophorus.
The food eaten by Cnemidophorus inornatus (Fig. 3) consists principally
of Lepidoptera and Coleoptera larvae and adult Coleoptera. Cnemidophorus
forage about the litter beneath vegetation for food. This species usually uses
its nose and front feet to turn over pieces of bark, dead leaves and twigs in its
quest for food. This method of feeding accounts for the high percentage of
Figure 3. Per cent by volume of food items of 73 stomachs of Cnemidophorus inor-
natus taken from the White Sands of New Mexico, during the summer of 1963.
1966
Summer Food of Lizards
5
UTA STANSBURIANA
Figure 4. Per cent by volume of food items of 45 stomachs of Uta stansburiana
taken from the White Sands of New Mexico, during the summer of 1963.
larvae in the diet, and perhaps a preference for this type of food. This method
of feeding indicates a lack of direct competition for food with Sceloporus and
Holbrookia, both of which pursue surface and flying arthropods.
A few Cnemidophorus were observed catching butterflies and skippers by
leaping in the air and capturing the insects as they alighted upon the flowers of
low prostrate plants. More often than not, they missed their desired target, but
occasionally they were able to catch a few insects by this method.
The major food item eaten by Uta stansburiana was Hymenoptera, con-
sisting principally of ants and small wasps. Uta are ground dwelling lizards
and none were observed climbing in vegetation. Wasps were not observed be-
6
Contributions in Science
No. 121
ing eaten by these lizards, but we assume that the wasps were captured by the
lizards while on the ground. Uta are abundant near the periphery of the dunes
in adobe flats that are densely covered with tar bush, rabbit bush, creosote
bush, iodine bush, alkali sacaton and gramma grass. Only a few other species
of lizards were seen in the latter plant association, and none of these were
abundant. Those species seen, but in small numbers, are Cnemidophorus
neomexicanus, Cnemidophorus inornatus, Sceloporus undulatus , Sceloporus
magister, Crotaphytus collaris, and Phrynosoma cornutum. The food prefer-
ences of the latter lizards are not known for this area, but Ota were abundant,
indicating a greater survival rate than for the other species.
A comparison of Uta food items (Fig. 4) with those of Sceloporus on
the dunes indicates that they feed upon the same quality of food. Although
Uta take less Coleoptera and no Lepidoptera, their absence from the dunes
may be attributed to competitive food gathering with Sceloporus and Hol-
brookia. However, the color and color pattern of the Uta population inhabit-
ing the fringe area of the dunes is identical to its counterpart population on
the adobe soils. Therefore they are at a distinct disadvantage in securing food
on the dunes without being seen by a predator. This factor may account for
the absence of Uta on the active dunes, rather than selective food competition
with other lizards adapted to the white gypsum soil.
Acknowledgments
We wish to thank Mr. Forrest M. Benson, Superintendent, and Mr.
Robert L. Morris, Chief Ranger, both of the National Park Service in charge
of the White Sand National Monument, for their invaluable aid in securing
permission for us to conduct our investigations on the National Monument
grounds; and we wish to acknowledge a grant from the Grants-in-aid of Re-
search Committee of the Society of Sigma Xi for travel and other expenses
incurred during the project.
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