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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 |
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Nos . |
91,95,104,108 |
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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 |
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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- 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 8 Vi 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
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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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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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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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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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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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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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
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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.
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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
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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.)
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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.
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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
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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.
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Contributions in Science
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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
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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- 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.
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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 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.
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David K. Caldwell 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.
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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
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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.
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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
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Late Miocene Mammals from Nevada
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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.
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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
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Late Miocene Mammals from Nevada
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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 |
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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.
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Late Miocene Mammals from Nevada
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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.
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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.)
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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
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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.
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Late Miocene Mammals from Nevada
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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.
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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.
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 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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1966
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.
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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.
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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.
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Contributions in Science
No. 93
Figure 23. Ophioderma panamense.
Figure 24. Ophiopsila californica.
Figure 25. Ophiopteris papillosa.
Figure 26. Ophiothrix spiculata.
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19
Figure 27 . Ophiothrix rudis.
Figure 28. Ophiopholis bakeri.
Figure 29. Ophiacantha phragma. Figure 30. Ophiacantha diplasia.
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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 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
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.
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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
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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.;
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Pliocene Birds from Chihuahua, Mexico
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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
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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
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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.
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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
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Pliocene Birds from Chihuahua, Mexico
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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.
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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
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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.
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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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Behavior of Wild and Captive Killer Whales
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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
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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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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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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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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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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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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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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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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
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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.
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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.
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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
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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.
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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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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.
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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.
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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
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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
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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
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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.
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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 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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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.
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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.
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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
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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.
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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
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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
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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)
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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.
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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
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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David K. Caldwell Editor
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.
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saiutsuiv
kM#hm
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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)
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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
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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.
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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
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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.
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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.
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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)
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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.
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$ . 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).
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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
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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.
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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.
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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,
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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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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
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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-
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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 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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David K. Caldwell 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
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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
o
G
U)
o3
p «
s g?
P cd , o
T3
cd
O
O T3 2 e
o g
x
<L> +->
<4_,
o g
£ -
g 13
P G
!Z c
o>
O G <u
CD g
jo cd
g <e
p _
+
-a
O
P
o
’a> _
> s
0) Cd
'O'-S
G
cd
CJ
+ I
+
Ph £ I O Q “TO m
+
+ +
I !
+ +
+ I + I
+ i
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
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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
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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.
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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-
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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.
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(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
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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.
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y o s e m i t e
1966
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9
y o Semite
16
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c?
yosemite
1966
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j a c i n t o
18
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vanduzeei
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21
calif ornicus
20
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22
23
cal if ornicus
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Cf
9
27
cali
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22
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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
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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
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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.
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New Genus of West American Diodora
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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)].
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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.
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New Genus of West American Diodora
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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-
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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
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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
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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David K. Caldwell Editor
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-
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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.
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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.
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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.
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“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).
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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.
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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 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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David K. Caldwell 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.
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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.
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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-
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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
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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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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
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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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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
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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
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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.
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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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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.
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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.
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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
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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
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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
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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.)
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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
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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.
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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.
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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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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.
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Literature Cited
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1965. The riddle of the ocean. Moscow: Young Guard, 175 p. (in Russian).
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1966. Observations on the behavior of wild and captive false killer whales, with notes on associated behavior of other genera of captive delphinids. Los Angeles County Mus., Cont. in Sci., 95 : 1-32.
Burt, William H., and Richard P. Grossenheider
1952. A field guide to the mammals. Boston: Houghton Mifflin Co., 200 p., 32 pis.
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1959. Rapport sur les petits Cetaces ouest-africains. Resultats des recherches entreprises sur ces animaux jusqu’au mois de mars 1959. Bulletin de 1’Institut Fran?ais d’Afrique Noire, 21, serie A, no. 4, pp. 1367-1409, pis. 1-31.
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1966. Behavior of the sperm whale, Physeter catodon L. In K. S. Norris, editor, Whales, dolphins and porpoises. Univ. Calif. Press, pp. 677-717 .
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1965. Marine mammals from the coast of Georgia to Cape Hatteras. J. Elisha Mitchell Scientific Soc., 81(1): 24-32.
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1965. Individualized whistle contours in bottlenosed dolphins ( Tursiops trun- catus ). Nature, 207(4995): 434-435.
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Caldwell, Melba C., David K. Caldwell, and William E. Evans
1966. Sounds and behavior of captive Amazon freshwater dolphins, Inia geoffrensis. Los Angeles County Mus., Cont. in Sci., 108.
Caldwell, Melba C., David K. Caldwell and I. B. Siebanaler
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.
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Elliot, Daniel G.
1901. A synopsis of the mammals of North America and the adjacent seas. Publ. Field Columbian Mus., Zool. Ser., 2: xv + 1-471.
1904. The land and sea mammals of Middle America and the West Indies. Field Columbian Mus., publ. 95, Zool. Ser., 4(1): xxi -f- 1-439 + xlix.
Flower, William H.
1874. On Risso’s dolphin, Grampus griseus (Cuv.). Trans. Zool. Soc. London, 8: 1-21, pis. 1-2.
Fraser, F. C.
1937. Whales and dolphins. In J. R. Norman and F. C. Fraser, Giant fishes, whales and dolphins. London: Putnam, pp. 201-349.
1950. Description of a dolphin, Stenella frontalis (Cuvier) from the coast of French Equatorial Africa. Atlantide Rept., 1 : 61-84, pis. 6-9.
Gunter, Gordon
1941. A record of the long-snouted dolphin, Stenella plagiodon (Cope), from the Texas coast. J. Mammal., 22(4) : 447-448.
1954. Mammals of the Gulf of Mexico. In P. S. Galtsoff, coordinator. Gulf of Mexico. Its origin, waters, and marine life. U.S. Fish and Wildlife Surv., Fish. Bull., 55(89) : 543-551.
Hall, E. Raymond, and Keith R. Kelson
1959. The mammals of North America. New York: The Ronald Press, vol. 2, viii + 547-1003 + 1-79.
Hoffmeister, Donald F.
1963. The golden bookshelf of natural history. Mammals. New York: Golden Press, 104 p.
Kay, Helen
1964. The secrets of the dolphin. New York: The Macmillan Co., 120 p.
Kellogg, Remington
1940. Whales, giants of the sea. Natl. Geogr. Mag., 67(1): 35-90.
Lauber, Patricia
1963. The friendly dolphins. New York: Random House, 81 p.
Lawrence, Barbara, and William E. Schevill
1956. The functionarl anatomy of the delphinid nose. Bull. Mus. Comp. Zool., Harvard, 114(4): 103-151, 30 figs.
Layne, James N.
1965. Observations on marine mammals in Florida waters. Bull. Florida State Mus., Biol. Sci., 9(4): 131-181.
Layne, James N., and David K. Caldwell
1964. Behavior of the Amazon dolphin, Inia geoffrensis (Blainville), in cap- tivity. Zoologica, 49(2): 81-108, pis. 1-3.
Lowery, George H., Jr.
1943. Check-list of the mammals of Louisiana and adjacent waters. Occ. Pap. Mus. Zool., Louisiana State Univ., 13: 213-257.
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Lyon, Marcus W., Jr., and Wilfred H. Osgood
1909. Catalogue of the type-specimens of mammals in the United States Na- tional Museum, including the Biological Survey Collection. Bull. U.S. Natl. Mus., 62: x + 1-325.
McBride, Arthur F.
1940. Meet mister porpoise. Nat. Hist., 45: 16-29.
McBride, Arthur F., and Henry Kritzler
1951. Observations on pregnancy, parturition, and post-natal behavior in the bottlenose dolphin. J. Mammal., 32(3) : 251-266.
Mahnken, Thomas, and Raymond M. Gilmore
1960. Suckerfish on porpoise. J. Mammal., 41 ( 1 ) : 134.
Millais, J. G.
1906. The mammals of Great Britain and Ireland. London: Longmans, Green and Co., 3: xii + 348 p.
Miller, Gerrit S., Jr.
1924. List of North American Recent mammals. Bull. U.S. Natl. Mus., 128: xvi + 1-673.
Miller, Gerrit S., Jr., and Remington Kellogg
1955. List of North American Recent mammals. Bull U. S. Natl. Mus., 205: xii -f 1-954.
Moore, Joseph C.
1953. Distribution of marine mammals to Florida waters. Amer. Midland Nat., 49(1): 117-158.
Nichols, John T.
1920. Red bat and spotted porpoise off the Carolinas. J. Mammal., 1 :87.
Nishiwaki, Masaharu, and M. Yabuuchi
1965. Whales and pinnipeds. Tokyo: Univ. Tokyo Press, 439 p. (in Japanese)
Palmer, Ralph S.
1954. The mammal guide. Garden City, New York: Doubleday and Co., Inc., 384 p., 40 pis.
Poole, Arthur J., and Viola S. Schantz
1942. Catalogue of the type specimens of mammals in the United States Na- tional Museum, including the Biological Surveys Collection. Bull. U.S. Natl. Mus., 178: xiii + 1-705.
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.
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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
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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
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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 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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David K. Caldwell Editor
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.
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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
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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.
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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
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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.
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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
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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
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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,
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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
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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
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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
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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.
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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.
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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.
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.
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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
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].
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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.
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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
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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).
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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.
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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 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
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.
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New Fossil Birds
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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
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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
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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.
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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-
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New Fossil Birds
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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.
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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 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
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
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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.
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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.
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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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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.
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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.
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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.
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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.
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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
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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.
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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.
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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.
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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
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“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-
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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
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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
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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
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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
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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
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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.
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Literature Cited
Allen, Ross, and Wilfred T. Neill
1957. White whales of the Amazon. Nat. Hist., 66(5) : 324-329.
Brown, David H., David K. Caldwell, and Melba C. Caldwell
1966. Observations on the behavior of wild and captive false killer whales, with notes on associated behavior of other genera of captive delphinids. Los Angeles County Mus., Cont. in Sci., 95 : 1-32.
Busnel, Rene-Guy and Albin Dziedzic
1966. Acoustic signals of the pilot whale Globicephala melaena and of the porpoises Delphinus delphis and Phocoena phocoena. In Kenneth S. Norris, editor, Whales, dolphins, and porpoises. Berkeley: Univ. Cali- fornia Press, pp. 607-646.
Busnel, Rene-Guy, Albin Dziedzic, and Soren Andersen
1963. Sur certaines caracteristiques des signaux acoustiques du Marsouin Phocoena phocoena L. Comptes Rendus des Academie des Sciences, Paris, 257: 2545-2548.
1965. Role de l’impedance d’une cible dans le seuil de sa detection par la sys- teme sonar du Marsouin P. phocaena. Comptes rendus dus seances de la Societe de Biologie, 159(1): 69-74.
Caldwell, Melba C., and David K. Caldwell
1964. Experimental studies on factors involved in care-giving behavior in three species of the cetacean family Delphinidae. Bull. So. Calif. Acad. Sci., 63(1): 1-20.
1965. Individualized whistle contours in bottlenosed dolphins ( Tursiops trun- catus). Nature, 207(4995) : 434-435.
Caldwell, Melba C., David K. Caldwell, and William E. Evans
In press. Preliminary results of studies on the sounds and associated behavior of captive Amazon freshwater dolphins, Inia geoffrensis. Proc. Third An- nual Conf. on Biological Sonar and Diving Mammals held at the Stan- ford Research Institute, California, on May 23-24, 1966.
Caldwell, Melba C., Ruth M. Haugen, and David K. Caldwell
1962. High-energy sound associated with fright in the dolphin. Science, 138 (3543): 907-908.
Curtis, Lawrence
1962. The Amazon dolphin, Inia geoffrensis, at the Fort Worth Zoological Park. Internatl. Zoo Yearbook, 4: 7-10.
Dempster, Robert P.
1965. Shipping a live Amazon dolphin. All-Pets Mag., 36(3) : 13-15,17.
Dreher, John J., and William E. Evans
1964. Cetacean communication. In William N. Tavolga, editor, Marine bio- acoustics. New York: Pergamon Press, pp. 373-393.
Evans, William E., and John 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.
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Evans, William E., and John H. Prescott
1962. Observations of the sound production capabilities of the bottlenose por- poise: a study of whistles and clicks. Zoologica, 47(3) : 121-128, pis. 1-4.
Fish, Marie P., and W. H. Mowbray
1962. Production of underwater sound by the white whale or beluga, Delphin- apterus leucas (Pallas). Sears Found.: J. Mar. Res., 20(2) : 149-162.
Herald, Earl S., and Robert P. Dempster
1965. Meet “Whiskers!’ Aquarium J., 36(5) : 213-215, 236-237.
Hill, Gary T.
1965. The Amazonian dolphin. Skin Diver Mag., 14(4) : 37.
Hofmeister, Max
1964. Our pink porpoise. Current number 1 attraction at the Toledo Zoo. Sa- fari (Toledo Zoo), 16(1): 6-7.
Kellogg, Winthrop N.
1961. Porpoises and sonar. Chicago: Univ. Chicago Press, xiv + 177 p.
Layne, James N.
1958. Observations on freshwater dolphins in the upper Amazon. J. Mammal., 39(1): 1-22.
1959. Feeding adaptations and behavior of a freshwater dolphin, Inia geof- frensis. Anat. Rec., 134(3) : 598. (abstract)
Layne, James N., and David K. Caldwell
1964. Behavior of the Amazon dolphin, Inia geoffrensis (Blainville), in cap- tivity. Zoologica, 49(2): 8-108, pis. 1-4.
Lilly, John C.
1962. Vocal behavior of the bottlenose dolphin. Proc. Amer. Philos. Soc., 106(6): 520-529.
McBride, Arthur F., and D. O. Hebb
1948. Behavior of the captive bottlenose dolphin, Tursiops truncatus. J. Comp. Physiol., 41(2): 111-123.
Morris, Robert A., and Louis S. Mowbray
1966. An unusual barnacle attachment on the teeth of the Hawaiian spinning dolphin. Norsk Hvalfangst-Tidende (The Norwegian Whaling Gazette), 55(1): 15-16.
Norris, Kenneth S.
1964. Some problems of echolocation in cetaceans. In William N. Tavolga, editor. Marine bio-acoustics. New York: Pergamon Press, pp. 317-336.
Norris, Kenneth S., and William E. Evans
1966. Directionality of echolocation clicks in the rough-toothed porpoise, Steno bredanensis (Lesson). Paper presented at a Symposium on Ma- rine Bio-Acoustics held at the American Museum of Natural History, New York, on April 13-15, 1966. (To be published)
Norris, Kenneth S., John H. Prescott, Paul V. Asa-Dorian, and Paul Perkins
1961. An experimental demonstration of echo-location behavior in the por- poise, Tursiops truncatus (Montagu). Biol. Bull., 120(2) : 163-176.
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Phillips, Craig
1964. The captive sea. Life behind the scenes of the greast modern oceanari- ums. Philadelphia: Chilton Books, xii -f 284 p.
Phillips, James D., Jr., and Garvin McCain
1964. Black-white visual discrimination in the Amazon porpoise: Inia geof- frensis. Amer. Psychologist, 19(7): 503. (abstract)
Powell, Bill A.
In press. Periodicity of vocal activity of captive Atlantic bottlenosed dolphins: Tursiops truncatus. Bull. So. Calif. Acad. Sci., 65(4).
Richardson, Hal
1965. (No title, in section entitled “Academically Speaking!’) Pacific Disc., 18(2): 33.
Schevill, William E.
1964. Underwater sounds of cetaceans. In William N. Tavolga, editor, Ma- rine bio-acoustics. New York: Pergamon Press, pp. 307-316.
Schevill, William E., and William A. Watkins
1962. Whale and porpoise voices. A phonograph record. Woods Hole, Massa- chusetts: Woods Hole Oceanographic Institution, 24-page booklet and phonograph record.
Simpson, George G.
1945. The principles of classification and a classification of mammals. Bull. Amer. Mus. Nat. Hist., 85: i-xvi -f- 1-350.
Walker, Ernest P.
1964. Mammals of the world. Baltimore: The Johns Hopkins Press, vol. 2, pp. i-viii -f 647-1500.
Walls, G. L.
1963. The vertebrate eye and its adaptive radiation. New York: Hafner Publ. Co., 785 p.
Watkins, William A.
1966. Techniques and pitfalls in sound analysis. Paper presented at a Sympo- sium on Marine Bio-Acoustics held at the American Museum of Natural History, New York, on April 13-15, 1966. (To be published)
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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 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 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.
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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 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.
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No. 109
Figure 1. Haliotis corrugata oweni, new subspecies, holotype (LACM). Long. 146, lat. 125, alt. 55 mm.
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Figure 2. Haliotis corrugata oweni, new subspecies, paratype (LACM). Long. 105, lat. 84, alt. 42 mm.
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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.
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COUNTY MUSEUM |
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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- 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
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
No. 110
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
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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
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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
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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.
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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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Contributions in Science
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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
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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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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.
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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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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-
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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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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.
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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
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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).
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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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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.
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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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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.
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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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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
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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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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
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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
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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$, Nogales, June 1, 1903 (Oslar; CU); 1 9, Picacho, Pinal Co., June 18, 1961 (F. D. Parker; UCD). SONORA: 1 9 , San Bernardo,
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Rio Mayo, July 6, 1935 (H. S. Gentry; LACM); 2 2$, 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.
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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-
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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).
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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.
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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
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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-
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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.
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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.
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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.
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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 $ !’
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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 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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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
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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
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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
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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.
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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-
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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.
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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.
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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-
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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.
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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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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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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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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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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
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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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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
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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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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-
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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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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.
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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-
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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.
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1959. Some Pleistocene amphibians and reptiles from Friesenhahn Cave, Texas. Southwestern Nat. 3: 17-27, 3 text-figs.
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1962. Generic classification of the high-altitude pelobatid toads of Asia ( Scutiger , Aelurophryne, and Oreolax). Copeia, 1962(2) :287-291.
Nevo, Abiatha
1956. Fossil frogs from a Lower Cretaceous bed in southern Israel (Central Negev). Nature (London), 178(4543) : 1 191-1 192, 2 text-figs.
Noble, G. K.
1924. A new spadefoot toad from the Oligocene of Mongolia with a summary of the evolution of the Pelobatidae. Amer. Mus. Nov., 132:1-15, 7 text- figs.
Pasteur, Georges
1958. Sur les tendances evolutives et la phylogenie des Pelobcites (Batriciens Anoures) actuels. Comptes Rendus Seances PAcademie Sciences, Paris, 247(14) : 1037-1039.
Ramaswami, L. S.
1935. The cranial morphology of some examples of Pelobatidae (Anura). Anatomischer Anzieger, 81(4/6) : 65-96, 14 text-figs.
26
Contributions in Science
No. 113
Reig, O. A.
1958. Proposiciones para una neuva macrosistematica de los anuros (Nota preliminar). Physis, 21(60) : 109-1 18.
Ritland, Richard M.
1955. Studies on the post-cranial morphology of Ascaphus truei. I. Skeleton and spinal nerves. J. Morph., 97 ( 1) : 1 19-178, 1 1 text-figs., 1 pi., 2 tables.
Simpson, George G.
1961. Principles of animal taxonomy. New York: Columbia Univ. Press, xii-f 1-247, 30 text-figs.
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1954. A Kansas Pleistocene herpetofauna. Copeia, 1 954 ( 3 ) : 2 17-22 1 .
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Wasserman, Aaron O.
1957. Factors affecting interbreeding in sympatric species of spadefoots (genus Scaphiopus) . Evolution, 11(3) : 320-338, 7 text-figs., 5 tables.
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1955. Die Fauna der miozanen Spaltenfiillung von Neudorf a.d. March (CSR), Amphibia (Anura) et Reptilia. Sitzungsberichten der Oster- reichische Akademie der Wissenschaften, Mathematischnaturwissen- schaftliche Klasse, 1(164)10:801-815, 2 pis.
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1956. Two pelobatid frogs from the Tertiary of North America and their rela- tionships to fossil and Recent forms. Amer. Mus. Nov., 1762:1-45, 25 text-figs., 1 table.
|
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.
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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. |
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|
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 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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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
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
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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
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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.
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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.
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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.
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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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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
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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
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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
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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.
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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.
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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
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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).
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1898. Die Leuchtorganeder Tiefsee-Fische Argyropelecus und Chauliodus. Zeitschrift fur Naturwissenschaften, 71:447-452.
Catesby, M.
1771. Natural history of Carolina, Florida, and the Bahama Islands. 2: ii + 120 + vi pp., 120 pis. London: Benjamin White, (not seen).
Danilchenko, P.
1960. Bony fishes of Maikopski deposits in the Caucasus. Works of Paleont. Inst., Acad. Sciences U.S.S.R., Moscow, 78:1-203. (in Russian).
David, L. R.
1943. Miocene fishes of Southern California. Geol. Soc. Amer., Spec. Paper 43, 193 p., pis. 1-16.
Durham, J. W.
1960. Palaeoclimates in “Physics and Chemistry of the Earth!’ 3:1-15. Lon- don: Pergamon Press.
Ege, V.
1948. Chauliodus Schn., a bathypelagic genus of fishes. Dana-Rept., 31 : 1-148.
Emery, K. O.
1960. The sea off Southern California. New York: John Wiley and Sons., Inc., 364 p.
Emiliani, C.
1954. Temperatures of Pacific bottom waters and polar superficial waters during the Tertiary. Science, 119:853-55.
Garman, S.
1899. Reports on an exploration off the west coast of Mexico, Central and South America, and off the Galapagos Islands. Mem. Mus. Comp. Zool., 24:431. (not seen).
28
Contributions in Science
No. 115
Gosline, W. A.
1960. Contributions toward a classification of modern isospondylous fishes. Bull. Brit. Mus. (Nat. Hist.)., 6(6) :327-365.
Greenwood, P. H., Donn E. Rosen, Stanley H. Weitzman, and George S. Myers
1966. Phyletic studies of teleostean fishes, with a provisional classification of living forms. Bull. Amer. Mus. Nat. Hist., 131(4) : 339-456, pis. 21-23.
Haffner, R.
1952. The zoogeography, biology and systematics of the Chauliodontidae. Unpublished Ph.D. Dissertation, Yale Univ., 114 p.
Hollister, G.
1934. Clearing and dying fish for bone study. Zoologica, 12:89-101.
Hoots, H. W.
1931. Geology of the eastern part of the Santa Monica Mountains, Los Angeles County, California. U.S. Geol. Surv., Prof. Paper 165-C: 83-134.
Jerzmanska, A.
1960. Ichtiofauna lupkow jasielskich z Sobniowa. (Ichthyofauna from the Jaslo shales of Sobniow) Acta Paleontologica Polonica, 5(4) :367-432.
Jordan, D. S.
1925. The fossil fishes of the Miocene of Southern California. Stanford Univ. Publ., Biol. Sciences, 4(1): 1-51.
Kummel, B.
1961. History of the earth. San Francisco: Freeman Co., 610 p.
Ladd, H. S.
1959. Ecology, paleontology, and stratigraphy. Science, 129:69-78.
Lagler, K. E., J. E. Bardach, and R. R. Miller
1962. Ichthyology, the study of fishes. New York: John Wiley and Sons, Inc., 545 p.
McCullouch, A. R.
1916. Report on some fishes obtained by the F.I.S. “Endeavor” on the coasts of Queensland, New South Wales, Victoria, Tasmania, South and Southeastern Australia. Dept. Trade and Customs, Fisheries, 4(4). (not seen).
Morrow, J. E., Jr.
1961. Taxonomy of the deep sea fishes of the genus Chauliodus. Bull. Mus. Comp. Zook, 125(9): 249-94.
1964. Family Chauliodontidae. In Fishes of the western North Atlantic. Mem. Sears Found. Mar. Res., 1(4) : 274-289.
Regan, C. T., and Ethelwyn Trewaves
1929. The fishes of the families Astronesthidae and Chauliodontidae. Dana- Rept., 5:1-39.
1930. The fishes of the families Stomiatidae and Malacosteidae. Dana-Rept., 6:1-143, pis. 1-13.
1966
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Sullwold, H. H., Jr.
1960. Tarzana fan, deep submarine fan of late Miocene age, Los Angeles County, California. Bull. Amer. Assoc. Petrol. Geol., 44(4) : 433-457.
Tchernavin, V. V.
1953. Summary of the feeding mechanism of a deep sea fish, Chauliodus sloani. Brit. Mus. (Nat. Hist.), London, 101 p., pis. 1-10.
Whitmore, F. C., Jr., and Robert H. Stewart
1965. Miocene mammals and Central American seaways. Science, 148: 180-185.
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
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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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I
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
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.
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Figure 1. Neadmete sutherlandi, new species. Holotype, LACM 1150. Long. 39.6, lat. 16.1 mm.
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Figure 2. Neadmete sutherlandi, new species. Holotype, LACM 1150. Long. 39.6, lat. 16.1 mm.
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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
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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.
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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.
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LOS ANGELES 1; COUNTY MUSEUM |
CONTRIBUTIONS IN SCIENCE |
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IJMBER 117 |
December 28, 1966 |
A NEW SPECIES OF ARCHITECTONICA FROM THE SANTA SUSANA MOUNTAINS, VENTURA COUNTY, CALIFORNIA
By J. Alden Sutherland
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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.
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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.-.
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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.
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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.
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LOS ANGELES |
CONTRIBUTIONS |
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COUNTY MUSEUM |
IN SCIENCE |
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Timber 118 |
December 28, 1966 |
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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.
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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
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Figure 3. Abedus parked Menke, new species, holotype. A, Dorsal view; B, Ventral view.
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Figure 4. Abedus breviceps Stal. A, Dorsal view; B, Ventral view.
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New Mexican Toe Biter
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Figure 5. Abedus signor eti sonorensis Menke. A, Dorsal view; B, Ventral view.
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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
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ADDITIONAL FISH REMAINS, MOSTLY OTOLITHS, FROM A PLEISTOCENE DEPOSIT AT PLAYA DEL REY, CALIFORNIA
I
By John E. Fitch
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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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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.
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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
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Pleistocene Fish Remains
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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
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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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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.
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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
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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.
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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.
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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-
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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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