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CONTRIBUTIONS
IN SCIENCE
DECEMBER 3 1, 1970
5 ? S
O'? Lies
TABLE OF CONTENTS
and
AUTHOR INDEX
1970
Nos. 176-205
LOS
M I
ANGELES
COUNTY
MUSEUM
FE8 1 81971
4'3RAR1£5.
Los Angeles County Museum of Natural History • Exposition Park
Los Angeles, California 90007
2
CONTRIBUTIONS IN SCIENCE
TABLE OF CONTENTS
No. 176.
No. 177.
No. 178.
No. 179.
No. 180.
No. 181.
No. 182.
No. 183.
No. 184.
No. 185.
No. 186.
No. 187.
No. 188.
No. 189.
The status and habits of Grauer's broadbill in Uganda (Aves:
Eurylaemidae), by Herbert Friedmann. 4 pp., January 30, 1970.
A new species of Neadmete (Neogastropoda) from the Pliocene of
California, by Jack D. Mount. 4 pp., 1 fig. February 25, 1970.
Description of a new species of net -winged midge from the Great
Basin, with a key to the North American species of the genus
Dioptopsis (Diptera: Blephariceridae), by Charles L. Hogue, 10 pp. ,
13 figs. February 25, 1970.
Geographic variation and systematic status of Mexican lyre snakes of
the Trimorphodon tau group (Colubridae), by Roy W. McDiarmid and
Norman J. Scott, Jr. 43 pp. , 7 figs. February 25, 1970.
Studies on North American bees of the genus Hylaeus. 5. The
subgenera Hylaeus, S. Str. and Paraprosopis (Hymenoptera: Colletidae),
by Roy R. Snelling. 59 pp. , 12 figs, February 27, 1970.
Type specimens of fossil invertebrata in the Los Angeles County
Museum of Natural History, exclusive of Paleoentomology, by
Edward C. Wilson and Donald E. Bing. 20 pp, February 27, 1970.
Aquatic adaptations of the water mouse, Rheomys underwoodi, by
Andrew Starrett and George F. Fisler. 14 pp, April 17, 1970,
Observations and records of Myotis (Pizonyx) vivesi Menegaux
(Chiroptera: Vespertilionidae), by Donald R. Patten and Lloyd T,
Findley, 9 pp. April 17, 1970.
Conocardium langenheimi Sp. N. (Mollusca: Bivalvia) in the Lower
Permian Series of the McCloud limestone, northern California, by
Edward C. Wilson. 14 pp, , 34 figs, April 17, 1970,
Five new epitoniid gastropods from the west coast of the Americas, by
Helen DuShane. 6 pp. , 5 ,figs. April 17, 1970.
A generic revision and catalog of the western hemisphere Glenurini
with the description of a new genus and species from Brazil (Neuroptera:
Myrmeleontidae), by L, A. Stange. 28 pp, , 8 figs. April 17, 1970.
Further studies on audible vocalizations of the Amazon freshwater
dolphin, Inia geoffrensis, by Melba C. Caldwell and David K.
Caldwell. 5 pp. , 2 figs. May 4, 1970.
Four new terebrid gastropods from the eastern Pacific jslands, by
Twila Bratcher and R. D. Burch. 6 pp. , 8 figs. May 4, 1970.
Reinstatement of the turrid genus Bellaspira Conrad, 1868
(Mollusca: Gastropoda) with a review of the known species, by
James H. McLean and Leroy H. PoOTman. 11 pp. , 16 figs.
May 4, 1970,
TABLE OF CONTENTS
3
No. 190.
No. 191.
No. 192.
No. 193.
No. 194.
No. 195.
No. 196.
No. 197.
No. 198.
No. 199.
No. 200.
No. 201.
No. 202.
No. 203.
No. 204.
No. 205.
A new species of fossil Kelletia (Molliisca: Gastropoda) from the
Lomita Marl, Late Cenozoic of San Pedro, California, by Leo G,
Hertlein. 8 pp. , 3 figs. May 20, 1970.
Studies on the frogs of the genus Leptodactylus (Amphibia:
Leptodactylidae). VI. Biosystem atics of the Melanonotus group,
by W. Ronald Heyer. 48 pp. , 11 figs. May 20, 1970.
System atics of the lizards of the gekkonid genus Phy llod actylus of
mainland South America, by James R, Dixon and Raymond B. Huey.
78 pp. , 14 figs. June 26, 1970.
Hadrosaurian dinosaur bills —morphology and function, by William J.
Morris. 14 pp. , 5 figs. July 1, 1970.
Another new night lizard (Xantusia) from Durango, Mexico, by Robert
G, Webb. 10 pp. , 2 figs. July 1, 1970.
The birds of the Kalinzu Forest, southwestern Ankole, Uganda, by
Herbert Friedmann and John G. Williams. 27 pp, July 1, 1970.
Two new species of catfishes of the genera Nannorhamdia and
Imparales (family Pimelodidae) from Central America, by William A,
Bussing. 11 pp. , 2 figs, July 1, 1970.
The social wasps of Lower California, Mexico (Hymenoptera: Vespidae),
by Roy R. Snelling. 20 pp, , 4 figs. July 28, 1970,
Additions to the known avifauna of the Bugoma, Kibale, and
Impenetrable Forests, west Uganda, by Herbert Friedmann and John G.
Williams, 20 pp, August 31, 1970.
Fish remains, mostly otoliths and teeth, from the Palos Verdes Sand
(Late Pleistocene) of California, by John E. Fitch, 41 pp, , 6 figs.
November 13, 1970,
A new species of Batrachoseps (slender salamander) from the desert of
southern California, by Arden H. Brame, Jr. 11pp., 5 figs.
November 13, 1970,
A new salamander (genus Oedipina) of the uniformis group from western
Panama, by Arden H. Brame, Jr. and William E. Duellman, 8pp,,
3 figs. November 13, 1970.
The ethology of the wasp, Pseudomasaris edwardsii (Cresson), and a
description of its immature forms (Hymenoptera: Vespoidea, Masaridae),
by Philip F. Torchio, 32 pp., 33 figs, November 24, 1970.
A review of the extinct avian genus, Mancalla, by Hildegarde Howard.
12 pp,, 1 fig. November 24, 1970,
A new moth fly of the genus Psychoda from crabholes on the Kenya
coast (Diptera: Psychodidae), by Charles L. Hogue, 5 pp, , 7 figs,
November 24, 1970,
Further information on the breeding biology of the honey guides, by
Herbert Friedmann, 5 pp, November 24, 1970,
4
CONTRIBUTIONS IN SCIENCE
Bing, Donald E.
Brame, Arden H, , Jr
Bratcher, Twila
Burch, R, D.
Bussing, William A.
Caldwell, David K.
Caldwell, Melba C.
Dixon, James R.
Duellman, William ]
DuShane, Helen
Findley, Lloyd T,
Fisler, George F.
Fitch, John E.
Friedmann, Herbert
Hertlein, Leo G.
Heyer, W. Ronald
Hogue, Charles L.
Howard, Hildegarde
Huey, Raymond B.
McDiarmid, Roy W.
McLean, James H.
Morris, William J,
Mount, Jack D.
Patten, Donald R,
Poorman, Leroy H.
Scott, Norman J., Jr
Snelling, Roy R.
Stange, L. A,
Starrett, Andrew
Torchio, Philip F.
Webb, Robert G,
Williams, John G.
Wilson, Edward C.
AUTHOR INDEX
No. 181
Nos. 200, 201
No. 188
No. 188
No. 196
No. 187
No. 187
No. 192
No. 201
No. 185
No. 183
No. 182
No. 199
Nos. 176, 195, 198, 205
No. 190
No. 191
Nos. 178, 203
No. 203
No. 192
No. 179
No. 189
No. 193
No. 177
No. 183
No. 189
No. 179
Nos. 180, 197
No. 186
No. 182
No. 202
No. 194
Nos. 195, 198
Nos. 181, 184
LOS
ANGELES
COUNTY
MUSEUM
CONTRIBUTIONS
IN SCIENCE
January 30, 1970
1. 7 J
THE STATUS AND HABITS OF GRAUER’S BROADBILL
IN UGANDA (Aves: Eurylaemidae)
By Herbert Friedmann
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 sep-
arately, and numbers run consecutively regardless of subject matter. Number 1 was
issued January 23, 1957. The series is available to scientific institutions and scien-
tists on an exchange basis. Copies may also be purchased at a nominal price. Inquiries
should be directed to Virginia D. Miller, Los Angeles County Museum of Natural
History, 900 Exposition Boulevard, Los Angeles, California 90007.
Robert J. Lavenberg
Managing Editor
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. Al-
though priority will be given to manuscripts by staff members, or to papers dealing
largely with specimens in the Museum’s collections, other technical papers will be
considered. All manuscripts must be recommended for consideration by the curator
in charge of the proper section or by the editorial board. Manuscripts must conform
to those specifications listed below and will be examined for suitability by an Edi-
torial Committee. They may also be subject to review by competent specialists out-
side the Museum.
Authors proposing new taxa in a CONTRIBUTIONS IN SCIENCE must
indicate that the primary type has become the property of a scientific institution of
their choice and cited by name.
MANUSCRIPT FORM.— (1) The 1964 AIBS Style Manual for Biological
Journals is to be followed in preparation of copy. (2) Double space entire manu-
script. (3) Footnotes should be avoided if possible. Acknowledgments as footnotes
will not be accepted. (4) Place all tables on separate pages. (5) Figure legends and
unavoidable footnotes must be typed on separate sheets. Several of one kind may
be placed on a sheet. (6) An abstract must be included for all papers. This will be
published at the head of each paper. (7) A Spanish summary is required for all
manuscripts dealing with Latin American subjects. Summaries in other languages
are not required but are strongly recommended. Summaries will be published at the
end of the paper. (8) A diagnosis must accompany any newly proposed taxon.
ILLUSTRATIONS. — All illustrations, including maps and photographs, will
be referred to as figures. All illustrations should be of sufficient clarity and in the
proper proportions for reduction to CONTRIBUTIONS page size. Consult the 1964
AIBS Style Manual for Biological Journals in preparing illustration and legend copy
for style. Submit only illustrations made with permanent ink and glossy photo-
graphic prints of good contrast. Original illustrations and art work will be returned
after the manuscript has been published.
PROOF. — Authors will be sent galley proof which should be corrected and
returned promptly. Alterations or changes in the manuscript after galley proof will
be billed to the author. Unless specifically requested, page proof will not be sent to
the author. One hundred copies of each paper will be given free to each author or
divided equally among multiple authors. Orders for additional copies must be sent
to the Editor at the time corrected galley proof is returned. Appropriate order forms
will be included with the galley proof.
Virginia D. Miller
Editor
THE STATUS AND HABITS OF GRAUER’S BROADBILL
IN UGANDA (Aves: Eurylaemidae)
By Herbert Friedmann^
Abstract: Five examples of the very rare broadbill, Pseudo-
calyptomena graueri, from the Impenetrable Forest, southwestern
Uganda, prove to be the same as topotypical east Congo birds,
and not to have consistently narrower bills as was earlier sus-
pected. The field notes accompanying the specimens cause a
complete revision of our knowledge of this bird. It is primarily
a denizen of the forest undergrowth, not of the tree-tops, and it
feeds largely on seeds, buds and flowers, plus such insect larvae
and beetles that may occur with them, and is not primarily an
aerial feeder on flying insects as has been assumed in the litera-
ture.
In the course of a second collecting survey of the vertebrate fauna of the
Impenetrable Forest, Kigezi, extreme southwestern Uganda, under the spon-
sorship of National Science Foundation grant GB 7787, to the Los Angeles
County Museum of Natural History Foundation, Robert Glen and Andrew
Williams obtained five specimens of Grauer’s green broadbill, Pseudocalyp-
tomena graueri, in May and June, 1969. These were particularly significant
because two years earlier A. L. Archer had collected one male in that area,
which had a very narrow, laterally compressed bill, only 8.5 mm broad at the
gape, and 5.2 mm wide at the proximal (basal) end of the nares, compared
with 10.6 to 1 1.5 mm at the gape and 7.3 to 7.6 mm at the basal edge of the
nostrils in eastern Congo examples. In discussing this specimen Friedmann
and Williams (1968: 22) stated that additional specimens would be of inter-
est to determine whether or not the Impenetrable Forest birds might prove to
be a racially distinct narrow billed population. The present five examples show
that this is not the case, that the first one was merely unusual in the narrowness
of its bill. The measurements in millimeters of the five 1969 examples, now in
The Los Angeles County Museum of Natural History, are as follows: 2 males—
width of bill at gape 13.2, 13.6; width of bill at basal edge of nostril 6.9, 7.1;
3 females— width of bill at gape 13.4, to 13.6; width of bill at basal edge of
nostrils 7 to 7.5. As may be noted from these figures, kindly measured for me
by J. R. Northern, the present series have slightly broader, not narrower, bills
than the eastern Congo topotypical examples measured earlier. No Congo
material was studied in connection with these five specimens, but it is obvious
that the variational range in bill width in the Kigezi birds completely overlaps
that of topotypical graueri, and that the two populations are not separable.
^Director, Los Angeles County Museum of Natural History, Los Angeles, Cali-
fornia 90007
1
Contributions in Science
No. 180
9
That the two are not different is actually easier to understand than was
the impression of a local difference suggested by Archer’s single specimen in
1967. The extreme limits of the entire, and relatively narrow, area of high-
land forest known to be inhabited by this broadbill are barely more than 150
miles apart, an area that seems to be too small and ecologically too continu-
ously uniform to allow for geographic differentiation. It is true that all the
eastern Congo records are from the mountains to the west of the deep gorge
of the Ruzizi river, but this valley does not constitute a barrier between those
highlands and the mountainous forests of Kigezi. The eastern Congolese high-
lands extend north from the northern end of the Ruzizi river along the west
shore of Lake Kivu to the Rwanda border and to that of extreme southwestern
Uganda, where they are coterminous with the so-called Impenetrable Forest.
Furthermore there is no reason for thinking that the broadbill is either very
local or sporadic within this small total area, even though the actual specimen
locality records are still few in number. Definite records from the Kivu dis-
trict, eastern Congo, are the type locality about 80 kilometers west of the
Ruzizi river, northwest of Lake Tanganyika; Miki, Muusi and Luvumba. In
southwestern Uganda, the bird is known only from the Bwindi and Ruhizha
areas of the Impenetrable Forest, Kigezi Province.
Knowledge of Grauer’s broadbill as a bird, and not merely as a museum
taxon, is drastically altered by the field observations by Glen and Williams
that accompanied the specimens. Archer, in his field notes on the first (1967)
Kigezi specimen, wrote that he shot the bird from a perch about 60 feet up in
a tall tree. Rockefeller and Murphy ( 1933: 28) similarly found this species
moving about in the upper branches of fairly tall trees. Chapin (1953: 23)
noted that the green broadbill was usually found 25 to 75 feet up in forest
trees near native plantations, and that in their behavior the birds were quite
flycatcher-like, making short but rapid dashes after insects, which they appar-
ently caught on the wing.
Chapin’s statements were based entirely on the notes of Rockefeller and
Murphy as at that time he had had no personal experience with the bird in life.
Therefore the observations of Rockefeller and Murphy are taken directly
from their paper. The first individual they saw was in a “. . . large tree with
abundant juicy berries, a feeding place for many other species of birds”; it was
not active and seemed not to be feeding. “When shot, it was catching insects
about twenty feet above the ground in a vine-draped forest tree . . .” Another
one was seen later “. . . slumped down close to the end of one of the uppermost
boughs, then with a hop and a flit of the wings it caught an insect over its
head, realighting on the perch it had just left . . .” These two observers wrote
that the “normal level” for Grauer’s broadbill was from 20 to 75 feet up in
trees.
The picture one gets from the reports of Glen and Williams is quite dif-
ferent. They found the species to be a denizen, not of the higher branches.
1970
The Status and Habits of Grauer’s Broadbill
3
but of the upper portions of the undergrowth, about eight feet from the
ground. When an individual of the species was seen high up in a tree it had
just flown there from the nearby lower vegetation. Glen considered its general
behavior and actions very much like those of a waxbill rather than a flycatcher.
The species looks extremely small in the field, and the short tail gives the bird
a crombec-like (Sylvietta) appearance in flight, especially when seen from
above. The birds were found, not always singly, but at times in loose groups of
2 or 3 individuals, searching about for food among the branches of a euphor-
biaceous tree, Neoboutonia sp., with apparent lack of concern about the ap-
proach of the observers.
The stomach contents of the five present specimens reveal not the slight-
est indication of any aerial feeding habits. The stomach of one of them con-
tained a single small beetle, some small seeds, and small flower buds; the sec-
ond contained only flower buds; the third had the remains of small pulpy fruits;
the fourth only small white flowers; and the fifth contained one small snail, one
beetle, several insect larvae and a white seed.
These stomach contents are in general agreement with the fact that
Rockefeller and Murphy saw one of these broadbills in a berry-laden tree,
even though they witnessed no feeding. In their paper they failed to report on
the contents of the stomachs of any of their specimens.
Inasmuch as the only other African broadbill genus, Smithornis, is known
to indulge in a specialized courtship flight behavior, one wonders if the state-
ment referred to above, of Pseudocalyptomena making short rapid flights
might relate to a similar courtship behavior and not merely to a feeding
pattern.
On one occasion Glen and Williams watched this species making short
flights through the upper undergrowth of the forest and noted that the flight
was of a slow and gliding nature. Even on short flights the birds rarely flapped
their wings after starting but seemed to glide by preference. In Smithornis
rufolateralis Chapin {loc. cit: 18) was convinced that the noises accompanying
the flights was produced in the wings. He noted that the primaries had unusually
stiff shafts and that the plane of their webs was twisted slightly on their distal
portions. “. . . four or five of the outer primaries (but not the outermost) have
unusually narrow webs toward the base, and it seems possible that air may
pass between them during this special flight . . .” He noted {loc. cit: 22-23)
that in Pseudocalyptomena the same remiges were fairly similar, but with
“. . . just a little of the downward twist of the inner web toward the tip . . .”
Whether this slight difference could be responsible for the absence of the
flight sound in the green broadbill is not at all clear, but as our collectors
made no entry in their notebooks of any such sound, it would seem that these
flights were unaccompanied by any special noises. The collectors did report
two types of call notes from Brauer’s broadbill: on one occasion they heard
one of these birds give a soft cree-cree repeated three times; and at another
4
Contributions in Science
No. 180
time they heard a one-syllabled high-pitched prrrp at intervals of about thirty
seconds.
The new field notes are also of interest in that they tend to allocate
Pseudocalyptornena ecologically more closely with Smithornis. The latter
genus is known to nest and to feed fairly low down in the forest vegetation,
the nests so far reported having been close to the ground or not more than
eight feet from it. The earlier observations that conveyed the impression that
the green broadbill was entirely a tree-top bird and an aerial feeder on insects
must be emended in the light of new knowledge.
Literature Cited
Chapin, J. P. 1953. The birds of the Belgian Congo. III. Amer. Mus. Nat. Hist. Bull.
75A: 1-821.
Friedmann, H., and J. G. Williams. 1968. Notable records of rare or little known
birds from western Uganda. Rev. Zool. Bot. Afr., 77, no. 1-2: 1 1-36.
Rockefeller, J. S., and C. B. G. Murphy. 1933. The rediscovery of Pseudocalyp-
tomena. Auk, 50: 23-29.
Accepted for publication December 16, 1969
LOS
ANGELES
COUNTY
MUSEUM
!
UMBER 177
CONTRIBUTIONS
iSf. IN SCIENCE
February 25, 1970
A NEW SPECIES OF NEADMETE (NEOGASTROPODA)
FROM THE PLIOCENE OF CALIFORNIA
By Jack D. Mount
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 Eos Angeles County Museum of Natural History. Issues are numbered sep-
arately, and numbers run consecutively regardless of subject matter. Number 1 was
issued January 23, 1957. The series is available to scientific institutions and scien-
tists on an exchange basis. Copies may also be purchased at a nominal price. Inquiries
should be directed to Virginia D. Miller, Los Angeles County Museum of Natural
History, 900 Exposition Boulevard, Los Angeles, California 90007.
Robert J. Lavenberg
Managing Editor
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. Al-
though priority will be given to manuscripts by staff members, or to papers dealing
largely with specimens in the Museum’s collections, other technical papers will be
considered. All manuscripts must be recommended for consideration by the curator
in charge of the proper section or by the editorial board. Manuscripts must conform
to those specifications listed below and will be examined for suitability by an Edi-
torial Committee. They may also be subject to review by competent specialists out-
side the Museum.
Authors proposing new taxa in a CONTRIBUTIONS IN SCIENCE must
indicate that the primary type has become the property of a scientific institution of
their choice and cited by name.
MANUSCRIPT FORM.— (1) The 1964 AIBS Style Manual for Biological
Journals is to be followed in preparation of copy. (2) Double space entire manu-
script. (3) Footnotes should be avoided if possible. Acknowledgments as footnotes
will not be accepted. (4) Place all tables on separate pages. (5) Figure legends and
unavoidable footnotes must be typed on separate sheets. Several of one kind may
be placed on a sheet. (6) An abstract must be included for all papers. This will be
published at the head of each paper. (7) A Spanish summary is required for all
manuscripts dealing with Latin American subjects. Summaries in other languages
are not required but are strongly recommended. Summaries will be published at the
end of the paper. (8) A diagnosis must accompany any newly proposed taxon.
ILLUSTRATIONS. — All illustrations, including maps and photographs, will
be referred to as figures. All illustrations should be of sufficient clarity and in the
proper proportions for reduction to CONTRIBUTIONS page size. Consult the 1964
AIBS Style Manual for Biological Journals in preparing illustration and legend copy
for style. Submit only illustrations made with permanent ink and glossy photo-
graphic prints of good contrast. Original illustrations and art work will be returned
after the manuscript has been published.
PROOF. — Authors will be sent galley proof which should be corrected and
returned promptly. Alterations or changes in the manuscript after galley proof will
be billed to the author. Unless specifically requested, page proof will not be sent to
the author. One hundred copies of each paper will be given free to each author or
divided equally among multiple authors. Orders for additional copies must be sent
to the Editor at the time corrected galley proof is returned. Appropriate order forms
will be included with the galley proof.
Virginia D. Miller
Editor
A NEW SPECIES OF NEADMETE (NEOGASTROPODA)
FROM THE PLIOCENE OF CALIFORNIA
By Jack D. Mount^
Abstract: Neadniete schwartzi, a new species of gastropod
belonging to the family Cancellariidae, is described from the
Upper Pliocene Fernando Formation near Newport Bay, Orange
County, California.
Introduction
During February, 1967, earth moving equipment on the east bluff of
upper Newport Bay, Orange County, California, uncovered a highly fossili-
ferous portion of the lower part of the Fernando Formation. This locality re-
mained exposed until August of the same year when further construction
destroyed it. Between these months extensive collecting at the exposure by me
and my associates yielded an interesting fauna of over 200 species of larger
invertebrates characterized by Nuculana fossa (Baird, 1863), Patinopecten
healeyi (Arnold, 1906), Pecten hemphilU Dali, 1879, Glycymeris grewingki
Dali, 1909, Exiloidea rectirostris hertleini Bentson, 1940, Neptunea tabiilata
colmaensis (Martin, 1914), Boreotrophon raymondi (Moody, 1916) and
My a tnuicata Linne, 1758. This assemblage suggests a Late Pliocene age
based on a two-fold division of the Pliocene of California (Vedder, 1960).
While studying the material several new species of mollusks were noted.
One of these belongs to the genus Neadniete Habe, 1961, which was only
recently recognized in the west American fauna (Kanakoff and McLean,
1966). This new species is described herein.
References to the Los Angeles County Museum of Natural History,
Section of Invertebrate Paleontology are hereafter abbreviated as LACMIP.
Acknowledgments
I am grateful for the assistance of the following persons. Dr. Robert F.
Meade, California State College at Los Angeles, and Dr. James H. McLean,
Los Angeles County Museum of Natural History, gave helpful advice and
critically reviewed the manuscript. Mr. Jack Schwartz, Mr. Boris Savic and
Mr. and Mrs. F. L. Grouard aided in collecting the type material. Photographs
are by Mr. Louis W. Kundrath and Mr. James W. Shuttleworth of Kundrath
Studio of Photography, Baldwin Park, California.
iDepartment of Geology, California State College, Los Angeles, 90032.
1
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Contributions in Science
No. 177
Figure 1. Neadmete sduvartzi, new species. Holotype, LACMIP 1779. Height 35.6,
diam. 16.9 mm.
1970
A New Species of NEADMETE
3
Systematic Description
Family Cancellariidae Gray, 1853
Genus Neadmete Habe, 1961
Neadmete schwartzi new species
Figure 1
Diagnosis: Shell relatively large; whorls 8, evenly convex, weakly tabu-
late; 16 spiral ribs on body whorl; outer lip reflected.
Description: Shell relatively large for genus, long, slender, fusiform; spire
elevated. Whorls 8, moderately inflated, evenly convex, weakly tabulate on
early whorls becoming obsolete on penultimate and body whorls. Nucleus
simple, smooth, whorls 2. Axial sculpture of approximately 16 low, rounded
ribs, becoming obsolete on last four whorls. Spiral sculpture of low, narrow,
rounded ribs, 3 on postnuclear whorls and 16 on body whorl; with single
smaller rib intercalated between major ribs on the apical whorls and the first
nine ribs of the body whorl; interspaces slightly wider than ribs. Sutures
moderately impressed. Aperture subeliptical, approximately one half the
height of the shell; outer lip smooth, thin, slightly reflected, with 15 internal
spiral lines; canal short and shallow. Columella long, straight, with two folds
and eight plicae formed by extension of primary spiral ribbing.
Type Material: Holotype, LACMIP 1779. Four paratypes: LACMIP
1780; California Academy of Sciences (CAS), Department of Geology type
collection 13203; University of California at Los Angeles (UCLA), Depart-
ment of Geology type collection 47313; and collection of Jack D. Mount
(JDM) 106. Measurements for the type specimens are listed in Table 1.
Type Locality: LACMIP locality 471; 33° 38' 21" N, 117° 53' 02" W; in
a 6 foot thick lens of sandy cobble conglomerate which occurs approximately
425 feet stratigraphically above the base of the formation. At the present time
it is located under the residence at 2161 Vista Entrada, Newport Beach,
California.
Age and Formation: Late Pliocene, Fernando Formation.
Discussion: Neadmete schwartzi is nearest to the Recent N. circum-
cincta (Dali, 1873:59) but differs from that species principally in the larger
size, more evenly convex, less tabulate whorls, more spiral ribs and in the
presence of the reflected outer lip. N. circumcincta is recorded from Alaska,
with a specimen in the Los Angeles County Museum of Natural History from
Hope Island, off the northern end of Vancouver Island, British Columbia.
N. sutherlandi Kanakoff and McLean, (1966:4) from the Lomita Marl (Plio-
Pleistocene) of Los Angeles County, California, is the only other fossil
species of Neadmete and it is readily distinguished from N. schwartzi by its
larger size, more pronounced tabulation, and smaller number of spiral ribs.
Etymology: This new species is named for Mr. Jack Schwartz, who
4
Contributions in Science
No. 177
collected the holotype, in recognition of his contributions to the mineralogy
of California.
Table 1.
Measurements (in mm) of the type specimens
of Neadmete schwartzi, new species.
Specimen no.
Height
Diameter
Height of
Aperture
LACMIP 1779
35.6
16.9
16.8
LACMIP 1780
28.2
14.8
13.3
CAS 13203
35.3
17.9
16.5
UCLA 47313
33.7
17.1
15.7
JDM 106
31.3
15.7
15.1
Literature Cited
Dale, 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.
Kanakoff, G. P., and J. H. McLean. 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. Los Angeles
County Mus., Cont. in Sci., 116: 1-6.
Vedder, J. G. 1960. Previously unreported Pliocene Mollusca from the southeastern
Los Angeles basin, in Short papers in the geological sciences. U. S. Geol. Survey
Prof. Paper 400-B, B326-B328.
Accepted for publication October 31, 1969.
'it
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LOS
ANGELES
COUNTY i“c?7.7j>
MUSEUM ^
CONTRIBUTIONS
IN SCIENCE
UMBER 178
February 25, 1970
DESCRIPTION OF A NEW SPECIES OF NET-WINGED MIDGE
FROM THE GREAT BASIN, WITH A KEY TO THE NORTH
AMERICAN SPECIES OF THE GENUS DIOPTOPSIS
(DIPTERA: BLEPHARICERIDAE)
By Charles L. Hogue
Los Angeles County Museum of Natural History • Exposition Park
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DESCRIPTION OF A NEW SPECIES OF NET-WINGED MIDGE
FROM THE GREAT BASIN, WITH A KEY TO THE NORTH
AMERICAN SPECIES OF THE GENUS DIOPTOPSIS
(DIPTERA: BLEPHARICERIDAE)
By Charles L. Hogue^
Abstract: The adult, pupa and larva of a new species of
Dioptopsis are described and figured from material collected in
Great Basin ranges in northeastern California and Nevada. The
adult is peculiar among its cogeners in lacking mandibles and
having infuscate wings. It is further characterized along with the
immature stages in a key presented for all the North American
member of the genus.
The amandibulate condition (with associated modifications
of other head structures) of this species and others is discussed
and a new term, colocephalous, is introduced to designate it.
Thanks to the exceptional field work of John F. Emmel and Oakley
Shields over the past three years, material has accumulated of a previously
undiscovered species of Dioptopsis in the Great Basin. For this contribution to
my studies on the Blephariceridae of North America, I wish to express my
appreciation to these spirited collectors. I wish also to thank J. A. Powell and
A. Stone and their respective institutions. University of California, Berkeley
— California Insect Survey Collection [CIS] and Agricultural Research Serv-
ice, U.S. Department of Agriculture — U.S. National Museum [USNM], for the
loan of additional and comparative material. The majority of the specimens
recorded are in the Los Angeles County Museum of Natural History [LACM].
Dioptopsis dismalea, new species
(Figs 1-13)
Dioptopsis n. sp. Hogue, in press. Bull. Calif. Insect Survey.
FEMALE (based on allotype; Figs. 6-8)
Size. — A medium sized blepharicerid. Measurements (lengths in mm):
Body: 3.6 Wing: 5.7
egs:
fore
mid
hind
femur
2.5
2.5
3.4
tibia
2.5
2.3
3.1
tarsus 1
1.2
1.0
1.4
2
0.3
0.3
0.4
3
0.25
0.25
0.25
4
0.2
0.2
0.2
5
0.4
0.4
0.4
^Senior Curator of Entomology, Los Angeles County Museum of Natural History,
Los Angeles, California 90007.
1
2
Contributions in Science
No. 178
Figures 1-8. Dioptopsis dismalea, n. sp., Adult. 1. Wing, 2. Terminal antennal seg-
ments, male. 3. Head, male. 4. Phallosome complex, male genitalia. 5. Male genitalia.
6. Terminal antennal segments, female. 7. Head, female. 8. Female genitalia.
1970
New Species of Net-Winged Midge
3
Figures 9-13. Dioptopsis dismalea, n. sp. 9. Larva. 10. Details of typical median
abdominal segment. 11. Pupa, lateral view of anterior end. 12. Pupa (female).
13. Terminal end of male pupa.
4
Contributions in Science
No. 178
Coloration. — General integument melanic, well sclerotized. Head, in-
cluding antennae, all black. Mesoscutum brownish laterad, large antero-
median rectangular area black, pruinose mesad, shiny black laterad; mesono-
tal sutures, supraanterior pronotal areas, supraalar, scutoscutellar notches and
postscutellum, all shiny black. Scutellum gray pollinose. Pleuron generally
brown, lower sternopleuron and metepimeron contrasting black. Haltere even
gray. Legs gray-black, darker at joints and distal half of tarsal segments 5.
Wing membrane tinged sooty-black. Abdomen evenly black, slightly pruinose.
Head. — Figure 7. Eyes widely separated; upper eye division small, 7-9
rows of ommatidia along maximum vertical arc; not well differentiated from
lower division, no unfacetted strip, ommatidia of both divisions equal in size.
Antenna short, 14 segmented, shape and length of flagellar segments subequal
except basal which is approximately 2.0 length of others, ultimate segment
slightly smaller than penultimate (Fig. 6) . Mouth parts short, mandibles
entirely lacking, hypopharynx truncate, with weak marginal teeth. Palpus
4-segmented, terminal segment (4 and 5) swollen ventromesad, subtriangular
in shape.
Thorax. — Scutellum with dense lateral setal patches, connected by a
single row of setae across caudal perifery; a few supraalar and prescutellar
bristles, anterior pronotum bare; other sclerites nude. Legs with tibial spurs
0-0-2, spurs of hind tibia unequal. Wing shape, venation and trichiation as
figured (Fig. 1).
Genitalia. — Figure 8. Oviscapt deeply cleft mid-posteriorly. Spermathe-
cae moderate in size, one smaller than other two and elongate-ovoid, others
elongate-ovoid, slightly constricted mesally.
MALE (based on holotype; Figs. 1-5)
General. — Slightly larger than female. Measurements (lengths in mm) :
Body: 3.6 Wing: 6.0
Legs:
fore
mid
hind
femur
2.9
3.3
4.3
tibia
3.1
2.8
4.1
tarsus 1
1.8
1.4
1.8
2
0.6
0.5
0.6
3
0.4
0.4
0.4
4
0.3
0.3
0.3
5
0.4
0.4
0.5
Coloration. — Generally darker than female. Essentially sooty-black all
over; abdominal pleural membranes contrasting white. Wing membrane tinged
sooty-black as in female.
Head. — Figure 3. Practically indistinguishable from that of female. Eyes
widely separated. Hypopharynx truncate, marginal teeth absent. Tip of labrum
1970
New Species of Net-Winged Midge
5
entire. Antenna short; 14-segmented, relative size and shape of segments as
in female.
Thorax. — Chaetotaxy as in female, a few bristles on anterior pronotum.
Legs with tibial spurs 0-0-2, hind pair very unequal. Wing as in female.
Genitalia. — Figures 4 and 5.
PUPA (Figures 11-13)
Size. — Small to medium. Measurements (overall length in mm) : 5 range
4. 9-5. 6, mean (n = 4) 5.1. $ range 5. 2-6.0, mean (n = 10) 5.5.
Coloration. — Dorsally all black, no pattern.
Structure. — Figures 1 1-13.
LARVA (final instar; Figures 9 and 10)
Variation: — Fortunately material has recently come to hand from a third
locality, in the Jarbidge Mountains, to indicate that this species is discordantly
variable in respect to several features of the male genitalia and one pupal
character. Formerly, specimens whose differences suggested the existence of
two geographical subspecies were only available from the Warner Mountains
of California and Ruby Mountains of Nevada.
In the genitalia of the male the following characters are notably variate.
( 1 ) Angle of projection of the 9th tergite lobes. The lobes tend to lie straight
and parallel, thus angled at about 90° from the posterior margin of the seg-
ment, in the Ruby Mountain populations and variously curved and con-
vergent in the other two populations. (2) Extent of bristles on the sides and
venter of the basistyle. These bristles are mostly lateral but a few stray
mesoventrad, especially in the Nevada populations. (3) Shape of tip of the
paramere. Usually the apex of this organ is rounded and slightly inturned with
the canal running nearly to the end. In some specimens from the Jarbidge
Mountains, the apex is extended beyond the canals to form a spatulate process.
(4) Shape of dorsal lobe of outer dististyle. In dorsal aspect this lobe appears
club shaped and straight in the Jarbidge Mountains specimens but tend to be
capitate and incurved in males from the Ruby and Warner Mountains. (5)
Length of ventral projection of the inner lobe of the dististyle. This process
variously exists as a short point hardly longer than the body of the main lobe,
to a long fingerlike projection curving up and beyond the latter.
A dimorphism occurs in the extent of dorsal integumentary stippling on
the pupa. In the California populations, these structures terminate abruptly
short of the lateral margins leaving a clear zone around most of the abdominal
periphery. Nevada pupae show the stippling strongly up to, and even carrying
ventrad of, the lateral edge of the abdominal segments.
Range: The species is now known only from extreme western and eastern
mountains in the Great Basin and Range geological system. It is certain to be
found throughout the intervening ranges where streams occur at elevations of
over 7000 feet.
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Contributions in Science
No. 178
Diagnosis'. The pigmented wing of this species is unique among Dioptop-
sis, the wing membrane of all other species being hyaline. Other characteristics
to distinguish adults and the immatures are given in the keys below (see
Discussion).
Type Locality and Material : The types of this species come from streams
draining Mount Bidwell through a large meadow in the Warner Mountains of
extreme northeastern California (Modoc County) called Dismal Swamp
( 119° 10' W, 41° 59' N) . Specific collections are as follows:
HOLOTYPE: ^ , (genitalia on slide No. CLH 6809-3) and ALLOTYPE,
$, (genitalia on slide No. CLH 6809-6) : Dismal Swamp, 7200 feet, 15-16
July 1968 (J. Emmel and O. Shields) [LACM].
PARATYPES: 5 $ $ , 6 $ $ : same data as holotype (one pair com-
pletely dissected, on slides) [LACM, USNM].
^ : creek just west of Dismal Creek, 7300 feet, 15 July 1968 (Emmel
and Shields) [LACM].
ADDITIONAL SPECIMENS: CALIFORNIA, Modoc County. 65
pupae and pupal skins: creek just west of Dismal Creek, 7300 feet, 15 July
1968 (Emmel and Shields) [LACM].
103 pupae: Dismal Creek, 7300 feet, 15 July 1968 ( Emmel and Shields)
[LACM].
3 pupae: stream west slope Mount Bidwell, 1 mi. S Dismal Swamp,
8000 feet, 16 July 1968 (Emmel and Shields) [LACM].
66 pupae, 75 larvae: Dismal Swamp, 8000 feet, 15 July 1948, 1950
(W. W. Wirth) [USNM, CIS].
NEVADA, Elko County. 27 pupae and pupal skins, 81 larvae: stream
connecting Island Lake and Lamoille Creek, Ruby Mountains, 8800-10,000
feet, 8 August 1967, 29 June 1968 (Emmel and Shields) [LACM].
16 S S ,15 pupae: Bonanza Gulch, 7800-8000 feet, Jarbidge Mountains,
23 July 1969 (Emmel and Shields) [LACM].
18 pupae: creek west of Angel Lake, 8600 feet. East Humboldt Range, 26
July 1969 (Emmel and Shields) [LACM].
Biological Notes : A]1 loca]ities where D. dismalea has been taken lie
between 7300 to 10,000 feet elevation. The habitats are accordingly subalpine
and alpine but vary to a considerable extent in several features important to
blepharicerid ecology. According to Emmel (personal communication) , “The
Dismal Swamp is a large wet meadow (not a swamp, really) with willow
thickets around the edges. There are scattered patches of thick Pine forest on
the surrounding slopes. The “swamp” is not in a basin or valley as such, but is
on the edge of a high plateau which slopes toward the east, just east of the crest
of the Warner Mts. Several very small streams flow into the Dismal Swamp
from the slopes above it on the west and south sides. Probably the largest of
these streams are two flowing from the slopes of Mt. Bidwell. The streams flow
through open aspen forest just above the edge of the large wet meadow; it is in
these areas of open aspen forest where blepharocerid material was most
1970
New Species of Net-Winged Midge
7
common. Larvae or pupae were not found more than several hundred yards
away from the Swamp (although the streams [terminated] not too far above
the areas where pupae stopped appearing), nor in the Swamp itself. Larvae
and pupae were found on stones ranging in size from 1" diameter up to 6"
diameter (which were probably the largest rocks). The stream rocks were
generally very dark in coloration, although not usually as dark as the pupae on
them. A few pupae were also found in a small stream high on the western
slope of Mt. Bidwell where there was no forest; small willow thickets
surrounded this stream. In all areas where blepharocerid material was
collected, the stream habitat here was in contrast to other blepharocerid stream
habitats in which we have collected, in that these streams were mere trickles
(although this was a very dry year) rather than rushing torrents of abundant
water as is usually the case. The observation that larvae and pupae were on
relatively small rocks (the only ones available) is also of interest. The habitat
of this species is noticeably different from that of . . . the Ruby Mountains
locality . . . [which] . . . looks very much like a typical stream in the high
altitudes of the eastern slope of the Sierra Nevada. Lamoille Canyon has been
well glaciated; many streams flow down its steep slopes in the early summer.
The stream in which larvae and pupae were found flows down an open slope
into Lamoille Creek; blepharocerid material was found in the swiftest water,
on dark rocks (probably granite). The whole area looks not unlike the higher
altitudes of the Sierra Nevada, and it is well watered. Aspens are more
abundant than conifers at this altitude; trees occur in scattered patches, never
as extensive forest.”
The site in the stream above Angel Lake in the East Humboldt Range is
like the preceding, a nearly vertical narrow cascade down a rough granite face
of a steep glaciated valley. There is scarcely any vegetation in the immediate
vicinity of the stream. On the other hand at Bonanza Gulch, in the Jarbidge
Mountains, the gently tumbling stream in the bottom of a broad ravine flows
through a jumble of log and debris jams and is heavily overgrown with tall
clump grass and willows.
Discussion : The new species clearly shows a complete lack of mandibles.
This is a normal condition in only one other North American blepharicerid,
Blepharicera ostensackeni Kellogg, but occurs consistently in many exotic
species and in populations (subspecies ?) of Dioptopsis aylmeri (Garrett) in
the eastern drainage of the Sierra Nevada (Hogue, in press) . Associated with
the amandibulate condition is a complete rearrangement in head proportions
including a much shortened labrum and other mouthparts, weak hypopharyn-
geal teeth, widely separated eyes and reduced upper eye divisions. This type
of head may be designated, “colocephalus” (Gr. Kolos, stunted, incomplete) .
The genetic and phylogenetic significance of colocephaly which is ana-
malous in some species and normal in others, deserves special study. I have
expressed (Hogue, in press), as has Edwards (1929:35-36), the opinion that
such a total reorganization of head construction could come about by pleio-
8
Contributions in Science
No. 178
trophic effects of simple mutations. Thus what would appear to be a profound
set of differences upon which new species or other taxa might be based
actually may easily and frequently arise in individuals or populations (even
becoming dominant in one or more geographic areas as has presumably
occurred with eastern Sierran D. aylmeri) of a single species. A trimorphic or
even quadrimorphic condition could exist wherein a species may have males
and females alternately with or without mandibles and with associated modi-
fied head structure. Such a hypothesis was long ago advanced by Muller but
was refuted by other dipterists (Alexander, 1958:814) on the grounds that
examples of dimorphic females actually belonged to different species. I believe
the phenomenon needs further study to determine its extent, cause, and
adaptive significance.
With the discovery of D. dismalea, the genus Dioptopsis in North Amer-
ica now consists of six species:
1. aylmeri (Garrett, 1923) . Can. Ent. 55:244 {cheaini Garrett, 1925, to be
placed in synonomy in forthcoming work by author) . Widespread in west-
ern Canada and the United States.
2. sequoiarum (Alexander, 1952) . Bull. Brook. Entomol. Soc. 47:91. Cali-
fornia, western Sierra Nevada.
3. dismalea Hogue, 1969. Great Basin Ranges.
4. arizonica Alexander, 1958. Bull. Brook. Entomol. Soc. 53:50-51. Arizona,
Sierra Ancha Mountains.
5. alpina Hogue, 1966. Los Angeles Co. Mus. Nat. Hist., Contrib. Sci., 120:
1-5. Northern and southern California.
6. markii (Garrett, 1925) . Seventy New Diptera, p. 5. Recorded only from
northern California, Washington and Alberta.
The first three species are very close relatives and may be united into an
Aylmeri group. The primary basis of the grouping is a common larval type. In
spite of the colocephalous heads of adult dismalea giving it a markedly different
character from aylmeri (with the exception of certain populations as men-
tioned above) and sequoiarum, the species are indistinguishable in the larval
stage. Arizonica and alpina together form a second natural grouping on the
basis of features I have outlined elsewhere (Hogue, 1966:5). Markii stands
alone with very different larval, pupal and adult characteristics.
The following keys will facilitate identification of the adults and im-
matures of all these species, with the exception of arizonica, whose female,
pupa, and larva are unknown.
KEY TO NORTH AMERICAN DIOPTOPSIS SPECIES
ADULTS
(Female of arizonica unknown)
1. General: Wing membrane infuscated. Palpus four segmented dismalea
General: Wing membrane hyaline. Palpus five segmented. 2
1970
New Species of Net-Winged Midge
9
2. General: Upper eye division at least one-fifth to one-fourth the area of
lower in males (at least 11-12 rows of ommatidia), greater than lower in
females 3
General: Upper eye division greatly reduced, much smaller than lower
in both sexes, a small area of 4-7 rows of ommatidia 4
3. Male genitalia: Paramere with conspicuous mesal spur. Mesal margin of
IX tergite lobe produced into a nude flange (i.e. without micro- or
macrochaetae) . Female genitalia : Spermathecae ovoid aylmeri (in part)
Male genitalia: Paramere simple, without spur. Mesal margin of IX
tergite lobe not produced. Female genitalia: Spermathecae elongate-
ovoid sequoiarum
4. General: Antenna with 15 segments. Mid-tibial spur present though
minute. Male genitalia: IX tergite lobe short, only 2 times width. Lobe of
outer dististyle short and projecting dorsomesad 5
General: Antenna with 14 segments or less. Midtibial spur entirely
absent. Male genitalia: IX tergite lobe elongate, length 5-6 times width.
Lobe of outer dististyle elongate and projecting subparallel to dististyle
face 6
5. Male genitalia: Tip of inner dististyle lobe recurved alpina
Male genitalia: Tip of inner dististyle lobe with cup-shaped depression
arizonica
6. General: Ultimate antennal segment smaller than penultimate. Female
mandible reduced to absent. Male genitalia: Paramere with spur. Female
genitalia: Spermathecae pear-shaped aylmeri (in part)
General: Ultimate antennal segment longer than penultimate. Female
mandible normal sized. Male genitalia: Paramere simple. Female geni-
talia: Spermathecae ovate markii
PUPAE
(Unknown for arizonica)
1. Branchial sclerite strongly lobed, projecting well beyond cephalic sclerite
in lateral aspect alpina
Branchial sclerite weakly lobed, cephalic sclerite visible from lateral
aspect 2
2. Branchial lamellae spreading, caudal pair strongly convergent, often
overlapping markii
Branchial lamellae all subparallel 3
3. Length/ width of anteriormost branchial lamella 1.1 or larger dismalea
Length/ width of anteriormost branchial lamella 1 .0 or smaller 4
4. Interbranchial index ( =interbranchial distance at base of lamellae/width
of anteriormost lamella) less than 0.7. Rugae of posterior region of
scutum forming a line running parallel to hind margin of sclerite
sequoiarum
10 Contributions in Science No. 178
Interbranchial index greater than 1.0. Rugae of posterior region of
scutum random, not forming a line. aylmeri
LARVAE
(Unknown for arizonica)
1. Dorsal tubercles and plates present alpina
Dorsal integument without tubercles or plates 2
2. Antenna indistinctly segmented, basal % membranous. Ventral gill tuft
of segment 6 diminutive, with only 2 filaments. Dorsal pseudopods
gently curving cephalolaterad markii
Antenna clearly 3 segmented, middle segment much smaller than others.
Ventral gill tuft of segment 6 subequal to others, with 5 filaments. Dorsal
pseudopods angular, tips project ventrolaterad
aylmeri, sequoiarum, dismalea
Literature Cited
Alexander, C. P. 1958. Geographical distribution of the net-winged midges
(Blepharoceridae, Diptera). Proc. Tenth Int. Congr. Entomol. 1 (1956): 813-
838.
Edwards, F. W. 1929. Blepharoceridae in Diptera of Patagonia and South Chile,
Part II, fasc. II, pp. 33-75, 3 figs., 4 plates.
Hogue, C. L. 1966. A new species of Dioptopsis from California (Diptera: Bleph-
aroceridae). Los Angeles Co. Mus. Nat. Hist., Contrib. Sci., 120: 1-5.
“. in press. The net-winged midges or Blepharoceridae of California. Bull.
Calif. Insect Survey.
Accepted for publication November 3, 1969.
j ;los
i ' ANGELES
I !
1 COUNTY
l:'
I MUSEUM
1"
ftlBER 179
CONTRIBUTIONS
IN SCIENCE
February 25, 1970
GEOGRAPHIC VARIATION AND SYSTEMATIC
STATUS OF MEXICAN LYRE SNAKES OF THE
TRIMORPHODON TAU GROUP (COLUBRIDAE)
By Roy W. McDiarmid and Norman J. Scott, Jr.
Los Angeles County Museum of Natural History
Los Angeles, California 90007
Exposition Park
CONTRIBUTIONS IN SCIENCE is a series of miscellaneous technical papers
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GEOGRAPHIC VARIATION AND SYSTEMATIC
STATUS OF MEXICAN LYRE SNAKES OF THE
TRIMORPHODON TAU GROUP (COLUBRIDAE)
By Roy W. McDiarmid^ and Norman J. Scott, Jr."
Abstract: Geographic variation in lepidosis and color pat-
tern was studied in the nominal species Trimorphodon collaris,
fasciolata, forbesi, latifascia, tan, and iipsilon. Specimens were
grouped into seven geographically separated samples which were
then compared. Ranges of most characteristics overlap broadly;
others vary independently and discordantly. Ventral-subcaudal
scales decrease from north to south. Snakes from the Balsas-
Tepalcatepec Basin have fewer dorsal bands than do snakes from
other areas. They also have a relatively uniform head pattern.
Specimens from other areas have a complex head pattern con-
sisting of prefrontal bars, interocular bars, and discrete parietal
marks in various combinations. Head pattern and body band
analyses indicate limited genetic exchange between the Balsas-
Tepalcatepec Basin samples and those from other areas, appar-
ently the result of secondary contact in southern Jalisco and
adjacent Michoacan. The Transverse Volcanic Range and the
Sierra Madre del Sur apparently are effective in restricting genetic
exchange.
The above nominal species are assigned to the synonymy of
Trimorphodon tan. Trimorphodon tan tan is maintained for the
foothills and plateau populations and T. tan latifascia (new com-
bination) is assigned to the Balsas-Tepalcatepec Basin popula-
tions. Trimorphodon tan is redescribed and its distribution
delimited. Character variation in a brood from Puebla approaches
the variation typical of the entire Puebla sample.
Introduction
In the years following 1935, rather extensive collections of amphibians
and reptiles were made in Mexico. Early reports on many of these collections,
culminating in the Mexican checklists by Hobart M. Smith and Edward H.
Taylor (1945, 1948, 1950), are extremely useful in elucidating the distribution
and relationships of the many components of this complex herpetofauna. Dur-
ing the past 15 years, reports on collections from various parts of Mexico have
continued to appear. Unfortunately, some of these reports have confused rather
than clarified our knowledge of the distributions and systematics of certain of
the Mexican species (see Duellman, 1966, for additional discussion). An
example of such confusion is found within the snake genus Trimorphodon.
^Research Associate in Herpetology, Los Angeles County Museum of Natural His-
tory and Collegiate Division of Biology, University of (Chicago. (Current address:
Dept, of Zoology, University of South Florida, Tampa.)
“Biological Sciences Group, University of Connecticut.
1
2
Contributions in Science
No. 179
All 12 nominal species of Trimorphodon have been reported from Mexico
(Smith and Taylor, 1945) . Smith ( 1941 ) divided the genus into two apparently
natural groups. The biscutatus group, characterized by large chevron-shaped
marks on the neck and head, included five nominal species {biscutatus, lambda,
lyrophanes, paucimaculatus and vandenburghi) that range generally from the
southwestern United States southward through the Pacific lowlands to Costa
Rica. The tau group, characterized by a light, transverse nuchal collar of vary-
ing width, included seven nominal species {collaris, fasciolata, latifascia, tau,
upsilon and vilkinsonii) whose ranges encompass upland and mountainous
portions of southeastern New Mexico, southwestern Texas and Mexico, north
of the Isthmus of Tehuantepec.
We became interested in the Trimorphodon tau species group in the course
of an extensive study of the herpetofauna of Sinaloa, Mexico (Hardy and
McDiarmid, 1969). In attempting to allocate specimens, we realized the need
for a revision of this group of snakes and so began a study of the geographic
variation and systematic relationships among the described forms, with empha-
sis on the zoogeographic implications of the distribution of specific character-
istics.
Historical Review
Cope (1869) described Trimorphodon tau from a single specimen col-
lected by Sumichrast, supposedly from the western part of the Isthmus of
Tehuantepec, and Trimorphodon upsilon from a specimen collected in Guada-
lajara, Mexico by Major (see Zweifel, 1959 for discussion of this collection).
Later in the same year, Peters (1869) described Dips as bis cut at a var.
latifascia from specimens collected in Puebla by Berkenbusch.
These descriptions were the first for representatives of the tau species
group, though specimens of these species were known previously. Jan (1863)
listed four specimens of Eteirodipsas biscutata, one from Mexcio. Nine years
later this same specimen was illustrated in Jan and Sordelli (1872) and listed
as part of the collection of “Westphal-Castelnau a Montepellier.” The Mexican
specimen definitely is not Trimorphodon biscutatus (Dumeril, Bibron and
Dumeril, 1854), nor is it representative of a species of the biscutatus group as
presently understood. In fact the color pattern, especially of the head (Jan and
Sordelli, 1872: vol. 3, livr. 39, pi. 1, no. 3) , is similar to the pattern of the holo-
type of Cope’s Trimorphodon upsilon.
Cope (1875) described a fourth species of the tau group, Trimorphodon
collaris. This specimen, also collected by Sumichrast, was from Orizaba, Vera-
cruz. In 1882 Sumichrast pointed out that the type specimen of Trimorphodon
tau was collected near Quiotepec, between Techuacan and Oaxaca and was
not from the western part of the Isthmus of Tehuantepec as stated by Cope
(1869) in the original description. In this same publication, Sumichrast (1882)
listed “los individuos tipicos” of Trimorphodon collaris from Tuxpango, near
1970
Geographic Variation of Mexican Lyre Snakes
3
Orizaba, Veracruz. However, in the description of T. collaris Cope (1875)
made no reference to specimens other than the type. The location of any addi-
tional specimens of T. collaris collected by Sumichrast is unknown.
Gunther (1895) pointed out the similarities between Trimorphodon tan,
collaris and upsilon, and referred his specimens to T. upsilon. Gunther con-
sidered Peters’ latifascia synonymous with Trimorphodon biscutatus. Boulen-
ger (1896) recognized Trimorphodon tan and T. upsilon as distinct species.
Boulenger followed Gunther’s suggestion and included Trimorphodon collaris,
as well as Trimorphodon biscutatus latifascia (Peters), in the synonymy of
Trimorphodon upsilon.
Taylor (1939) reviewed Mexican species of Trimorphodon and revived
Peters’ name latifascia for specimens from Puebla and Morelos. Taylor recog-
nized T. upsilon and T. tau as distinct species but considered T. collaris synony-
mous with T. latifascia.
In 1941 Smith surveyed the genus Trimorphodon and established the two
species groups. He considered Trimorphodon latifascia, T. upsilon, T. tau and
T. collaris distinct species. In addition Smith ( 1941 ) described two more forms
in the tau group: Trimorphodon fasciolata from Tzararacua Falls, Michoa-
can (one specimen) and Trimorphodon forbesi from San Diego, Puebla (one
specimen) .
After examining specimens of Trimorphodon from Michoacan, Schmidt
and Shannon (1947) suggested that T. fasciolata might be conspecific with
T. latifascia. Davis and Smith (1953) and Peters (1954) demonstrated that
these two forms probably were representative of a single species.
Smith and Darling (1952) examined some specimens from Tamaulipas
and San Luis Potosi and regarded Trimorphodon upsilon as a subspecies of
Trimorphodon tau. Martin (1958) confirmed the conspecificity of these forms.
Duellman (1961) concluded that data were inadequate to support the
retention of upsilon as a subspecies of Trimorphodon tau. He regarded Tri-
morphodon tau as a monotypic species that has a highly variable color pattern.
Thus, if the systematic changes since Smith’s revision (1941) are accepted,
there are currently four species in the tau group, Trimorphodon tau, collaris,
forbesi and latifascia.
Jones and Findley (1963) suggested that T. vilkinsoni might be related
to T. lambda, a member of the biscutatus group.
Methods and Materials
We have examined all known specimens, including holotypes, of the
following nominal species of the Trimorphodon tau group: tau, upsilon, col-
laris, latifascia, fasciolata and forbesi. Locality, sex (determined by dissection),
and the following features were recorded for each of the 197 specmiens: num-
ber of ventral scales (counted according to the method outlined by Dowling,
1951), number of subcaudal scales including the tail tip, total number of
4
Contributions in Science
No. 179
ventral and subcaudal scales, nature of the anal plate (divided or entire),
number of supralabials, supralabials that contact eye, number of infralabials,
the infralabial that is first separated from the chin shields by small gular scales,
number of loreal scales (all scales between the preocular and nasal scales),
number of preocular and postocular scales, and the number of primary and
secondary temporal scales. The dorsal scale rows were counted one head
length behind the parietal scales, at midbody and one head length in front of
the anus. The number of scales in the light collar counted on the midline was
recorded, as was the number of body and tail bands or blotches. The first cross
band that showed any indication of splitting (the presence of a light center on
the midline) was recorded. For each band-interspace-band sequence the num-
ber of scales on the midline was counted beginning with the first dark band-
light interspace-second dark band, for the seventh dark band-light interspace-
Figure 1. Distribution of Trimorphodon tan in Mexico. Each dot represents a locality
or localities of one or more specimens. Solid dots indicate specimens examined; circles
indicate additional specimens or literature records. Samples analyzed are lettered A
through G.
1970
Geographic Variation of Mexican Lyre Snakes
5
eighth dark band, and at equivalent intervals along the body of the snake. In
all instances the band-interspace-band sequence immediately anterior to the
vent was counted. Detailed drawings of the dorsal head pattern were made for
each specimen.
Specimen localities were plotted on a topographical map of Mexico and
grouped into seven geographical units (Fig. 1). These seven samples, here-
after referred to as populations A through G, include specimens from the
following areas :
Population A (25 specimens, — from localities in the foothills of the Sierra
Madre Occidental of southern Sonora and northern Sinaloa. These localities
are separated from population B to the south by nearly 480 kilometers.
Population B (15 specimens) — from localities in the upper drainage of
the Rio Santiago and its tributaries in Nayarit and Zacatecas and from the
general vicinity of Guadalajara and Lago de Chapala in Jalisco and adjacent
Michoacan. These snakes probably are continuously distributed in suitable
habitats northward through Sinaloa, as indicated by two specimens from
western Durango, but are separated into two populations, A and B, because
of the lack of adequate material from the intervening area.
Population C (14 specimens) — from localities in Tamaulipas, San Luis
Protosi and Hidalgo in the eastern drainage systems of the Rio Panuco and
Rio Tamesi. There are six specimens known from within the 400 kilometer
hiatus that separates population C from population B, the latter of which is
restricted to western drainages.
Population D {20 specimens) — from localities in central Oaxaca. This
sample is situated about 480 kilometers southeast of population C with only a
few specimens known from the intervening area. The uplifted southern section
of the Mexican Plateau effectively separates populations C and D.
Population E {51 specimens) — a large sample from the vicinity of Chil-
pancingo, Guerrero located in the central part of the Sierra Madre del Sur.
These localities are isolated from sample D by the eastern extension of the
Sierra Madre del Sur and the mountains in northwestern Oaxaca.
Population F (25 specimens) — from areas in northeastern Guerrero and
adjacent Mexico and Morelos and from southwestern Puebla. These localities
are in areas drained by the northeastern tributaries and headwaters of the Rio
Balsas. Population F is separated from E by the arid Balsas Basin and from D
by the mountainous areas of northwestern Oaxaca. Population F is separated
from sample C south of the volcanic peaks and high southern portions of the
Mexican Plateau.
Population G (19 specimens)— from parts of Colima, southern Jalisco
and western Michoacan. These localities are situated in areas drained by the
Rio Armeria and Rio Coahuayana ( Autlan and Contla, Jalisco and Colima) or
by the major northwestern tributary of the Rio Balsas, the Rio Tepalcatepec
(Michoacan). This sample is isolated to the northwest of E by the arid Balsas
6
Contributions in Science
No. 179
Basin and to the south of B by the Transverse Volcanic Range and western
portions of the Mexican Plateau.
Specimens removed from the geographic range or topographically iso-
lated from these units were considered separately. In other words, we did not
extend the geographical coverage of our units to accommodate individual
specimens.
After the seven groups were established, the meristic, scale and pattern
characteristics were compared within each unit and among the seven units.
Geographical Variation
LEPIDOSIS
Ventral and subcaudal scales. — Ventral scales vary from 20-231 in males
and from 210-243 in females. Subcaudal scales range from 61-85 in males and
from 55-80 in females. Females average 2-11 more ventrals than males in the
various populations. However, males on the average possess 8-1 1 more sub-
caudals than do females in the same population. The relatively shorter body and
longer tail of males, as indicated by fewer ventrals and more subcaudals, appar-
ently is the result of their more anteriorly placed cloacal opening. This condi-
tion is probably an accommodation for the relatively long hemipenes in the
tail. The converse probably is true of females which have a longer, more
bulky body to accommodate eggs (Klauber, 1956:167).
The number of ventral and subcaudal scales in snakes is indicative of the
number of vertebral segments (Dowling, 1951:99). To test the assumption
that there is little or no sexual dimorphism in the total number of vertebral
elements in members of the same geographic population, we analyzed the
number of ventral plus subcaudal scales for males and females in each of
the seven samples using the Student’s t-test. No significant differences (95 per-
cent level) were found. Therefore, for purposes of analysis, we combined the
ventral and subcaudal counts of males and females in each population.
The ventral-subcaudal scales range from 265-319. The higher numbers
are characteristic of the northwestern population in Sonora and Sinaloa and
the lower numbers are from the southern population in Oaxaca. Variation in
numbers of ventral-subcaudal scales in the seven populations is illustrated
(Fig. 2) . The relative geographic positions of specimens from localities outside
of the geographic limits of the prescribed populations are plotted by number
in Figure 2.
There is a north to south reduction in the number of ventral-subcaudal
scales within population A. The specimens from southern Sonora have 298-
319, X = 307 (N=ll). Specimens from the Rio Fuerte in northern Sinaloa
have 300-306, x = 303 (N=4). Specimens from north central Sinaloa have
293-308, X = 303 (N=10). Two specimens from Ventanas, Durango (300
and 302 scales), and a specimen from the vicinity of San Bias, Nayarit (294
scales) are the only known specimens from the 480 kilometer distance between
1970
Geographic Variation of Mexican Lyre Snakes
7
populations A and B (Fig. 2, Nos. 1-3). Their counts are intermediate and
suggest that additional material from the intervening area would show that
the north to south reduction in the number of ventral-subcaudal scales ex-
hibited in population A continues into B. A specimen from La Cumbre, Jalisco
(Fig. 2, No. 4) has 292 ventral-subcaudal scales.
Within population B, three high counts (299, 299, 303) are characteristic
of specimens from northwestern localities of population B, at Barranquitas,
Nayarit, and near Magdalena and Bolanos, Jalisco, respectively. Two speci-
mens from the southeastern parts of population B near Emiliano Zapata,
Michoacan have low counts (282, 287).
Figure 2. Statistical analysis of the number of ventral plus subcaudal scales in popula-
tions A through G of Trimorphodon tau. The number in parenthesis indicates sample
size of each population. The horizontal lines indicate sample range and the vertical
lines sample mean. The black rectangles represent one standard deviation on each
side of the mean; the white rectangles represent 2.646 standard errors on each side of
the mean (Eberhardt, 1968). Dots represent specimens from the following inter-
mediate localities outside the geographic range of the samples: 1, 2, Ventanas,
Durango; 3, N of San Bias, Nayarit; 4, La Cumbre de los Arrastrados, Jalisco; 5, 7 mi
S Rincon de Romos, Aguascalientes; 6, 7, Tacicuaro, Michoacan; 8, 10, Guanajuato,
Guanajuato; 9, Mt. Cubilete, Guanajuato; 11, Jalapa, Veracruz; 12, Tuxpango, Vera-
cruz; 13, San Diego, Puebla; 14, Quiotepec, Oaxaca.
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Contributions in Science
No. 179
Specimens from population C in eastern Mexico have fewer ventral-
subcaudal scales than specimens from B and in this character indicate a con-
tinuation of a dine. The trend is continued within population C in that
northern specimens have higher totals than do southern specimens. The highest
counts (290, 300) are found in northern specimens from Tamaulipas, while
the lowest counts (278, 277) are from snakes from more southern localities in
San Luis Potosi and Hidalgo.
Two specimens from Tacicuaro, Michoacan on the interior drainage of
the Rio Grande de Morelia and three from the vicinity of Guanajuato, Guana-
juato in the drainage of the Rio Lerma are situated between populations B and
C. Four of these five specimens exhibit ventral-subcaudal counts that lie on or
between the means for populations B and C, and thus reflect their intermediate
geographic position (Fig. 2, Nos. 6-10).
Specimens from population D have the lowest ventral-subcaudal counts
and are from localities at the southern limit of the range. Most of these locali-
ties lie within the Pacific drainage of the Rio Verde althought some are from
localities in the Atlantic drainage of the Rio Papaloapan. These eastern speci-
mens, when considered together with two other specimens from eastern locali-
ties north of population D (Fig. 2, Nos. 11, 14), suggest a possible connection
between samples C and D along the eastern edge of the Mexican Plateau. The
specimen from Jalapa, Veracruz (Fig. 2, No. 11) and the type of Trimorpho-
don tan from Quiotepec, Oaxaca (Fig. 2, No. 14) have ventral-subcaudal
counts that are within the range of variation of population D but slightly higher
than the average and thus approach populations C and F in this character.
Snakes from the largest sample, population E, average about 13 more
ventral-subcaudal scales than specimens in population D and, in this respect,
are nearly identical to those in population C (Fig. 2). Sample C is located
nearly 400 kilometers north of E and separated from it by the Balsas Basin and
the Mexican Plateau.
Population F averages slightly more ventral-subcaudal scales than E, but
this difference is not significant at the 95 percent level. All known specimens in
sample F are from localities in the upper Balsas Basin. The holotype of
Trimorphodon collaris from western Veracruz (Fig.. 2, No. 12) and the holo-
type of T. forbesi from southern Puebla (Fig. 2, No. 13) are both from locali-
ties in eastern drainage systems. These two specimens have ventral-subcaudal
counts similar to those of population F. Thus, specimens from southeastern
Puebla and adjacent Oaxaca and from Veracruz have two patterns in the
number of ventral-subcaudal counts. The holotypes of Trimorphodon collaris
and T. forbesi (Fig. 2, Nos. 12, 13) are more similar to specimens from popu-
lation F, while the holotype of T. tan and the Jalapa specimen (Fig. 2, Nos.
11, 14) are more similar to snakes from population D.
Snakes from population G have more ventral-subcaudal scales than are
found in either E or F. In this respect sample G approaches sample B which
1970
Geographic Variation of Mexican Lyre Snakes
9
lies immediately to the north. There is no significant difference (95 percent
level) between samples B and G in the numbers of ventral-subcaudal scales.
This analysis shows that there is a clinal trend in total ventral-subcaudal
scales within population A and among populations A, B, C and D, with higher
counts in the north and lower counts in the south. Populations E, F, and G
are more closely related to each other than they are to either of the two
geographically adjacent populations, B in the north and D in the south. How-
ever, these three Balsas-Tepalcatepec Basin samples (E, F, G) have the same
trend of higher totals in the north and lower totals in the south. In this instance,
however, the trend is somewhat less marked.
Anal scale. — The anal scale is divided in all specimens except one. A male
from Morelos (TCWC 7390) has an entire anal scale.
Loreal scales. — The number of loreal scales varies from two to five on
each side of the head, with the majority of specimens having either two or
three loreal scales (Table 1 ) . Two loreals are more common in specimens from
populations B, C and D, while three loreals are typical of populations A, E, F
and G.
Preocular scales. — Although preocular scales may vary in number from
one to four, the majority of snakes in all populations have three (Table 1).
Specimens with two preocular scales are found in all populations except D. A
single preocular is present in one specimen from population F.
Postocular scales. — Three postocular scales are characteristic of most
specimens (Table 1). All snakes from sample F have three postoculars. Two
postocular scales occasionally occur in the other six samples. Four scales are
known only from specimens in populations A and E.
Temporal scales. — Most specimens have either two or three primary
temporals and three or four secondary temporals. The majority of specimens
in populations A, E, F and G have three primary and four secondary temporal
scales, while specimens from population B, C, and D usually have two primary
and three secondary temporals (Table 1). The differences may reflect the
smaller sample sizes of these populations. A single primary temporal occurs
only once, in population B; five primary temporals occur only once, in popula-
tion E. In five instances snakes from sample D have two secondary temporals.
A comparison of the total number of lateral head scales (loreals, oculars
and temporals) for the specimens from each sample shows that the higher
numbers and means are characteristic of populations A (28-37, x = 30.9,
N=23), E (26-39, x = 30.8, N = 48) and F (26-34, x = 31.2, N = 25), while
the lower totals and means are found in samples B (23-31, x = 28.2, N=14),
C (25-31, X = 27.9, N=12) and D (24-32, x = 28.2, N=20). Population G
(24-32, X = 29.0, N=19) is somewhat intermediate in total head scales but is
slightly closer to the lower than to the higher values. There is considerable
variation in the number of head scales within some populations and some
variation among several populations. We do not consider the different head
10
Contributions in Science
No. 179
scale characteristics to be of systematic importance in distinguishing any of
the samples.
Labial scales. — Most snakes have either eight or nine supralabials, except
in population C where all specimens have eight (Table 1 ) . Eight is the more
common number of supralabials in all populations except F, in which nine
occurs more frequently. Only in sample D are specimens with seven supra-
labials found. The type specimen of Trimorphodon tau has six supralabials on
each side, the lowest number known.
Although the fourth and fifth supralabials border the eye in the majority
of specimens examined, some variation was noted. In populations A, D, E, and
F occasional individuals have the fifth and sixth supralabial bordering the eye.
One specimen from each of populations F and G has the third, fourth and fifth
supralabials bordering the eye on one side while the fourth and fifth border the
eye on the other. The fifth supralabial borders the eye on one side in two
specimens from population F; the usual condition is found on the other side.
There is considerable variation in the number of infralabials from popu-
lation to population and within populations (Table 1). Twelve is the most
common number, although ten, eleven, and thirteen infralabials are found in
nearly all samples. Population D has the lowest recorded number of infra-
labials (nine) and has a lower mean than do the other six samples.
The small gular scales usually separate the infralabial series from the chin
shields at the level of the fifth or sixth infralabial scale. In population A this
separation occurs in the majority of snakes at infralabial six. In population D
this separation occurs in most specimens at infralabial five. In the other five
populations the separation occurs about equally at the fifth or sixth infra-
labial. Occasionally the infralabial series may be separated from the chin
shields at a level as far anterior as the fourth infralabial or as far posterior as
the seventh.
Dorsal scales. — The dorsal scales of Trimorphodon are smooth, with
paired apical pits. The scales are arranged in diagonal rows and vary con-
siderably in number. The anterior count at one head length behind the head
ranges from 21-27 rows; 75 percent of the individuals have 22, 23, or 24
rows. The number of scale rows at midbody varies from 17-25; 80 percent of
the snakes have 22 or 23 rows. At one head length in front of the vent, there
are between 14 and 18 rows, and at this level 83.33 percent of the sample have
15 or 16 rows.
The reduction in scale rows is similar to the reduction found in Lepto-
deira (Duellman, 1958:20). The reduction involves changes in the para-
vertebral row. The level (ventral scale) at which reduction in scale rows take
place is highly variable with no geographic consistency.
PATTERN AND COLORATION
Cross bands. — The number of dorsal body bands (body and tail blotches)
in individuals with complete tails ranges from 18-46, though evidence suggests
1970
Geographic Variation of Mexican Lyre Snakes
11
Table 1.
Range and Percent Distribution of Various Head Scales in
the Seven Populations of Trimorphodon
No. of specimens
A
23
B
15
C
14
D
20
E
51
F
25
G
19
2
6.5%
66.6%
75.0%
60.0%
18.0%
12.0%
44.7%
3
87.0
33.4
25.0
35.0
79.0
86.0
55.3
Loreals
4
6.5
—
—
5.0
2.0
2.0
—
5
—
—
—
—
1.0
—
—
1
2.0
2
6.5
13.3
14.3
—
8.9
2.0
15.8
Preoculars
3
93.5
83.3
85.7
97.5
88.1
90.0
84.2
4
—
3.4
—
2.5
3.0
6.0
—
2
19.2
3.5
10.7
7.5
1.9
10.5
Postoculars
3
76.6
96.5
89.3
92.5
94.1
100.0
89.5
4
4.2
—
—
—
4.0
—
—
1
3.5
2
19.6
55.2
39.3
60.0
23.5
24.0
31.6
Primary
3
78.3
41.3
57.2
37.5
71.6
74.0
68.4
Temporals
4
21.0
—
3.5
2.5
4.0
2.0
—
5
—
—
—
—
0.9
—
—
2
—
—
—
12.5
—
—
—
Secondary
3
26.1
44.8
63.0
50.0
25.7
12.0
47.4
Temporals
4
60.9
55.2
37.0
35.0
72.3
86.0
52.6
5
13.0
—
—
2.5
2.0
2.0
—
7
10.0
—
8
60.9
69.0
100.0
72.5
63.9
39.5
57.9
Supralabials
9
37.0
31.0
15.0
29.9
58.1
39.5
10
2.1
—
—
2.5
6.2
2.4
2.6
9
—
—
2.5
—
—
—
10
2.1
3.3
9.5
27.5
2.1
4.6
—
11
28.4
30.0
28.6
50.0
14.9
23.3
26.5
Infralabials
12
47.8
43.4
47.6
20.0
45.8
44.2
44.1
13
21.7
20.0
14.3
—
35.1
25.6
20.6
14
3.3
—
—
2.1
2.3
8.8
12
Contributions in Science
No. 179
that higher numbers do occur (see below). The higher values are found in
samples A, B, C and D while the lower numbers are characteristic of the three
Balsas-Tepalcatepec Basin populations, E, F and G. The numbers of body
bands for the seven populations and for geographically intermediate specimens
are shown in Figure 3.
There is a decrease of dorsal body bands within population A from north
to south. Specimens from Sonora have 27-36, x = 32 (N=ll); specimens
from northern Sinaloa have 29-33, x = 30 (N=14); specimens from central
Sinaloa have 25-31, x = 28 (N=10). Snakes from population B on the aver-
age have more body bands than specimens from population A, and snakes of
population C tend to have more bands than those from B. Specimens that are
geographically intermediate between samples A and B have an intermediate
number of bands. The same is true for two specimens from the area between
samples B and C (Fig. 3, Nos. 8, 9) .
In population C the Hidalgo specimens apparently have more bands than
Figure 3. Statistical analysis of the number of cross bands in populations A through
G of Trimorphodon tan. See legend to Figure 2 for explanation of diagrams and
symbols.
1970
Geographic Variation of Mexican Lyre Snakes
13
do the Tamaulipas specimens. The only Hidalgo specimen with a complete tail
has 43 body bands. Three other snakes from Hidalgo appear to have 40, 42
and 49 bands or more. These three specimens have damaged tails and, for this
reason, are not included in Figure 3. The higher number of bands indicated
for southern specimens in population C may account for the high numbers
found in two of the intermediate specimens between B and C (Fig. 3, Nos. 7,
10). Both of these specimens (FMNH 105193, USNM 11370) are from
localities slightly farther south than most of the localities in either B or C.
Specimens from population D, like population B, have fewer dorsal
blotches than those from population C. Three of the four snakes from locali-
ties intermediate between C and D in eastern Mexico are more similar to D
than to C. The fourth snake (Fig. 3, No. 12), the holotype of Trimorphodon
collaris, is much closer in band number to specimens from sample F.
Snakes from populations E and F average 11 to 13 fewer blotches than
specimens from population D, the sample that is closest geographically. They
are most similar to snakes from population A which is the farthest removed
geographically. In band count, samples E and E are most similar to sample G.
These populations (E, F, G) are significantly different from all of the other
populations. Although populations B and G are in close geographic proximity,
there is no overlap in the number of body bands. In fact these two samples are
separated by a mean difference of 15 dorsal body bands.
The lengths of the bands may vary considerably on a single individual and
among individuals of the same or different populations. In order to evaluate
the differences in band size, the lengths on the middorsal scale row (as indi-
cated by the number of included scales) of three band-interspace-band se-
quences were compared. The first sequence behind the collar (anterior) , a mid-
body sequence, and the last sequence befor the vent (posterior) were used.
These values are presented in Table 2.
In all populations the first dark cross band usually is the longest. The
second cross band is longer on the average than either of the midbody bands,
and they, in turn, always average longer than the posterior body bands. The
mean band length of specimens from population A is greater than that of
specimens from B, and mean band length in B is greater than in C (Table 2).
In all instances the bands on snakes from population D average longer than
bands in C but are shorter than bands on snakes in B.
Bands of specimens from populations E, F and G average longer than
those of any of the other four samples. The only exception is the mean first
band length of sample A which is longer than in sample E but never as long
as in samples E or G. In general, specimens from population G have longer
bands than specimens from E, and specimens from F have longer bands than
specimens from E. The longest mean band lengths at all three body positions
are characteristic of specimens from population G while the shortest band
lengths are always found in population C.
The Range and Mean of Band-Interspace-Band Lengths at Three Middorsal
Positions on Specimens from the Seven Populations
14
Contributions in Science
No. 179
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^ oo ^ vn ON
^ 2 ^ q ^
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^'^^ON^qcd^qrd^ioio
00
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band R 5.5-11.0 4.0-8.0 3. 5-6.0 3.0-6.0 5.5-14.0 6.0-12.5 8.0-14.0
X 7.7 5.8 4.5 4.6 8.6 8.8 11.1
1970
Geographic Variation of Mexican Lyre Snakes
15
There is an average decrease in interspace length from anterior to
posterior in all samples except F and G. In the former the midbody interspaces
average shorter than the posterior interspaces while in the latter the midbody
interspaces average slightly longer than either the anterior or posterior inter-
spaces which are the same (Table 2). Specimens from populations A and B
have the smallest mean interspace lengths. Specimens from population F have
the longest interspace lengths.
There is no consistent number or location of bands which show signs of
splitting or fusion (indicated by light centers) in any population. However,
there is some consistency within certain populations in the extent of the light
central area. For example, in Oaxaca the light areas within the darker bands
are relatively large and diffuse. In Sinaloa the central light areas often form
discrete spots.
Collar length. — The length of the light nuchal collar varies among indi-
viduals from the same population. In some specimens the first body blotch
contacts the posterior margin of the head cap or the parietal scales on the
midline or is separated from them by only half a dorsal scale. Others may have
Figure 4. Statistical analysis of the collar lengths in populations A through G of
Trimorphodon tan. See legend to Figure 2 for explanation of diagrams and symbols.
16
Contributions in Science
No. 179
as many as six dorsal scales separating the first body blotch from the head cap
or the parietal scales. However, the collars among the seven samples have
nearly the same average lengths as indicated by the number of light scales on
the midline (Fig. 4). Only population F has a noticeably higher number of
dorsal scales in the nuchal collar. In addition, snakes from population F have
the greatest interspace length (Table 2). Most of the specimens from inter-
mediate localities have intermediate collar lengths which fall within the range
of variation of the two samples concerned. Notable exceptions, however, are
the collar lengths for the type specimens of T. collaris and T. forbesi which are
plotted between populations C and D (Fig. 4, Nos. 12, 13) . Both of these have
collar lengths most similar to those characteristic of sample F. The type of
Trimorphodon forbesi has four scales, very near the mean for F, while the
type of T. collaris has five and is near the upper end of the range of sample F.
Dorsal head cap. — The pattern of dark and light colors on the head
generally is consistent within a geographic area. Because of this geographic
consistency, a detailed analysis of the head patterns is extremely useful in
determining the relationships among the seven samples. Although individual
head patterns may be very complex (Figs. 5, 6), they can be analyzed in terms
of the following components: snout — light or dark rostral and internasal
scales; presence or absence of a prefrontal bar — light bar extending across the
prefrontal scales with dark anterior and posterior borders; presence or absence
of an interocular (interorbital) bar — a light bar extending between the eyes,
usually including the supraocular scales and the frontal scale; presence or
absence of a parietal mark — a light mark of various design that may or may
not connect to the interorbital bar or the light collar; presence or absence of an
occipital spot — a medial, dark rounded spot located on the posterior portion
of each parietal scale and the adjacent dorsal scales in the collar; posterior
border — the shape of the posterior margin of the dark cap where it is bordered
by the light nuchal collar.
Most snakes from population A have dark snouts (a few specimens from
near Alamos, Sonora have light snouts) and a light prefrontal bar (blending
into the snout of the light snouted forms) that is restricted to the anterior third
of the prefrontal scales (Fig. 5A) . Occasionally (three specimens) the pre-
frontal bar is connected with the interorbital bar by a straight or irregular
medial area. The interocular bar is always present, usually has irregular bor-
ders, and generally is continuous between the orbits (interrupted in two speci-
mens) . A parietal mark is always present but extremely variable in shape and
extent (Fig. 7). Frequently the parietal mark is shaped like an irregular-armed
trident, a bell, a teardrop, or consists of two oblong parts arranged at right
angles to form an inverted T. The parietal mark (connected medially to the
interorbital bar in one specimen) begins at the posterior edge of the frontal
scale and continue posteriorly to connect with the collar (56 percent) . In the
remaining specimens the parietal mark is separated from the light collar. An
1970
Geographic Variation of Mexican Lyre Snakes
17
occipital spot is present in 12 of the 14 specimens with the parietal mark-collar
connection. The posterior border of the head cap may have a single median
indentation (16 specimens), a lateral indentation on either side of a median
posterior extension (five specimens), or it may be nearly straight (three
specimens) .
The two specimens from Ventanas, Durango have complete interorbital
Figure 5. Representative dorsal head cap patterns for specimens of Trimorpliodon
tan from A) Terreros, Sinaloa in population A; B) between Hostotipaquillo and
Magdalena, Jalisco in population B; C) S Ciudad Victoria, Tamaulipas in population
C; D) intermediate locality at Jalapa, Veracruz.
18
Contributions in Science
No. 179
bars and light snouts that include the area of the prefrontal bar (Fig. 7). The
simple parietal marks are restricted to the suture lines between the parietal
scales. The parietal mark is connected to the light collar in one specimen and
separated in the other. An occipital spot is present in the snake wdth the
parietal mark-collar connection.
The specimen from near San Bias, Nayarit has a dark snout, a prefrontal
bar, an interorbital bar, a parietal mark connected to the collar, and an occipital
spot. It is very similar to several Sinaloan specimens.
Most snakes from population B have dark snouts (light in the Zacatecas
specimen and two Michoacan specimens) and distinct light prefrontal bars
(66 percent of individuals) . The interorbital bar is present (faint in KU 67735)
and complete in all specimens except one (UMMZ 118950), where it is inter-
rupted medially. The arms of the parietal mark, which is usually Y-shaped,
edge the frontal scale posteriorly (Fig. 5B; Cope, 1900: fig. 315). The upright
base extending posteriorly along the parietal suture may connect to the collar
(six specimens) or remain separate from it (five specimens). In one specimen
(FMNH 105194) the arms of the parietal mark connect with the interorbital
bar on the supraocular scales forming a triangle (Taylor, 1939: pL 35, fig. 2).
The parietal mark is oblong in three snakes and heart-shaped in one. The
occipital spot is less common than in population A, being found in only two
specimens. The posterior border of the head cap is rounded in two specimens,
forms a straight edge in one, is indented medially in five, and has a medial
posterior projection in the remaining seven snakes.
A snake from La Cumbre, a locality southwest of population B, has a head
pattern very similar to many specimens from B in that it has an interocular bar
and a medial posterior projection of the head cap. The snout is somewhat
lighter than the rest of the head cap and includes the prefrontal bar. The
parietal mark is V-shaped and outlines the frontal scale posteriorly.
Two specimens from Tacicuaro, Michoacan (southeast of population B)
have dark snouts, interorbital bars, and irregular Y-shaped parietal marks that
connect to the light collar. An occipital spot is present in one specimen but
absent from the other (Fig. 7) . Although these two snakes lack prefrontal bars,
they are similar in head pattern to other snakes from population B.
Three specimens from Guanajuato (east of population B) have inter-
ocular bars and Y-shaped parietal marks characteristic of most specimens from
B but resemble the Tacicuaro specimens in lacking the prefrontal bars. The
parietal mark connects to the collar in two specimens and is separate from it
in the third. The snout, including the prefrontal bar, is light in one specimen
and dark in two.
An Aguascalientes specimen from a locality north of population B has a
light snout that includes the prefrontal. In this respect the Aguascalientes snake
is similar to one of the Guanajuato specimens and to the Zacatecas specimen
in population B. The interorbital bar is complete and the parietal mark, which
1970
Geographic Variation of Mexican Lyre Snakes
19
is roughly Y-shaped, does not connect to the light collar. The posterior border
of the head cap is rounded.
Specimens from sample C have a very complex head pattern. Half of
the individuals examined have a light snout that encompasses the prefrontal
area. The other specimens have a dark snout with either a well-developed or
very faint prefrontal bar. Many specimens, especially those with the lighter
snout have a patch of dark color in the center of each prefrontal scale. These
patches may be joined medially or separated by a lighter suture line between
the prefrontal scales. About half of the specimens have a second light bar on
the posterior edges of the prefrontals. The interocular bar is always present and
usually expanded on the frontal scale. In one-third of the specimens the inter-
orbital bar is connected anteriorly to the prefrontal bars and posteriorly to the
parietal mark. In these specimens the dark head color actually is restricted to
six spots, one in the center of each prefrontal scale, one at the anterior edge of
each supraocular and adjacent frontal scale, and one on the posterior part of
each supraocular and adjoining frontal and parietal scales.
In most of the specimens from Tamaulipas and San Luis Potosi, the arms
of the parietal mark form a V and outline the posterior edges of the frontal
scale. The posterior part of the parietal mark bifurcates and encloses a large
occipital spot before joining the collar (Fig. 5C) . In these specimens the
posterior border of the head cap is indented medially.
The parietal marks of the four specimens from Hidalgo differ from the
more northern representatives of sample C. One specimen (LACM 53023) has
a Y-shaped mark outlining the frontal edges and connecting to the collar. It
lacks an occipital spot and has a postero-medially indented head cap. In two
other specimens (AMNH 93432 and ANSP 14770) the parietal mark is lyre-
shaped and not connected to the light collar. The arms of the lyre extend to the
frontal borders, connecting with the interorbital bar on one side in one speci-
men. They do not outline the postero-medial edges of the frontal scale. In both
specimens the occipital spot is incorporated into the head cap, thereby extend-
ing the posterior border of the cap medially. In the fourth specimen (USNM
1 10401 ) the interocular bar extends posteriorly to the tip of the frontal scale.
The parietal mark is fragmented into three parts. On the left side the mark
borders the frontal anteriorly and extends posteriorly nearly to the light collar
forming a shallow, laterally concave arc. On the right side only the anterior
half of the arc is present. The posterior half consists of a light rounded spot on
the posterior part of the right parietal scale, just above the light collar. The
posterior border of the head collar is nearly straight medially but rounded
laterally.
A specimen from Jalapa, Veracruz has a light snout extending over the
anterior half of the prefrontals. Each nasal scale is very faintly pigmented. The
interorbital bar is complete and connects medially to the light snout along the
prefrontal suture. The parietal mark is irregular in shape, restricted to the
20
Contributions in Science
No. 179
parietal suture (Fig. 5D) and connected to the light collar. The occipital spot
is well defined.
The type specimen of Trimorphodon collaris lacks pigment on the inter-
nasal scales. Its interocular bar is broken about the middle of each supraocular,
and it has a more rectangular-shaped parietal mark connected to the collar. In
other respects it is very similar to the Jalapa specimen.
The type specimens of Trimorphodon tan and T. forbesi also are from
localities in the Atlantic versant and generally are similar to the two specimens
mentioned above. The type of T. tail has dark circular marks in the center of
each internasal scale. The light snout is connected medially and laterally with
the prefrontal bar which is also connected with the complete interocular bar
along the suture between the prefrontals. The lighter color of the collar con-
tinues onto the temporal parts of the head connecting with the postero-lateral
portions of the interocular bar. As a result, the dark parietal portion of the
head cap is greatly restricted and completely surrounded by the lighter colora-
tion. The parietal mark is connected to the collar. Posteriorly the head cap is
nearly straight. The occipital spot is present but small. The type of T. forbesi
also has a lighter snout that includes the prefrontal area. The interocular bar is
complete and connects along the prefrontal suture to the lighter snout. The
parietal mark is rounded and broadly joined to the light collar. The occipital
spot is small. The posterior border of the head cap is rounded laterally and in-
dented medially. In general these four specimens from eastern Mexico (Vera-
cruz, Puebla, Oaxaca) are very similar in overall head pattern.
The head patterns of specimens from population D generally are more
uniform than are those from population C. Most of the specimens (75 per-
cent) have a dark snout and lack a prefrontal bar. Two of the five individuals
with a light snout have dark pigment in the central parts of the internasal
scales and a suggestion of a prefrontal bar. In this respect they are similar to the
type of Trimorphodon tau. In several specimens, including light-snouted and
dark-snouted individuals, the edges of the internasal and prefrontal scales are
lighter than the central portions (Fig. 6A). Ten specimens have a light bar
along the suture between the prefrontals. Often this bar is connected to the
interocular bar (50 percent) forming the inverted T-pattern characteristic of
Trimorphodon tau (Cope, 1869: 152) . The interocular bar is always present
and frequently (50 percent) extends laterally to the orbits. In some specimens
(eight of 20) the interocular bar forms a shallow arc, concave posteriorly.
The parietal mark has a simple configuration, rarely touching the frontal scale,
and is connected with the light collar in all but two specimens (Taylor, 1940:
fig. 8). In one specimen (AMNH 1 0093) the mark is irregular in shape (two
lateral arms on each side) and contacts the posterior part of the frontal scale.
An occipital spot is present in only five specimens.
The head cap is indented medially in all but two specimens from sample
D. In these two specimens the parietal mark and the collar do not connect, and
1970
Geographic Variation of Mexican Lyre Snakes
21
thus the head cap extends postero-medially. The white collar extends anteriorly
along the sides of the head in these specimens, together with 16 of the 18
indented head cap forms, giving the head cap a rounded appearance. In two
(Fig. 7) the postero-lateral edges of the cap are nearly straight.
We examined the head patterns of 48 snakes from population E. In only
eight of these does the light snout include the prefrontal bar area. Although the
Figure 6. Representative dorsal head cap patterns for specimens of Trimorphodon
tan from A) 3 mi NE Mitla, Oaxaca in population D; B) Chilpancingo, Guerrero in
population E; C) Coalcoman, Michoacan in population G; D) 4 mi S Autlan, Jalisco
in population G.
22
Contributions in Science
No. 179
snout in these few specimens is somewhat lighter than the rest of the cap, it is
still well pigmented and not the white or very light tan characteristic of certain
light-snouted specimens from populations A and C. None have a defined pre-
frontal bar, although three individuals have a pair of light spots on the anterior
prefrontal scales. Three other specimens have a medial light spot on the pre-
frontal suture. An interorbital bar is lacking in 23 specimens, reduced to a
median frontal spot or to two lateral supraocular spots in 21 specimens, and
complete in four specimens. None has a connection between the interorbital
bar and parietal mark. In all specimens, except four, the parietal mark is con-
nected to the light collar (Fig. 6B) or is absent. Of these four, two have a small
circular spot on the parietal suture, one has a small central and two small lateral
spots, and the fourth has two lateral spots that appear to be remnants of a V-
shaped mark. Of the remaining 44 specimens, eight have no indication of a
parietal mark; the posterior margins of their head caps are rounded laterally
(Fig. 7) . Most of the others have a median extension of the collar connecting
to the parietal mark. Usually the parietal mark or anterior end of the collar
extension is rounded, but occasionally it may be Y- or V-shaped or may narrow
rapidly at its anterior apex near the frontal scale. Occasionally (six specimens)
the collar has a double extension onto the parietal region, and the head cap
extends posteriorly on the midline. The head cap is rounded laterally in nearly
all of these specimens. An occipital spot is present in only nine.
In twelve of 24 specimens from sample F, the snout is lighter than the
rest of the head cap and includes the prefrontal bar area. In this respect they
are similar to the light-snouted individuals of population E. Only two of the
dark snouted specimens have a prefrontal bar. When a complete interorbital
bar is present (three specimens), it is faint. In twelve other snakes, the bar is
fragmented or reduced to one, two, or three circular spots or short bars. There
is not interorbital bar in nine specimens (Taylor, 1940: pi. 52) . Four specimens
lack parietal marks and have a head cap with a rounded posterior margin.
Three specimens have shallow V-shaped parietal marks that outline the frontal
scale and are not connected with the light collar. In these specimens the
posterior edge of the head cap has a single medial indentation, two lateral
indentations, or a postero-medial extension. In the remaining snakes (17) the
parietal mark is connected to the collar, and thereby, the posterior edge of the
head cap is indented medially. The extended collar-parietal mark may be
broadly Y-shaped (Fig. 7), triangular, narrow and restricted to the parietal
suture, or wishbone-shaped. There is a large distinct occipital spot in the collar
extension in most (12 of 17) of these specimens.
The head patterns are remarkably consistent in population G. None, ex-
cept the type of T. fasciolata, have a light snout or show any trace of a pre-
frontal bar. In the type specimen of T. fasciolata the snout and the edges of the
prefrontal scales are somewhat lighter than the rest of the head cap. An inter-
orbital bar is lacking in all specimens (Fig. 6C) except three. In two of these
1970
Geographic Variation of Mexican Lyre Snakes
23
(UMMZ 112516 and the type of T. fasciolata) the supraocular scales have
light centers. The third specimen from Autlan, Jalisco (KU 27191) has two
light circular spots in the interorbital area. Distinct parietal marks are lacking
from all except the Autlan specimen (Fig. 6D) and a specimen from near
Apatzingan (UMMZ 112515). In the latter there are two small spots on the
posterior third of each parietal scale. The posterior margin of the head cap is
indented in one specimen from Apatzingan (FMNH 37079) and the type of
T. fasciolata and shallowly indented in a third (UMMZ 104696) . Occipital
spots, lacking in other specimens, are present in these three snakes. The
posterior margin of the head cap is doubly indented in nine specimens (Fig.
6C), rounded in four, nearly straight in two, and slightly convex in one.
The geographic distribution of the different head patterns is illustrated
in Figure 7.
Coloration. — These snakes have considerable variation in coloration from
sample to sample and within some samples. Because adequate color descrip-
tions and pictures of specimens from many areas are available in the literature
(Taylor, 1939: 364-367; 1940: 474-479; Smith, 1941: 161-167; Peters, 1954:
33; Davis and Dixon, 1957: 24; Duellman, 1961 : 111; Brown and Brown,
Figure 7. Geographic distribution of representative head cap patterns for specimens
of Trimorphodon tan from localities within the seven samples and selected inter-
mediate localities.
24
Contributions in Science
No. 179
1967: 325), no detailed color description will be included here. Instead we
will summarize this information and present it together with color notes on
living specimens from Sinaloa and Puebla.
The ground color of adults ranges from various shades of brown to fawn,
pale tan or gray, usually becoming darker posteriorly. Juveniles often have a
more reddish or grayish ground color than do the adults. The cross bands of
both age groups are darker than the ground color and often yellow-brown or
gray-brown to black. In many specimens the center of each band is lighter. The
edges of each band are usually bordered with dark brown or black. Juveniles
apparently have darker bands than adults from the same area. Laterally,
smaller dark blotches are present in the interspaces of some specimens. The
head cap is brown, gray-brown or black and nearly the same color as the cross
bands. The lips, chin and throat usually are tan or white. The venter may be
tan or creamy white, and often with a reddish or salmon tinge posteriorly. In
some snakes the venter is mottled by ventral extensions of the cross bands and
interspace blotches; in others the venter may be spotted. Some have a nearly
immaculate belly. In some juveniles the ventral surface of the tail may be
nearly black.
The iris may be yellow-brown, gray-brown or grayish-tan.
Discussion
CHARACTER ANALYSIS
Much similarity exists among the seven samples. Some characters indicate
independent, discordant patterns of variation. Even though most characters are
not diagnostic, three are particularly useful in deducing the relationships among
the seven samples: number of ventral plus subcaudal scales, number of dorsal
body bands, and dorsal head cap pattern.
The number of ventral-subcaudal scales decreases from north to south in
samples A through C. The trend continues southward into population D, where
the lowest counts are recorded. Two specimens (Fig. 2, Nos. 11, 14) from
intermediate areas between C and D are more similar to D than to C in this
character, while two other specimens from intermediate areas (Fig. 2, Nos.
12, 13) are more similar to C and F than they are to D. This suggests a con-
nection among the samples through the high river valleys of southeastern
Puebla and Oaxaca. Population G has the highest counts and is the most
northern of the three Balsas-Tepalcatepec Basin samples. In this character
sample G approached sample B, its closest geographic neighbor. The ventral-
subcaudal scale counts will not allow for absolute distinction between speci-
mens from geographically adjacent samples.
An analysis of the number of dorsal cross bands (Fig. 3) gives a slightly
different picture of the relationships among the seven samples. Among popula-
tions A, B, and C there is an increase in number of dorsal blotches from north-
west to southeast. The geographically intermediate specimens generally con-
1970
Geographic Variation of Mexican Lyre Snakes
25
firm this trend. Sample D has fewer bands than does C and thus has a slightly
different relationship to A, B, and C in this character than it did in terms of
ventral-subcaudal scales. In this respect, however, the number of dorsal
blotches of sample D approaches those characteristic of samples E and F. Of
the four specimens from eastern Mexico that are geographically intermediate
among samples C, D and F, three (Fig. 3, Nos. 11, 13, 14) have counts most
similar to the counts of D while the fourth (Fig. 3, No. 12) has a count most
like those of sample F.
In dorsal cross bands E, F and G are closely related to each other and
well differentiated from sample D. The most striking thing about the dorsal
band counts is the difference between samples G and B which do not approach
each other even though the two populations are closely adjacent (Fig. 3). This
is a notable exception to their similarity in the number of ventral-subcaudal
scales.
The same general relationship is present in the seven samples for both
dorsal head patterns and dorsal cross bands. An interocular bar, usually com-
plete, and a discrete parietal mark are always present in specimens from
samples A, B, C, and D. The remnants of an interocular bar are present in 43
of 91 specimens from samples E, F and G. Only seven specimens from the
Balsas-Tepalcatepec samples have a complete interocular bar. A discrete parie-
tal mark, that is one not connected to the collar, is found only in nine of the
91 specimens in samples E, F and G. A prefrontal bar or light snout including
the area of the prefrontal bar is found in most snakes from samples A, B and C.
In sample D a prefrontal bar is usually lacking, as it is in most specimens from
samples E and G. Approximately 50 percent of the specimens from sample F
have light snouts that include the area of the prefrontal bar and thus are some-
what intermediate between samples A, B, C and D, E, G.
In summary, the Balsas-Tepalcatepec Basin samples (E, F and G) have
a darker, more uniform head cap than is found in specimens from samples A,
B, C and D. The Balsas-Tepalcatepec samples generally lack the prefrontal and
interocular bars and the discrete parietal marks characteristic of the other
samples. In general they lack the ornate head marking characteristic of the
other four populations. Except for two snakes (KU 27191 and USNM
110400), there is no indication of any similarity between samples B and G in
characters of head pattern. Some specimens from samples E and F show an
apparent relationship to samples C and D by possessing remnants of an inter-
ocular bar and parietal mark. The head patterns of specimens from areas geo-
graphically intermediate between samples A, B, C and D have intermediate
head patterns.
TYPE SPECIMENS
It seems appropriate to discuss briefly the six type specimens with refer-
ence to our seven populations to clarify our systematic treatment of the various
samples.
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Contributions in Science
No. 179
The type specimen of Trimorphodon tau is a small snake in relatively poor
condition. It was adequately described by Cope (1869) and redescribed by
Taylor (1939). Although from a locality northwest of sample D, in all char-
acteristics, it is typical of specimens from that sample.
The holotype of Trimorphodon upsilon is typical in all respects of speci-
mens from sample B and was included in that unit. Zweifel (1959) noted that
many of the specimens collected by Major probably did not come from
Guadalajara, Jalisco as the locality information indicated but possibly were
from Colima. However, Zweifel did point out that the type of Trimorphodon
upsilon may actually have been from Guadalajara as additional specimens of
this species are known from the same general area. Our analysis supports
Zweifel’s contention that Guadalajara probably is the type locality as originally
stated.
The cotypes of Trimorphodon latifascia from Puebla have the character-
istics typical of Puebla specimens from sample F. However, they were not
included in sample F because the specific type locality is unknown (Peters,
1869).
For purposes of nomenclatural stability and in accord with Article 74,
International Commission on Zoological Nomenclature (1964), we designate
ZMB 6652a, the larger of the two syntypes (588 mm total length), as the
lectotype. The smaller specimen (417 mm total length), ZMB 6652b, is desig-
nated the paralectotype in accord with recommendation 74E. Both specimens
are males and have 211, 209 ventrals, 79, 74 subcaudals, and 21, 22 dorsal body
bands. The snout is slightly lighter than the head cap in both specimens. In one
the interocular bar consists of two short lateral spots and in the other of a
median spot. The posterior border of the head cap is indented medially and
rounded laterally in both. A faint occipital spot is present in the smaller
paralectotype.
The holotype of Trimorphodon collaris, like the type of T. tau, is a
juvenile. Cope’s description (1875) is accurate for the characters listed. Some
confusion exists concerning the type locality and number of specimens (see
Historical Review and Smith, 1941 : 166) . This specimen, although from a
locality somewhat intermediate among samples C, D and F, is typical of speci-
mens from sample F in the important characteristics of ventral plus subcaudal
scales, collar length, and number of dorsal blotches. The band-interspace-band
lengths also are closest to those characteristic of sample F. The head pattern is
similar to the patterns found on specimens from both D and F, although it is
more similar to those from D. We consider the specimen to be most similar to
snakes from sample F.
The type specimen of Trimorphodon fasciolata is a large male collected
from a locality on the northeastern periphery of sample G. The specimen was
adequately described by Smith ( 1941 : 160-162) . In all characteristics except
certain details of head pattern (see previous description), it is typical of rep-
1970
Geographic Variation of Mexican Lyre Snakes
27
resentatives of sample G, in which it was included. These differences in head
pattern emphasize the marginal nature of the locality and may reflect its
proximity to specimens from sample B and the intermediate specimens from
Tacicuaro, Michoacan.
Finally, the type of Trimorphodon forbesi is an exceptionally large male
that is relatively well preserved but badly faded. In general our findings, on
examination of the type, agree well with Smith’s description (1941: 163-165)
with two exceptions. We recorded five more ventrals and two more subcaudals
than listed in the type description. Unfortunately the ventral scales are dam-
aged in some areas and accurate counts are difficult to make. This may account
for the discrepancies. However, we believe our count of 297 scales to be
accurate.
The other point with which we disagree is Smith’s (1941: 164-165) dis-
cussion of and reference to the “pattern reversal” characteristic of Trimorpho-
don forbesi. It is quite obvious that this specimen is faded; apparently the brown
color of the head pattern faded more rapidly than did the dark borders along
the interocular bar and nuchal collar, thus giving the appearance of a light head
pattern with a darker interocular bar and nuchal collar. We cannot accept the
contention that this is anything other than a badly faded but originally nor-
mally colored head pattern. The pattern itself is similar to the pattern of the
type of Trimorphodon tau and other specimens in sample D.
The remaining characters of forbesi that Smith (1941: 165) used to dis-
tinguish it from other species include a divided anterior loreal, the fifth and
sixth labials entering the eye, a faintly marked belly and white subcaudal sur-
face, the number of cross bands, the narrow black borders of the cross bands,
and the length of the cross bands on the mid-dorsal line. All of these charac-
teristics are known from other samples (see previous description of geographic
variation) and therefore are of little value in distinguishing this specimen. In
ventral-subcaudal scales and collar length this specimen is more similar to
sample F. In number of dorsal blotches and in head pattern, it is more similar
to sample D. Certain characteristics of head scales and blotch length are inter-
mediate between those for samples D and F.
Thus, the type of T rimorphodon forbesi has some characters typical of
specimens from sample D, some typical of specimens from sample F, and
some that are intermediate. Because of its intermediate locality and character-
istics, we consider the specimen to represent a population of Trimorphodon
that apparently bridges the gap between samples D and F.
In order to evaluate the usefulness of the three characters, ventral-sub-
caudal scales, body bands and head cap pattern, as indications of phylogenetic
relationship, we attempted to correlate their geographic variation with various
environmental parameters. Klauber (1941 ) showed that desert specimens con-
sistently tend to have more ventral scales than coastal specimens of the same
species. He also found that specimens of Trimorphodon vandenburghi from
28
Contributions in Science
No. 179
the desert had more dorsal blotches than those along the coast. Klauber at-
tributed these disparities to differences in humidity, and possibly temperature.
Fox (1948) showed how different temperature regimes can influence scutella-
tion in developing embryos of Thamnophis. Gravid females from the same
natural population kept at cooler temperatures give birth to young with
significantly fewer ventral and subcaudal scales than young from females
maintained at warmer temperatures. Stebbins (1949) suggested that physical
factors of humidity, temperature and light are important considerations deter-
mining the coloration and pattern of the salamander Ensatina.
In our analysis we could find no consistent correlation between environ-
mental factors of temperature, rainfall or elevation and the geographic varia-
tion in ventral-subcaudal scales, dorsal cross bands or head pattern. In some
samples there appeared to be a positive correlation between ventral-subcaudal
scales and temperature; in other samples the same character indicated an
inverse relationship to mean temperature. These same results generally were
found in other comparisons. Unfortunately, the lack of precise microenviron-
mental information for many areas prevents detailed analysis of the possible
correlation between other environmental factors and morphology.
Band lengths (Table 2) and band numbers (Fig. 3) have an inverse re-
lationship. As the average band length increases, the number of bands de-
creases, and the snakes have a greater dark to light ratio. This is found in
samples E, F, and G and is exemplified also by their darker head pattern. The
Balsas-Tepalcatepec Basin is characterized by both high mean annual tempera-
tures and high mean annual rainfall (Vivo Escoto, 1964). This suggests that a
detailed analysis might show a correlation between mean annual temperature,
rainfall, and/or light intensity and the number of dorsal bands and head
pattern. Details of vegetation type and density would also have to be considered.
Variation in band numbers among the samples may be the result of
splitting or fusion of some of the cross bands. For example, there is an average
increase of five body bands from population A to B, but a decrease of about
15 scale lengths (difference between totals of six band lengths in Table 2).
Although the light centers of many bands suggest splitting or fusion, there is no
consistent location or number of bands which have light centers within any
population. In addition, we were unable to derive the various band patterns
from one another. It seems unlikely, therefore, that the light centers charac-
teristic of many cross bands are indicative of fusion or division, a possible
means of decreasing or increasing the band number. Differences in cross band
numbers and lengths may represent adaptive responses to different selective
pressures that increase or decrease the overall ratios between dark and light
areas on the snakes.
SYSTEMATIC TREATMENT
Our data indicate overall similarities among the seven samples and the
type specimens in nearly all characteristics examined. The analyses reveal
1970
Geographic Variation of Mexican Lyre Snakes
29
a geographic trend towards a southern reduction in ventral-subcaudal scales
among samples A, B, C and D and among samples E, F and G, and also point
to the similarities among samples A, B, C and D and samples E, F and G in
number of body bands and head pattern. Specimens with characteristics that
are intermediate between these two groups indicate that only a single species
is involved. Therefore, we refer our seven samples, their intermediates, and
the types of Trimorphodon tau, upsilon, collaris, Jatifascia, fasciolata and
forbesi to one species. After careful consideration and consultation, we con-
clude that Gunther’s (1895) use of Trimorphodon upsilon did not constitute
the action of the first revisor as defined in Article 24, International Code of
Zoological Nomenclature. Therefore, the name Trimorphodon tau is applicable
because of the action of Smith and Darling (1952) who gave priority to tau
rather than upsilon, by their action as first revisor.
TRIMORPHODON TAU Cope
Trimorphodon tau Cope, 1869, Proc. Amer. Phil. Soc., 11, p. 152 (type
description; type locality “western part of the Isthmus of Tehuantepec, Mex-
ico,” in error); 1875, J. Acad. Natur. Sci. Philadelphia, ser. 2, 8, p. 131. Sumi-
chrast, 1892, La Naturaleza, 6(2), p. 44 (type from Quiotepec, between Tehu-
acan and Oaxaca). Cope, 1886, Proc. Amer. Phil. Soc., 23, p. 286; 1887, Bull.
U.S. Natl. Mus., 32 p. 68; 1892, Proc. U.S. Natl. Mus., 14* p. 678. Gunther,
1895, Biologia Centrali-Americana, Reptilia, p. 174. Boulenger, 1896, Cata-
logue of the snakes in the British Museum, vol. 3, p. 56. Mocquard, 1899, Bull.
Soc. Phil., ser. 9, l,p. 157. Cope, 1900, Rep. U.S. Natl. Mus. for 1898, p. 1101.
Gadow, 1905, Proc. Zool. Soc. London, 1905, p. 224. Mocquard, 1908, in
Dumeril and Bocourt, Mission Scientifique au Mexique et dans I’Amerique
Centrale, 3, pp. 908, 912. do Amaral, 1929, Mem. Inst. Butantan, 4, p. 202.
Taylor, 1939, Univ. Kansas, Sci. Bull., 25, pp. 366-67; 1940, Univ. Kansas Sci.
Bull., 26, pp. 474-77, fig. 8, plate 51. Smith, 1941, Proc. U.S. Natl. Mus., 91,
pp. 166-67; 1943, Proc. U.S. Natl. Mus., 93, p. 494. Smith and Taylor, 1945,
Bull. U.S. Natl. Mus., 187, p. 148. Taylor, 1949, Univ. Kansas Sci. Bull., 33,
pp. 210-11. Davis and Dixon, 1957, Southwestern Natur., 2, p. 24. Martin,
1958, Misc. Publ. Mus. Zool. Univ. Michigan, 101, p. 74. Duellman, 1961,
Univ. Kansas Publ. Mus. Nat. Hist., 15, p. 112; 1965, Univ. Kansas Publ. Mus.
Nat. Hist., 15, pp. 655, 657, 660-61, 666, 685. Lynch and Smith, 1965, Herpe-
tologica, 21, p. 168. Brown and Brown, 1967, Texas J. Sci., 29, p. 325.
Eteirodipsas biscutata (in part), Jan, 1863, Elenco sistematico degli
ofidi . . ., p. 105. Jan and Sordelli, 1872, Iconographic generate des ophidiens,
vol. 3, livr. 39, plate 1, fig. 3. Cope, 1892, Proc. U.S. Natl. Mus., 14, p. 679
(footnote). Gunther, 1895, Biologia Centrali-Americana, Reptilia, p. 175.
Trimorphodon upsilon Cope, 1869, Proc. Amer. Phil. Soc., 11, p. 152
(type description; type locality “Guadalaxara, West Mexico”); 1875, J. Acad.
30
Contributions in Science
No. 179
Natur. Sci. Philadelphia, ser. 2, 8, p. 131; 1886, Proc. Amer. Phil. Soc., 23,
p. 286; 1887, Bull. U.S. Natl. Mus., 32, p. 68; 1892, Proc. U.S. Natl. Mus., 14,
p. 678. Gunther, 1895, Biologia Centrali-Americana, Reptilia, pp. 174-75.
Boulenger, 1896, Catalogue of the snakes in the British Museum, vol. 3, p. 55.
Mocquard, 1899, Bull. Soc. Phil., ser. 9, 1, p. 157. Cope, 1900, Rep. U.S. Natl.
Mus. for 1898, pp. 1 104-1105, fig. 315. Gadow, 1905, Proc. Zool. Soc. London,
1905, pp. 196, 231, 233; 1910, Zool. Jahrbuch, 29, pp. 697, 701. Mocquard,
1908, in Dumeril and Bocourt, Mission Scientifique an Mexique et dans
I’Amerique Centrale, 3, pp. 908, 910-11. do Amaral, 1929, Mem. Inst. Butan-
tan, 4, p. 202. Taylor, 1939, Univ. Kansas Sci. Bull., 25, pp. 365-66, plate 35,
fig. 2. Smith, 1941, Proc. U.S. Natl. Mus., 91, pp. 162-163; 1943, Proc. U.S.
Natl. Mus., 93, p. 494. Smith and Taylor, 1945, Bull. U.S. Natl. Mus., 187,
p. 148. Zweifel and Norris, 1955, Amer. Midland Natur., 54, p. 245. Duellman,
1961, Univ. Kansas Publ. Mus. Nat. Hist., 15, p. 112.
Dips as biscutatus var. latifascia Peters, 1869, Monatsber. Akad. Wiss.
Berlin, p. 877 (type description; type locality, Puebla, Mexico) .
Trimorphodon collaris Cope, 1875, J. Acad. Natur. Sci. Philadelphia, ser.
2, 8, p. 131 (type description; type locality “Orizaba, Vera Cruz”). Sumichrast,
1882, La Naturaleza 6(2), p. 44 (type(s) came from Tuxpango, near Ori-
zaba) ; Cope, 1886, Proc. Amer. Phil. Soc., 23, p. 286; 1887, Bull. U.S. Natl.
Mus., 32, p. 68; 1892, Proc. U.S. Natl. Mus., 14, p. 679. Gunther, 1895,
Biologia Centrali-Americana, Reptilia, p. 174. Cope, 1900, Rep. U.S. Natl.
Mus. for 1898, p. 1101. Taylor, 1939, Univ. Kansas Sci. Bull., 25, pp. 364-65.
Smith, 1941, Proc. U.S. Natl. Mus., 91, pp. 165-66; 1943, Proc. U.S. Natl.
Mus., 93, p. 493. Smith and Taylor, 1945, Bull. U.S. Natl. Mus., 187, p. 147.
Sibon biscutatus, Garman, 1883, Mem. Mus. Comp. Zool. Harvard Univ.,
8, pp. 16, 134 (part).
Sibon biscutatum var. latifasciatum, Garman, 1883, Mem. Mus. Comp.
Zool. Harvard Univ., 8, pp. 16-17, 134.
Sibon upsilon, Garman, 1883, Mem. Mus. Comp. Zool. Harvard Univ.,
8, p. 134.
Trimorphodon biscutatus, Gunther, 1895, Biologia Centrali-Americana,
Reptila, p. 174 (part).
Trimorphodon upsilon var. collaris, Mocquard, 1908, in Dumeril and
Bocourt, Mission Scientifique au Mexique et dans I’Amerique Centrale, 3,
plate 74, fig. 2, a, b.
Trimorphodon latifascia, Taylor, 1939, Univ. Kansas Sci. Bull. 25, pp.
364-65, plate 36, fig. 2; 1940, Univ. Kansas Sci. Bull., 26, p. 479, plate 52.
Smith, 1941, Proc. U.S. Natl. Mus., 91, p. 160. Smith and Taylor, 1945, Bull.
U.S. Natl. Mus., 187, p. 148. Schmidt and Shannon, 1947, Fieldiana Zool. Chi-
cago Nat. Hist. Mus., 31, p. 83. Hall, 1951, Univ. Kansas Sci. Bull., 34, pp.
207-08. Davis and Smith, 1953, Herpetologica, 8, p. 140. Peters, 1954, Occ.
Papers Mus. Zool. Univ. Michigan, 554, p. 32. Davis and Dixon, 1957, South-
1970
Geographic Variation of Mexican Lyre Snakes
31
western Natur., 2, p. 23. Webb and Fugler, 1957, Herpetologica, 13, p. 33.
Duellman, 1958, Occ. Papers Mus. Zool. Univ. Michigan 589, p. 12. Davis
and Dixon, 1959, Proc. Biol. Soc. Washington, 72, p. 88. Duellman, 1961,
Univ. Kansas Publ. Mus. Natur. Hist., 15, pp. 111-12. Fouquette and Ross-
man, 1963, Herpetologica, 19, p. 199. Duellman, 1965, Univ. Kansas Publ.
Mus. Nat. Hist., 15, pp. 655-56, 659, 680, 699.
Trimorphodon fasciolata Smith, 1941, Proc. U.S. Natl. Mus., (type de-
scription; type locality “near Zararacua [sic] Falls, 6 kilometers southeast of
Uruapan, Michoacan.”) 91, pp. 160-62; 1943, Proc. U.S. Natl. Mus., 93, p.
493. Smith and Taylor, 1945, Bull. U.S. Natl. Mus., 187, p. 147. Schmidt and
Shannon, 1947, Fieldiana Zool. Chicago Nat. Hist. Mus., 31, p. 83. Davis and
Smith, 1953, Herpetologica, 8, p. 140. Peters, 1954. Occ. Papers Mus. Zool.
Univ. Michigan, 554, pp. 32-34. Davis and Dixon, 1957, Southwestern Natur.
2, p. 23.
Trimorphodon forbesi Smith, 1941, Proc. U.S. Natl. Mus., 91, pp. 163-65
(type description; type locality “from San Diego [about 5 mites south of
Tehuacan], Puebla”); 1943, Proc. U.S. Natl. Mus., 93, p. 493. Smith and
Taylor, 1945, Bull. U.S. Natl. Mus., 187, p. 147.
Trimorphodon tau tau, Smith and Darling, 1952, Herpetologica, 8, p. 85.
Trimorphodon tau upsilon, Smith and Darling, 1952, Herpetologica, 8,
p. 85. Zweifel, 1959, Amer. Mus. Novitates, 1949, pp. 3-4, 6-8.
Description. — A species of Trimorphodon with 201-231 ventral and 61-
85 subcaudal scales in males; 210-243 ventrals and 55-80 subcaudals in fe-
males; 265-319 total ventral-subcaudal scales; a divided anal scale; usually 3
(2-5) loreal scales, 3 (1-4) preocular, 3 (2-4) postocular, 3 (1-5) primary
temporal, 4 (2-5) secondary temporal, 8 (7-10) supralabial with the fourth
and fifth (third-sixth) bordering the eye, and 12 (9-14) infralabial scales
usually separated from the chin shields at the sixth (fourth-seventh) infra-
labial; dorsal scales usually in 22, 23, or 24 (21-27) rows at one head length
behind the head, 22 or 23 (17-25) rows at midbody, and 15 or 16 (14-18) rows
at one head length in front of the anus.
The snakes are usually gray, tan or light brown in ground color and
crossed by 18-46 body bands that usually reach the edges of the ventral and
subcaudal scales. The cross bands are various shades of brown or black and
often darkest along their margins. Most dark cross bands have lighter central
areas that approach the ground color. These central areas may be large and
diffuse or concentrated into discrete spots. They suggest the splitting of one
band or fusion of two bands. The cross bands decrease in length on the midline
from anterior to posterior. Often the first cross band is considerably larger
than the second. The cross bands are widest dorsally and taper to a broad or
narrow base laterally. Laterally, small, irregular interband spots may be
present. Frequently the body bands continue onto the abdominal scales giving
32
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No. 179
the ventral surface an irregular blotched pattern. However, in some specimens
the abdominal scales are irregularly spotted or nearly immaculate.
The head pattern is extremely variable (see discussion). It consists of a
dark head cap extending from the snout to the posterior parts of the parietals
where it usually is separated from the first cross band by a light nuchal collar.
In some specimens the snout is light. There may be a prefrontal bar, an inter-
ocular bar, a parietal mark and an occipital spot or any combination of these
components. In some specimens most or all of the head cap components may
be lacking, and the head is uniformly dark brown.
The nuchal collar sometimes extends medially onto the parietal area to
connect with the parietal mark. In this instance the anterior border of the
collar is chevron-shaped. Often the nuchal collar is straight or slightly indented
anteromedially. The posterior border of the collar is nearly always straight. The
first body blotch occasionally contacts the head cap or may be separated from
it one-half to six dorsal scales. The collar is usually three scales long on the
midline.
Subspecies. — The numbers of cross bands, the cross band lengths, and the
head pattern indicate that gene flow between samples B and G is limited. The
geographic barrier of the Transverse Volcanic Range separating these two
samples may be partly or wholly responsible for this apparent restriction of
gene flow. Presumably these two populations have come into secondary con-
tact after experiencing separate histories. In our opinion, these two samples
do not represent a continuum that was split into two components by the uplift
of the Transverse Volcanic Range. It seems more likely that population G
evolved from ancestors that moved into the Tepalcatepec Basin from the
south after the Transverse Volcanic Range reached its present height in Jalisco
and Michoacan (see discussion below). The similarities between the Balsas-
Tepalcatepec samples E, F and G and their differences from A, B, C and D
support this view. The latter four samples are similar in several characteristics
and probably are representative of a continuum, as indicated by specimens
from intermediate localities. There appears to have been some genetic exchange
between the two groups through samples F and D in some of the river valleys
of southeastern Puebla, northeastern Oaxaca and eastern Veracruz.
The similarities and differences between A B, C and D and between E, F
and G have been thoroughly discussed. Consideration of these points, com-
bined with the apparent geographic barriers between these two groups, at this
time leads us to recognize them as subspecies. The specimens representative of
samples A, B, C and D and their intermediates are referred to Trimorphodon
tau tau. The specimens from samples E, F and G in the Balsas-Tepalcatepec
Basin are referred to Trimorphodon tau latifascia (new combination).
Distribution and ecology. — Trimorphodon tau is widely distributed along
the coastal slopes and foothills of the Sierra Madre Occidental and Sierra
Madre Oriental (Fig. 1 ) . Occasaional specimens have been taken on the coastal
1970
Geographic Variation of Mexican Lyre Snakes
33
plains. The species is found in suitable habitats to the north of the Transverse
Volcanic Range and the high southern portions of the Mexican Plateau.
Trimorphodon tau has also been recorded from scattered localities to the east
and south of the Mexican Plateau, from the Valley of Oaxaca south and east of
the Sierra Madre del Sur. The species is widely distributed in the foothills of
the Balsas and Tepalcatepec Basins. Although the species is reported from
localities ranging from 100 to 2600 meters in elevation, most of the range of
the species lies between 1000 and 2100 meters.
The lack of adequate material from localities between samples A and B,
between samples D and E, and between samples E and G probably reflects the
inaccessibility of these areas as well as inadequate sampling of suitable habitat.
Only four specimens are known from the eastern versant between samples C
and D. We suspect that local climatic factors (high annual precipitation and
abundant cloud cover) probably restrict the distribution of Trimorphodon to
drier, less accessible habitats in this area.
Trimorphodon tau is a nocturnal species that occupies semi-arid to season-
ally dry habitats. Specimens have been collected in arid tropical scrub, thorn
woodland, tropical deciduous forest, mesquite grassland and dry pine-oak
woodland (vegetation classifications follow Leopold, 1950). The species is
terrestrial and commonly found in rocky areas of irregular relief. Individuals
have been collected from beneath rocks, in holes and under bark on fence posts
and dead trees during the day and in a flooded rice field at night. Most speci-
mens were found on roads at night. The species feeds primarily on lizards, al-
though frogs and small mammals probably are eaten also. Unidentifiable
species of Sceloporus and Cnemidophorus were found in specimens from
Guerrero, Michoacan and Oaxaca.
Eggs and young. — Because very little is known concerning the eggs and
young of oviparous snakes, the following data for Trimorphodon tau are ap-
propriate.
On July 18, 1966 a large female Trimorphodon tau (MVZ 81354) was
collected 26 miles NW of Huajapan de Leon on Mexico Highway 190 in
Puebla (Sample E) by J. L. LaPointe. Sometime between the date of capture
and July 25, this snake deposited seven eggs. The eggs were incubated follow-
ing the technique outlined by Zweifel (1961: 112-113). On September 25, one
of the eggs was opened and found to be developing. On October 10, the eggs
averaged 26.1 mm in length, 16.8 mm in width and 4.0 gm in weight. Eive of
the remaining six eggs hatched between October 29 and November 1, at least
96 days after laying. The hatchlings made from 2-7 (x = 5.4) longitudinal
slits in the egg shell to escape. The sixth egg did not hatch and was found, on
November 4, to contain a well developed, but dead, female (MVZ 81361).
Pertinent information for the mother and five hatchlings are presented in
Table 3.
The hatchling snakes are typical of sample E in the characteristics studied.
34
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No. 179
Table 3.
Data for Adult Female and Five Offspring of
Trimorphodon tau from Puebla, Mexico
Specimen No. (MVZ) :
Sex
Weight - gms.
Total length - mm.
Snout-vent length - mm.
tail length
total length
Ventrals
Subcaudals
Ventrals & Subcaudals
Umbilical Scar *
Body Bands
Supralabials
Infralabials
First ventral scale with scar
81354
81359
81360
Ad. $
$
$
—
2.73
2.95
752
199
207
633
168
176
15.8
15.6
14.3
217
218
216
69
67
68
217
218
216
—
186
182
28
24
?
8-9
8-8
9-9
13-13
13-12
13-13
81356
81357
81358
$
S
2.39
2.86
2.86
199
213
219
165
174
179
17.1
18.3
18.3
195
205
203
80
84
78
195
205
203
170
174
177
22
22
24
8-8
9-8
8-8
11-11
13-12
12-12
Five hatchlings have 275-289 (x = 282.8) total ventral-subcaudal scales. These
counts are very similar to counts from other Puebla specimens from the same
general area (275-290, x = 284.75, N = 8). The same similarities exist in
comparing the total body band counts. Four hatchlings have 22-24 (x = 23.0)
dorsal cross bands while other Puebla specimens have 19-26 (x = 22.3, N = 9)
cross bands. The hatchlings also agree with other Puebla specimens of Tri-
morphodon in their supralabial and infralabial counts (Table 2).
Evolutionary History
Some brief comments concerning speciation in the genus Trimorphodon
seem appropriate. Because no fossil remains are known, our interpretation of
the evolutionary history of the species is based primarily on an analysis of the
distributions and relationships of living forms correlated with the geohistory
and paleogeography of the area.
Savage (1966) considered snakes of the genus Trimorphodon to be rep-
resentative of the Middle American Element. Duellman (1958, 1966) sug-
gested that Trimorphodon diverged early from the evolutionary stock that gave
rise to Hypsiglena, Leptodeira, Eridiphas and Cryophis. We concur, and sug-
gest that the ancestor of Trimorphodon diverged early in the Cenozoic and
probably occupied areas of relatively low relief in northern and central Mexico.
Axelrod (1958) has indicated that most of this region was dominated by a
Neotropical Tertiary Geoflora. Based on indirect evidence several authors have
suggested that a broad ecotone of Arcto-Tertiary Geoflora extended into the
mountains of Mexico at this time (Axelrod, 1960; Brame and Wake 1963;
Savage, 1966).
1970
Geographic Variation of Mexican Lyre Snakes
35
As a result of the general cooling trend in the early Tertiary, the northern
borders of the tropics began to move towards the equator. In areas of north-
western Mexico and the southwestern United States, elements of a Madro-
Tertiary Geoflora, that developed in situ, began to spread. The Miocene was a
period of major mountain building and general uplift (Maldonado-Koerdell,
1964). The Mexican Plateau, the Sierra Madre Occidental, the Sierra Madre
Oriental, and the Sierra Madre del Sur were uplifted to their present height
beginning in the Miocene and continuing to the present. This period of orogeny
initiated major volcanic action, especially during the Pliocene and Recent, that
formed the Transverse Volcanic Range. By the end of Miocene, in response to
the gradual Tertiary temperature depression and increasing aridity, much of
the tropical vegetation, except the more xeric type, was eliminated from cen-
tral and western Mexico. In northern Mexico the derivatives of the Madro-
Tertiary Geoflora were segregating into their component parts in the late
Cenozoic. The mixed coniferous and deciduous derivatives of the Arco-
Tertiary Geoflora were found throughout most of the mountainous areas of
Mexico.
We suggest that the ancestral Trimorphodon stock gave rise to three
distinct lineages, each of which was markedly affected by the continuing
trends of cooling and aridity and by the late Tertiary period of mountain
building and volcanic activity.
The first lineage probably occupied much of central Mexico and gave
rise to Trimorphodon tail. As the climatic changes continued, this species
adapted to dry habitats throughout much of the present Mexican Plateau per-
haps as far south as Oaxaca. Tamayo and West (1964) suggested that much
of this area, originally drained by the Lerma River included the lake basins
drained by the present Rio Lerma as far southeast as the Valley of Mexico.
With the late Tertiary volcanic activity that gave rise to the Mesa Central
(West, 1964) and the Transverse Volcanic Range and the resultant cooler
temperatures, this ancestral species was eliminated from the central part of its
range and restricted to a horseshoe pattern of distribution north, east and south
of the Mesa Central. One segment of this population gave rise to the subspecies
latifascia in the eastern Balsas Basin. Subsequently, when conditions were
suitable, the Balsas population apparently expanded into the Tepalcatepec
drainage system and approached the range of the northern segment of the
original population. These two ends of the horseshoe are now separated by the
western portions of the Transverse Volcanic Range. At the same time that the
Balsas-Tepalcatepec populations were expanding, the population ancestral to
the subspecies tau was expanding northward along the slopes and foothills of
the Sierra Madre Occidental and Sierra Madre Oriental.
The second major lineage of Trimorphodon evolved in association with
the developing Madro-Tertiary Geofloral derivatives and gave rise to the
species found in the arid lowlands of the southwestern United States and north-
36
Contributions in Science
No. 179
western Mexico today. We suspect that Trimorphodon vandenburghi, T.
lambda and T. lyrophanes, if indeed they are distinct species, were derived from
this second major lineage. Trimorphodon vilkinsonii, today found in eastern
New Mexico, western Texas and adjacent parts of Mexico, likely was derived
from the ancestral lyrophanes stock and isolated east of the Rocky Mountain-
Sierra Madre Occidental mountain system, where it is found today in moun-
tains and foothills on the margin of the Chihuahuan Desert.
The third lineage gave rise to the species T. biscutatus and probably
evolved in a xeric tropical lowland environment in the Pacific coastal lowlands
of southern Mexico and northern Central America. Savage (1966) referred to
this species in his discussion of the history of the Western Mesoamerican
Complex. Trimorphodon biscutatus subsequently dispersed northward along
the developing Pacific lowland route as far north as southern Jalisco.
Areas of sympatry or near sympatry between Trimorphodon biscutatus
and T. lambda occur in the southwestern corner of Jalisco. Trimorphodon tau
is sympatric with Trimorphodon biscutatus at Apatzingan, Michoacan and
Huajintlan, Morelos and with Trimorphodon lambda at Terreros, Sinaloa and
about 10 miles west of Alamos, Sonora. Although T. tau has been reported
from Chihuahua, Chihuahua (Cope 1900) near the type locality of Trimor-
phodon vilkinsonii, this record needs to be verified.
Acknowledgments
We would like to thank the following people and institutions for allowing
us to examine material in their care: Charles M. Bogert and Richard G. Zwei-
fel, American Museum of Natural History, New York (AMNH) ; Edmond V.
Malnate and James E. Bohlke, Academy of Natural Sciences, Philadelphia
(ANSP); M. J. Fouquette, Arizona State University, Tempe (ASU); Alice
G. C. Grandison, British Museum (Natural History), London (BM); Richard
B. Loomis, California State College, Long Beach (CSCLB); Robert F. Inger
and Hymen Marx, Field Museum of Natural History, Chicago (FMNH);
William E. Duellman, Museum of Natural History, University of Kansas,
Lawrence (KU); John W. Wright, Los Angeles County Museum of Natural
History, Los Angeles (LACM); Douglas A. Rossman, Louisiana State Uni-
versity, Museum of Zoology (LSUMZ); Ernest E. Williams, Museum of
Comparative Zoology, Harvard University, Cambridge (MCZ); Walter W.
Dalquest, Midwestern University, Wichita Falls, Texas (MU); Robert C.
Stebbins, Museum of Vertebrate Zoology, University of California, Berkeley
(MVZ); George S. Myers, Museum of Natural History, Stanford University,
Palo Alto (SU — collections now housed at California Academy of Sciences,
San Francisco); James R. Dixon, Texas Cooperative Wildlife Collection,
Texas A & M University, College Station (TCWC); Charles H. Lowe, Uni-
versity of Arizona, Tucson (UAZ) ; Walter Auffenberg, Florida State Museum,
University of Florida, Gainesville (UF); Hobart M. Smith, University of
1970
Geographic Variation of Mexican Lyre Snakes
37
Illinois Museum of Natural History, Urbana (UIMNH); Charles F. Walker,
University of Michigan Museum of Zoology, Ann Arbor (UMMZ) ; James A.
Peters, United States National Museum, Washington, D.C. (USNM); and
Gunther Peters, Zoologisches Museum Humboldt-Universitat, Berlin (ZMB).
Robert L. Bezy and C. Jay Cole, and Richard D. Worthington also provided
specimens for study. The information on eggs and hatchlings was made avail-
able by Joseph L. LaPointe, Ray B. Huey and Ted Papenfuss.
Evie Templeton illustrated the head patterns in Figures 5, 6, and 7.
Mercedes McDiarmid and Richard D. Worthington provided technical assist-
ance with the statistical analysis. Allene J. Brown typed the manuscript. David
B. Wake and Hymen Marx read the manuscript in final form and made several
helpful suggestions. Fred F. Gehlbach kindly relinquished his interest in this
group of Trimorphodon. Carl L. Hubbs, W. I. Follett and Lillian Dempster
offered their advice concerning the proper use of the name Trimorphodon tau.
The Society of Sigma Xi provided financial assistance to the first author
for field work in Sinaloa in 1962.
To all of these people and institutions we extend our thanks.
Specimens Examined
MEXICO
Aquascalintes: 7 mi S Rincon de Romos (UIMNH 27566).
Colima: Colima (AMNH 12777); 4.3 mi SW Colima, 575' (UMMZ 114479).
Durango: Ventanas (BM 83.4.16.68 and 69).
Guanajuato: Guanajuato (SU 4412, USNM 11370), Mt. Cubilete (AMNH 93433).
Guerrero: 15 mi NE Acapulco (LACM 7114); Acahuizotla (TCWC 22142); 3 mi
WSW Axixintla (UMMZ 126541); Chilpancingo (FMNH 38412-32; KU 24100-03;
MCZ 33657-58, 33663-5; UIMNH 34991-93); near Chilpancingo (UMMZ 85769-
74); vicinity of Chilpancingo (AMNH 72522-24); 4 mi W Chilpancingo, 5800'
(TCWC 9575); 13 m S Chilpancingo, 3750' (KU 67733); 7 mi E Chilpancingo
(UIMNH 19139); 40 mi N Chilpancingo (LACM 53026); 25 mi NNE Iguala,
3800' (TCWC 12608); Ojito de Agua, 2.5 mi S Almolonga, 5600' (TCWC 11602-
03); Ojito de Agua, 3 mi S Almolonga, 5400' (TCWC 11601); Omilteme, Sierra de
Burro Mtns. (MCZ 42677); Palo Blanco, 4800' (TCWC 9574); 15 mi SE Tonatico
(UMMZ 126542); Taxco (MCZ 33901).
Hidalgo: 10 km N Jacala (USNM 1 10401 ); Zacualtipan (ANSP 14770); 5.6 mi N
of road to Zimapan on Hwy 85 (LACM 53023 ) ; 5 mi S Zimapan (AMNH 93432).
Jalisco: 2 mi ENE Acatlan, 5300' (KU 67735) ; 4 mi S Autlan (KU 27191); 2 mi E
Bolanos, 3500' (KU 91428); 1 mi NE Contla, 3800' (KU 80762-63); La Cumbre de
los Arrastrados, 8500' fTalpa Mascota] (BM 92.10.31.63); Guadalajara (MCZ
46891; USNM 31358-type of upsilon); between Hostotipaquillo and Magdalena
(AMNH 19582) ; 10.4 km NNW Ixtlahuacan del Rio (KU 102986) ; Jamay (AMNH
38
Contributions in Science
No. 179
19841); near Magdalena, 1300 m (FMNH 105194); 1 mi NW El Molino, 5100'
(AMNH 96652).
Mexico: Ixtapan de la Sal, 5250' (AMNH 71360).
Miahoacan: Apatzingan, 1000' (FMNH 39078-79); 7.6 mi E Apatzingan, 1600'
(UMMZ 112514); 14 mi E Apatzingan, 1700' (UMMZ 112516); 17.6 mi E Apat-
zingan, 1600' (UMMZ 112515); 3.9 mi S Apatzingan (UlMNH 73730); Coal-
coman, 945 m (UMMZ 104696-97); 6 mi E Emiliano Zapata, 5350' (UMMZ
118950); 7 mi E Emiliano Zapata, 5400' (UMMZ 118949); 4.3 mi N Lombardia,
2850' (UMMZ 118951); 8.7 mi S Lombardia (UMMZ 124039) ; Nueva Italia, 1250'
(UMMZ 118952); 43 mi S Nueva Italia on Playa Azul Road, 1000' (UAZ 27060);
Tacicuaro, 2000 m (FMNH 105193; UIMNH 19138); 5.5 m W Tangamandapio,
5800' (UMMZ 118948); 1.4 mi N Rio Tepalcatepec (UIMNH 73729); Tzararacua
Falls, 6 km SE Uruapan (USNM 110400-type of fasciolata).
Morelos: Alpuyeca, 3500' (TCWC 4127); 4 mi S Alpuyeca, 3600' (UMMZ 114478);
Campo Agricola, Progreso (UIMNH 26073); camp near Cuernavaca (UIMNH
19135); Huajintlan (UIMNH 19137); km 128, near Puente de Ixtla (FMNH
105100); 12 mi S Puente de Ixtla (UIMNH 19134); Tepoztlan (UIMNH 19136);
6 mi W Yautepec (TCWC 7390).
Nayarit: Barranquitas (AMNH 75585); 5 mi SE Ixtlan del Rio (TCWC 12609); N
of San Bias (LACM 53025).
Oaxaca: Juquila Mixes [^zXuquila Mixes] (AMNH 100649); 3 mi NE Mitla, 5800'
(AMNH 97985); ca. 2 mi W Mitla, La Fortaleza (AMNH 89629); 17 mi SE
Nochixtlan, 6900' (TCWC 12640); Oaxaca (FMNH 105305, 105350); 21 mi SE
Oaxaca, 5350' (UMMZ 112522); 59 mi NW Ciudad Oaxaca, (UMMZ 121824);
Quiotepec (USNM 30338-type of m^/); San Felipe del Agua, 6500' (AMNH 97988,
100932-33); 2 km W San Felipe del Agua, (UIMNH 60799); Cerro San Felipe
(UIMNH 53121, 73654, 74462-64) ; San Lucas Camotlan (AMNH 89630); 3 mi N
San Miguel del Valle, 6950' (AMNH 97986-87).
Puebla: no specific locality (ZMB 6652-syntypes of latifascia); 11.4 mi NW Acatlan
(LACM 38206); 16 mi N Acatlan (LACM 9509); 6 mi SE Acatlan (KU 31680);
10.1 mi NW Amatitlan (UF 11337); 10 mi NW Huajuapan de Leon on Hwy 190
(MVZ 81355); 26 mi NW Huajuapan de Leon on Hwy 190 (MVZ 81354); 3 km W
Izucar de Matamoros (KU 39625); 17 mi SE Izucar de Matamoros (AMNH
89631); San Diego, ca. 5 mi S Tehuacan (USNM 110402-type of forbesi); 6.7 mi
NNW Tehuitzingo (UF 11335); 6.3 mi SE Tehuitzingo (UF 11336).
San Luis Potosi: 10.5 mi W Rioverde, 4000' (UMMZ 126193); 2.5 mi S Pendencia
(LSUMZ 298).
Sinaloa: 16 km NNE Choix, 1700' (KU 68754); 22.1 mi N Culiacan (LACM
53024); 4 mi NNE El Fuerte, 300' (FMNH 71531) 2-3 mi ESE Rio Fuerte Dam
[8 mi NNE El Fuerte, 300'] (FMNH 71532-33) 8.3 mi N Guacamil [=Gua-
muchil] (UF 12829); 1.6 mi S Guacamil [=Guamuchil] (UF 12830); Terreros
(LACM 9510); near Terreros (LACM 7112); 2.5 mi NW Terreros (LACM 7108);
5 mi NW Terreros (LACM 7109); 7 mi NW Terreros (LACM 7111); 10 mi NW
Terreros (LACM 7 110); 10.2 mi SE Terreros (LACM 7113).
Sonora: La Aduana, 5 mi W Alamos (LACM 53028) ; Alamos (AMNH 75119); W
1970
Geographic Variation of Mexican Lyre Snakes
39
side Alamos (KU 24119); 3.5 mi W Alamos (LACM 9157); 9.7 mi W Alamos,
1300' (LACM 53029); 8 mi SSE Alamos on Rio Cuchujaqui, 900' (LACM 53030);
Guirocoba (MVZ 50833-34); ca. 15 mi SE Navojoa (MVZ 76372); 18 mi SE
Navojoa (MVZ 76373); 20 mi E Navojoa (LACM 53027).
Tamaulipas: 10 km N Antiguo Morelos (MU 196); S Ciudad Victoria, on Hwy 85
between C. Victoria and Llera (UMMZ 1 1 1255) ; 13 mi S Ciudad Victoria (UIMNH
19284); 14 mi S Ciudad Victoria (UIMNH 27164); 23 mi S Ciudad Victoria
(AMNH 72399) ; 24 mi S Ciudad Victoria (AMNH 72400) ; Gruta de Quintero, near
Quintero (AMNH 58224) ; La Joya de Salas, 1550 m (UMMZ 110882).
Vera Cruz: Jalapa (BM 81.10.31.65); Tuxpango, near Orizaba (USNM 26499-type
of collaris).
Zacatecas: Hacienda San Juan Capistrano (USNM 46334).
Locality Unknown: “Mexico” (FMNH 42066; USNM 9911-12, 25361, 26138-9
[see Taylor, 1939: 366 for comments] ).
“Southern Mexico” (BM 95.1.4.7 A and B)
“Districto Federal” (AMNH 19718 [seeZweifel, 1959: 7 for comments]).
Additional Specimens {examined but data not included)
Colima: 4.3 mi SSW Colima, 1300' (UF 24783).
Hidalgo: 10.4 mi NNW Ixmiquilpan, 7500' (UF 27378).
Jalisco: 2 mi NE El Molino (UAZ 27023).
Sonora: 6 mi W Alamos (ASU 6651, 6684); 9 mi W Alamos (ASU 6377); 11 mi W
Alamos (ASU 6663, 6712); 14 mi W Alamos (ASU 6648).
Literature Records (specimens not examined)
Chihuahua: Batopilas (Cope, 1887: 68; Klauber, 1928: plate 22); city of Chihuahua
(Cope, 1900: 1105).
Jalisco: “Jalisco” (Mocquard, 1899: 157); Guadalajara [USNM 12419] (Taylor,
1939: 366; Smith, 1941: 163).
Michoacan: San Salvador [BM 1914.1.28.136] (Peters, 1954: 32-34).
Morelos: between Cuernavaca and Tepoztlan; near Huajintlan (Taylor, 1941: 479).
Nayarit: Sierra del Nayarit (Mocquard, 1899: 157).
Tamaulipas: 14 mi SW Jimenez (Brown and Brown, 1967: 325).
Resumen
La variacion geografica en lepidosis y tipo de coloracion han sido es-
tudiadas en las siguientes especies nominales Trimorphodon collaris, fasciolata,
forbesi, latifascia, tau y upsilon. Los ejemplares fueron agrupados en siete
diferentes muestras geograficas que fueron luego comparadas entre si. La
mayoria de las caracteristicas tienen una amplia distribucion y se confunden
mas con otras; otras caracteristicas varian independientemente. Las escamas
ventrales y subcaudales disminiiyen en numero de norte a sur. El numero de
bandas dorsales en el cuerpo es menor en culebras del Valle de Balsas-
40
Contributions in Science
No. 179
Tepalcatepec, las cuales tienen tambien coloracion uniforme en la cabeza.
Ejemplares de otras regiones tienen una compleja coloraeion cefaliea que
consiste en bandas prefrontales, bandas interoculares y marcas parietales
discretas en varias combinaciones. Estudios de los bandas en el cuerpo y
coloracion cefaliea indican un intercambio genetico limitado entre los ejem-
plares del Valle de Balsas-Tepalcatepec y aquellos de otras regiones, posi-
blemente como resultado de contacto secundario en el sur de Jalisco y la region
adyascente de Michoacan. La Cadena Volcanica Transversa y la Sierra Madre
del Sur aparentemente restringen el intercambio genetico.
Estas especies nominates se asignan al sinonimo Trimorphodon tau. El
nombre Trimorphodon tau tau se conserva para las poblaciones de las colinas
y la meseta y T. tau lati fascia (nueva combinacion) se asigua a la pobla-
cion del Valle de Balsas-Tepalcatepec. Trimorphodon tau es redescribo y se
delimita su distribucion. La variacion en caracteristicas de un grupo procedente
de Puebla es semejante a la variacion tipica que se encuentra en la muestra
total de Puebla.
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Jan, G. 1863. Elenco sistematico degli ofidi descritti e disegnati per ITconografia
generale. Milano, 143 p.
Jan, G. and F. Sordelli. 1872. Iconographia generale des ophidiens. Milan, vol. 3,
livr. 39, plate 1, fig. 3.
Jones, C. and J. S. Findley. 1963. Second record of the lyre snake Trimorplwdon
vilkinsonii in New Mexico. Southwestern Natur., 8(3) : 175-177.
Klauber, L. M. 1928. The Trimorplwdon (lyre snake) of California, with notes on
the species of the adjacent areas. San Diego Soc. Nat. Hist., Trans., 5(11):
183-194, 2 plates.
. 1941 . The correlation between scalation and life zones in San Diego County
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. 1956. Rattlesnakes. Their habits, life histories and influence on mankind.
2 vols., Univ. California Press, Berkeley and Los Angeles, 1476 p.
Leopold, A. S. 1950. Vegetation zones of Mexico. Ecology, 31(4) : 507-518.
Lynch, J. D. and H. M. Smith. 1965. New or unusual amphibians and reptiles from
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Maldonado-Koerdell, M. 1964. Geohistory and paleogeography of Middle Amer-
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3-32.
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1-102, 7 plates.
Mocquard, M. F. 1899. Reptiles and batraciens recueillis au Mexique par M. Leon
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Geographic Variation of Mexican Lyre Snakes
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113.
Accepted for publication June 30, 1969.
■■ik. k
i
i
LOS
ANGELES
COUNTY
MUSEUM
CONTRIBUTIONS
IN SCIENCE
UMBER 180
February 27, 1970
STUDIES ON NORTH AMERICAN BEES OF THE GENUS
HYLAEUS. 5. THE SUBGENERA HYLAEUS, S. STR. AND
PARAPROSOPIS (HYMENOPTERA: COLLETIDAE)
By Roy R. Shelling
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 sep-
arately, and numbers run consecutively regardless of subject matter. Number 1 was
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should be directed to Virginia D. Miller, Los Angeles County Museum of Natural
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STUDIES ON NORTH AMERICAN BEES OF THE GENUS
HYLAEUS. 5. THE SUBGENERA HYLAEUS, S. STR. AND
PARAPROSOPIS (HYMENOPTERA: COLLETIDAE)
By Roy R. Swelling^
Abstract: The Nearctic representatives of the two subgen-
era Hylaeus, s. str. and Paraprosopis are considered in this paper.
Within Hylaeus, s. str,, eleven nominate species are recognized,
one of these with a morphologically differentiated subspecies.
One species {H. granulatus) is removed from synonymy, one new
species {H. sejunctus) is described from the southwestern United
States, and one name {H. gaigei) is transferred to the subgenus
Prosopis. The distribution and morphological variation of several
species are treated in detail, and the hypothesis is advanced that
one species {H. stevensi) may be adventive from the Palearctic
fauna. A key is given for all Nearctic species and facial views of
the heads of both sexes of most species, and the male apical ven-
trites are illustrated.
Fourteen Nearctic species of Paraprosopis are recognized
and included in the key to the species. Two new species {H. luni-
craterius, H. timberlakei) are described from Idaho and Cali-
fornia, respectively. The species H. hydrangeae is transferred
from the subgenus Metziella to Paraprosopis as a synonym of
H. georgicus. The distribution of most species is discussed and
facial characteristics and male apical ventrites of all species are
figured.
The present paper is a continuation of those comprising the series in
Literature Cited (Snelling, 1966 a, b, c; 1968), and includes all the species
assigned to the nominate subgenus and to Paraprosopis. One additional paper,
treating the subgenus Prosopis in greater detail than my earlier paper (1966a)
is in preparation. Hopefully, the series will conclude with a treatment of the
species of Mexico and Central America.
HYLAEUS S. STR.
The nominate subgenus is, in the Nearctic Region, one of the larger
groups in numbers of species. Except for the ubiquitous H. cressoni (Cock-
erell), however its component species are less abundant than those of Prosopis
and Paraprosopis. As is the case of Prosopis, Hylaeus s. str., is Holarctic and
most of the species are Palaearctic. Our fauna is closely allied to that of the
Old World, and one species, H. ellipticus (Kirby) shows definite affinities with
a small complex of Old World species. Its nearest relative seems to be
^ Entomology Section, Los Angeles County Museum of Natural History, Los Angeles,
California 90007.
1
CMITUOomi A
2
Contributions in Science
No. 180
H. cardioscapus Cockerell, of Siberia. The females, especially, are difficult to
separate.
Our species are definitely of boreal origin and seem to have only com-
paratively recently invaded the more arid regions of the Southwest. Thus, of
the eleven species here recognized, only one is a true xerophile. One other
species has evolved a xeric subspecies. Of the remainder all save one are
essentially northern and/or montane in distribution. The one exception seems
to be restricted to the relatively cool sand dune areas along the California coast.
That the invasion of the New World is a phenomenon of long standing is
attested by the extensive southern distribution of the group and by the develop-
ment of several rather distinctive groups of species. The most primitive of our
species seems to be H. ellipticus which extends from as far north as the Firth
River in the Northwest Territories south to California, Arizona and New
Mexico. From Alaska on the west H. ellipticus ranges eastward across Canada
to the Atlantic Coast in Nova Scotia. From there it ranges south as far as
North Carolina and Tennessee at the high elevations in the Appalachians.
Another essentially boreal species is H. verticalis (Cresson). The north-
ern limit of this species is unknown to me although I have a few specimens
from scattered localities in southern Alaska. It occurs as far south as Cali-
fornia, Arizona and New Mexico in the western states, and North Carolina and
Tennessee in the eastern. At its southern extremities H. verticalis occurs only
at high elevations.
Two species comprise the H. conspicuus group : H. conspicuus (Metz)
and H. maritimus Bridwell. Both are highly modified from the usual pattern
of the H. cressoni group, but seem, nonetheless, to be related to that assemblage
through H. rudbeckiae (Cockerell and Casad) . Hylaeus conspicuus is the more
common and widely distributed, extending along the Pacific Coast states from
British Columbia south to southern California. Eastward the range includes
northern Nevada and Utah, Idaho, Wyoming and Montana. Although often
locally abundant H. conspicuus is of sporadic occurrence and is seldom rep-
resented in collections by large series. Far more restricted in distribution,
H. maritimus is known only from coastal sand dunes of California from
Sonoma County south to Santa Barbara County.
The one completely isolated species in the Nearctic fauna is H. stevensi.
This species seems to be unrelated to any species in the Nearctic fauna. This
species is most abundant in the north-central and northwestern states, but does
not extend very far into Canada. In many samples which I have seen from
these areas it is obviously the dominant species of Hylaeus. Beyond this region
its occurrence is spotty but it may be locally very abundant. For example, it is
quite common along the San Joaquin River, ten miles west of Turlock, Stan-
islaus County, California, where it frequents Melilotus spp. At Turlock, where
I collected intensively for about ten years, it appears to be very rare.
The remaining group is apparently entirely Nearctic in distribution. This
1970
Studies on North American Bees
3
is the H. cressoni group which includes, in addition to H. cressoni itself,
H. fedorica (Cockerell), H. labiatifrons (Cockerell), H. rudbeckiae (Cock-
erell and Casad), H. saniculae (Robertson) and H. sejunctus, n. sp.
The most common, and most widely distributed, member of this group is
H. cressoni which extends transcontinentally from southern Canada to north-
ern Mexico. Its elevational amplitude exceeds that of all other species in the
subgenus, extending from sea level in California to altitudes in excess of 9500
feet in the same state and at the same latitude. Perhaps in response to this
extensive distribution, H. cressoni has produced a welter of varietal forms, a
number of which have been formally named. I recognize only the form which
inhabits the southwestern deserts as a valid subspecies. Several additional
species are very closely related to H. cressoni and their females are extremely
difficult to separate from those of H. cressoni. Of these species, only H. rud-
beckiae is at all common and widely distributed, and its range is nearly as
extensive as that of H. cressoni, the species being absent only from very arid
regions. The only other member of the group other than H. cressoni mesillae
to occur in desert areas is the new species H. sejunctus; this species, obviously
derived from H. cressoni, occurs sporadically in desert regions in southern
California and Nevada, western Arizona and extreme northwestern Mexico.
Three species, H. fedorica, H. labiatifrons and H. saniculae are found in the
midwestern and eastern states. All are rare and very poorly known.
It is not very difficult to visualize the evolution of the H. cressoni group
from a more primitive bee such as H. ellipticus. Such requires only a reduction
in size and in punctation, together with some modification of the male sec-
ondary characters (scape configuration, extent and orientation of facial
maculae). By a similar reasoning, the H. conspicuus group can be derived
from the same source, through an intermediary form such as H. rudbeckiae.
The larger size, larger and more clearly defined sculpturation and Holarctic
distribution of the H. ellipticus group favor the conjecture that this is the more
primitive of the two groups. The H. verticalis group appears to be still less
specialized, and our one representative is structurally a more variable species.
From the above sketchy commentary on our species of Hylaeus, s. str., it
should be obvious that the species of this subgenus will prove difficult to
identify. This particularly is true of those which I have assigned to the H.
cressoni group. In the key which follows one should not experience any diffi-
culty with the females until those couplets dealing with the H. cressoni group
are reached (Nos. 7-12) . I have been forced to rely on characters which are,
at best, exceedingly difficult to work with, and the results are far from satis-
factory. During the preparation of this portion of the key, an AO-Spencer
binocular microscope, with 18x ocular eyepieces and 6x objectives, was used.
A five unit measuring reticle, divided into twenty subunits was used, this cali-
brated against a stage micrometer for conversion into millimeters. Facial
measurements were made with the head in full-face view, i.e., the head so
4
Contributions in Science
No. 180
positioned as to produce maximum distance between the clypeal apex and the
posterior head margin. Head length is a useful character although even slight
overall size increase usually results in a disproportionate increase in the devel-
opment of the vertex; this does not become apparent until one has examined
many specimens. Far less variable, and hence more useful for some purposes,
is the maximum measurable distance between the clypeal apex and the anterior
margin of the anterior ocellus, along the mid-line of the face. This I have called
the facial length. With the head positioned as indicated above, a measurement
of the maximum width, across the compound eyes, is made to yield the head
width. This measurement is subject to some allometric variation and hence is
less useful than desired, but its alternative, facial width (maximum measurable
distance between the eyes at the level of the antennal sockets) is less easily
made with precision and is only slightly less variable.
Characters of punctation have also been used to separate females within
the H. cressoni group. These are subtle and difficult to express, and until one
has become somewhat familiar with the various species, they are very difficult
to appreciate. Unfortunately, there is a considerable amount of variation,
especially in H. cressoni, and many (about 15% of those I have studied) fe-
males cannot be keyed satisfactorily. The alternative was to bring all females
of this group to a single couplet without any attempt to separate them. Others
may wish to adopt this alternative and stop keying at couplet 7. Tentative
determinations may then be made on the basis of associated males, although
this is hazardous since there are few areas where but a single species is found,
although in any area one species will be dominant, usually H. cressoni.
The portion of the key separating the males should present few difficul-
ties. Two forms (H. cressoni mesillae and H. sejunctus) are separated by
differences in the shape of the ninth ventrite since no reliable external features
have been discovered. All specimens from the southern California deserts,
southwestern Nevada, the western third of Arizona and adjacent portions of
northwestern Mexico must be separated in this manner. Specimens from the
remaining areas of the southwestern deserts will all be H. cressoni mesillae or
intergrades to the typical form (these are particularly common in Utah and
Texas) . Males of the other species may also be identified simply by comparison
with the figures of the faces; these are quite distinctive for each species.
KEY TO HYLAEUS, S. STR.
1. Females 2
Males 13
2. Anterior coxae not distinctly angulate laterally; facial foveae ending no
more than one-fourth of distance between eyes and lateral ocelli; pro-
notal collar distinctly shorter medially than laterally; pronotal collar
usually maculate 3
Anterior coxae distinctly angulate laterally; facial foveae ending about
1970
Studies on North American Bees
5
one-third of distance between eyes and lateral ocelli; pronotal collar
usually as long medially as laterally; pronotal collar immaculate
verticalis (Cresson)
3. Oblique propodeal carina evanescent or absent; lateral carina evanes-
cent or absent above; mesopleura very densely tesselate, punctures
obscure; if mesopleura densely tesselate above only, then genae broader
than eyes 4
Oblique and lateral propodeal carinae usually well-defined; mesopleu-
rae frequently punctate and shiny above, if densely tesselate and im-
punctate, than genae narrower than eyes 5
4. Entire mesopleuron densely tesselate, dull, punctures very obscure;
genae, seen from side, distinctly narrower than eyes; median length of
basal zone of propodeum much shorter than median length of post-
scutellum; clypeus usually extensively maculate conspicuus (Metz)
Mesopleuron moderately shiny and punctate on lower half; genae, seen
from side, distinctly broader than eyes; median length of basal zone of
propodeum subequal to median length of postscutellum; clypeus
immaculate maritimus (Bridwell)
5. Mesopleural punctures extremely coarse, several times diameter of
those of vertex, distinctly larger than those of mesocutum, integument
dull, densely tesselate; postscutellum with deep, distinct coarse punc-
tures, frequently appearing rugose stevensi (Crawford)
Mesopleural punctures little, if any coarser than those of vertex or
mesocutum; punctures of postscutellum usually obscure, never coarse.... 6
6. Lateral and oblique carinae very strongly developed; transverse carina
present, high; entire basal area coarsely rugose; mesopleura deeply,
uniformly punctate; postscutellum dull, densely tesselate and rough-
ened; larger species, length to apex of second tergite 5. 5-6.7 mm; cly-
peus frequently with transverse, apical macula ellipticus (Kirby)
Lateral and oblique carinae variable, but usually rather faint; transverse
carina usually absent; basal area rarely entirely rugose; if rugose, then
mesopleura tesselate on upper half, punctures sparse, obscure; post-
scutellum usually slightly shiny, often with evident punctures; smaller,
slender species, length to apex of second tergite 2. 7-4. 2 mm 7
7. Head width 1 .40 mm or more, usually about 1.5 mm; facial length 1.2
mm or more rudbeckiae (Cockerell and Casad)
Head width not exceeding 1.31 mm; facial length not exceeding 1 .2 mm 8
8. Punctures of mesocutum fine, separated by 1.5 or more times a punc-
ture diameter; interspaces very densely tesselated so that integument
is dull 9
Punctures of mesocutum coarser, separated by about a puncture diam-
eter or less; interspaces usually somewhat shiny; if densely tesselate,
punctures quite close 10
6
Contributions in Science
No. 180
9. Punctures of mesocutellum medially distinctly closer than those on
either side, interspaces somewhat polished sejunctus Snelling
Punctures of mesocutellum medially little, if any, closer than those on
either side, interspaces slightly shining, distinctly tesselated
- - cressoni mesillae (Cockerell)
10. Clypeus and lower half of supraclypeal area tesselate, but slightly
shining with extremely fine longitudinal striolae ........11
Clypeus and lower half of supraclypeal area very densely tesselate,
appearing almost granular, without fine longitudinal striolae
fedorica (Cockerell)
11. Pronotal collar immaculate 12
Pronotal collar maculate; tegulae maculate; clypeus either maculate
apically or strongly suffused with light reddish
- cressoni cressoni (Cockerell)
12. Facial maculae well-developed; tubercules and basal two-thirds of front
tibiae maculate cressoni cressoni (Cockerell)
Facial maculae greatly reduced or absent; tubercules immaculate; front
tibiae with small basal spot saniculae (Robertson)
13. Sides of face without conspicuous oval concavity; clypeus rarely marked
with black 14
Sides of face, slightly above level of antennal sockets, with a conspicu-
ous oval concavity; lateral face marks narrow, elongate; clypeus usually
partially infuscated along lower margins saniculae (Robertson)
14. Antennal scape broad, its greatest width equal to one-half or more its
length ....15
Antennal scape narrower, its greatest width not exceeding 0.47 times its
greatest length 19
15. Scape more or less dilated, strongly flattened to concave beneath; lateral
face marks variable... 16
Scape robust, thick, hardly, if at all, flattened beneath; lateral face marks
curving away from eye margins over antennal sockets (Fig. 2H)
verticalis (Cockerell)
16. Lateral face marks above well-removed from inner orbits and con-
spicuously dilated or clavate above (Figs. 2C, D, J) ; transverse propo-
deal Carina evanescent or absent ......17
Lateral face marks above not separated from eye margin, but with
pointed median extension ending over antennal sockets (Fig. 2C) ;
transverse propodeal carina sharply defined ....ellipticus (Kirby)
17. Scape no more than 0.58 times as broad as long; concave surface black,
with at most a narrow yellow stripe along lower margin; mesopleura
distinctly punctate 18
Scape at least 0.70 times as broad as long, concave under surface
wholly yellow, with broad yellow stripe along lower margin which
1970
Studies on North American Bees
7
expands above onto convex surface, (Fig. 2J); mesopleura densely
tesselate and roughened, without conspicuous punctures
conspicuus (Metz)
18. First flagellar segment subequal to second; rugulae of basal area of
propodeum not attaining posterior margin; the rugulae longitudinal,
widely spaced; oblique carina absent; lateral face marks slightly con-
stricted above leval of clypeus, separated from inner orbits by less than
their width at constriction (Fig. 2D) maritimus Bridwell
First flagellar segment half, or less, as long as second; basal area of
propodeum rugoso-reticulate to posterior margin; oblique carina
present, face marks strongly constricted, separated from inner orbits by
much more than their minimum width at constriction (Fig. 2F)
rudbeckiae (Cockerell and Casad)
19. Mesopleural punctures little, if any, coarser than those of frons; trans-
verse and oblique carinae usually absent; lateral face marks constricted
above (Figs. 2 A, G, I) ; smaller species, length to apex of second tergite,
2.7-3.5 mm 20
Mesopleural punctures much coarser than those of frons; transverse and
oblique carinae present, distinct; lateral face marks not constricted
above (Fig. 2B) ; larger species, length to apex of second tergite, 3. 0-4. 2
mm stevensi (Crawford)
20. Lateral face marks expanded above constriction, or continued upward
and curving inward above antennal sockets (Figs. 2G, I) 21
Lateral face marks not expanded above constriction, rarely extending
beyond level of upper margin of antennal sockets, never curving inward
above sockets (Fig. 2A) fedorica (Cockerell)
21. Punctures of mesoscutum and mesoscutellum coarse, separated by one-
half a puncture diameter or less, the interspaces densely tesselate, and
dull; rugulae of basal area of propodeum usually extending to summit
of declivity 22
Punctures of mesoscutum and mesoscutellum fine, mostly separated by
one and one-half times a puncture diameter or more, interspaces lightly
tesselated and moderately shiny; rugulae of basal area evanescent be-
yond middle, rarely attaining summit of declivity 23
22. Tibiae and tarsi entirely yellow; tegulae maculate; scape largely yellow
beneath labiatifrons ( Cockerell )
Middle and hind tibiae yellow on basal-half only; tegulae often black;
scape often entirely dark beneath cressoni (Cockerell)
23. Apical lobes of ninth ventrite shorter than pedicle
cressoni mesillae (Cockerell)
Apical lobes of ninth ventrite distinctly longer than pedicle
sejunctus Snelling
8
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Hylaeus (Hylaeus) verticalis (Cresson) , NEW STATUS
Figs. IG, 2H, 3B, 5B
Prosopis verticalis Cresson, 1869. Proc. Boston Soc. Nat. Hist. 12:271. $ .
Prosapis tridentulus Cockerell, 1896. Psyche 7 (sup.): 27. $ NEW
SYNONYMY.
Prosapis tridens Cockerell, 1896. Op. cit.\ 27. $ .
Prosopis subtristis Swenk and Cockerell, 1910. Ent. News 21 :70 9.
NEW SYNONYMY.
H
Figure 1. Hylaeus (//.) spp., female faces: A, H. fedorica; B, H. stevensi; C, H.
ellipticLis', D, H. maritimus; E, H. riidbeckiae; F, H. cressoni cressoni', G, H. verticalis;
H, H. sejunctus; I, H. conspiciius. Figures by Ruth A. DeNicola.
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Studies on North American Bees
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Prosopis melitina Lovell, 1911. Ent. News 22:214. $.
Prosopis tridentulus, Metz, 1911. Trans. Amer., Ent. Soc. 37:116-117.
Prosopis episcopalis var. subtristis, Metz, 1911. Op. cit.\ 131. 9 {in part) .
Hylaeus verticalis, Cockerell and Sumner, 1931. Amer. Mus. Nov. 490:
3-4.
Hylaeus verticalis tridentulus, Linsley, In Mueseback, et ah, 1951.
U.S.D.A., Agr. Monog. No. 2:1050.
Hylaeus verticalis verticalis, Linsley, Op. cit.: 1052. Mitchell, 1960. N. C.
Agr. Exp. Sta. Tech. Bui. 141 :76.
With the exception of H. cressoni no species of Hylaeus is more widely
distributed than H. verticalis. Throughout most of its range this species is quite
common and highly variable. Most of the variation occurs in the females.
This variation assumes several different forms. In western populations there
is a strong tendency for the apical one-third of the clypeus to be suffused with
reddish, rather than entirely black. This variant form is found together with
females which have the clypeus entirely black, but there is a noticeable trend
toward the maculate condition in those populations which occur at lower
elevations in the Sierran and Cascade ranges, and it is a dominant feature in
the populations of the Coast Ranges in California. The maculate clypeus is
rare in northern and eastern populations. Very rarely are specimens found
which have small yellow maculate on the pronotal collar. I have seen only
three such specimens among the hundreds I have examined; all were females,
one from Idaho, one from Colorado and one from California.
Variation, too, occurs in the shape of the pronotal collar. In the usual
condition the pronotal collar, when viewed from above is very nearly as long
in the middle as at the sides. In profile it is convex and evenly rounded toward
the anterior pronotal neck. Some females, however, have the middle portion
flat and somewhat compressed so that it is much shorter here than at the sides.
Such specimens, on the basis of the material I have seen, occur only in the
western populations, and are especially common in samples from Oregon,
California and Nevada. No specimens presenting an intermediate condition
are known to me, and these individuals may actually represent a cryptic sibling
of H. verticalis. I have not, however, been able to associate them with any
male variant, so am content to assign them to H. verticalis for the present.
I have examined the types of H. tridentulus, H. tridens and H. subtristis
and can find no justification for their continued recognition. The first two, both
based on males from Colorado, were named because of slight differences in
the color of the maculae and slight differences in punctation from the typical
form described from Pennsylvania. With large series of males from the entire
range of the species available, it is evident that they fall well within the total
range of variation exhibited by any one population. The females described by
Swenk and Cockerell from Nebraska as H. subtristis are perfectly typical of
the western populations of H. verticalis; they were merely the, until that time.
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undescribed opposite sex of this species. The females which Metz (1911) had
called H. episcopalis subtristis were a conglomerate association which in-
cluded, in addition to H. verticalis, H. rugulosus rugulosus (Cockerell), H. r.
episcopalis (Cockerell) and H. modestus citrinifrons (Cockerell). I have not
seen the type of H. melitina, but as there is nothing in its description to indicate
merit for its further consideration, I have accepted its synonymy as published
by Mitchell (1960).
Hylaeus {Hylaeus) cressoni cressoni (Cockerell)
Figures IF; 2H; 6.
Prosopis pygmaea Cresson, 1869. Proc. Boston Soc. Nat. Hist. 12:272.
$ . Preoccupied by Schenck, 1853.
Prosopis cressoni Cockerell, 1907. Annals and Magazine of Natural
History (7) 20:131. New name for P. pygmaea Cresson, not P. pygmaea
Schenk. Metz, 1911. Trans. Amer. Ent. Soc., 37:105. $ $.
Prosopis pasadenae Cockerell, 1910. Annals and Magazine of Natural
History (8) 5:30. $ $.
Prosopis teleporus Lovell, 1911. Ent. News, 22:213. $ NEW SYN-
ONYMY.
Hylaeus mesillae race pasadenae, Meado-Waldo, 1923. In Wytsman,
Gen. Insect., fasc. 181:30.
Hylaeus laciniatus Cockerell and Sumner, 1931. Amer. Mus. Novitates,
490:9-10. $ . NEW SYNONYMY.
Hylaeus repolitus Cockerell and Sumner, 1931. Op. cit., 13-14. $ . NEW
SYNONYMY.
Hylaeus {Hylaeus) cressoni cressoni, Linsley, 1951. IN Muesebeck, et al.,
Hymen. Amer. N. Mex., Synoptic Cat., Monogr. 2, U.S.D.A., p. 1050;
Mitchell, 1960, N. C. Agr. Expt. Sta. Tech. Bull. 141:60, 62, 70.
Hylaeus {Hylaeus) cressoni pasadenae, Linsley, 1951. IN Muesebeck,
et al. Hymen. Amer. N. Mex., Synoptic Cat., Monogr. 2, U.S.D.A., p. 1050.
Hylaeus {Hylaeus) teleporus, Linsley, 1951. IN Muesebeck, et al.,
Hymen. Amer. N. Mex., Synoptic Cat., Monogr. 2, U.S.D.A., p. 1052;
Mitchell, 1960, N. C. Agr. Expt. Sta. Tech. Bull. 141:60, 62, 75; fig. 11.
This species extends transcontinentally from southern Canada to north-
ern Mexico. Over most of this range H. cressoni is a common species. The
populations from eastern, northern and montane regions commonly have the
clypeus and pronotal collars immaculate. Some individuals within these popu-
lations may also lack maculae on the pronotal lobes and the tegulae. Clinal
variation occurs uniformly from east to west, from north to south and with
decreasing elevation. This clinal variation is manifested in two concordant
characters, an increase in the tone and extent of the pale maculae and a de-
crease in the size and density of metasomal punctation, so that populations at
opposite poles of the dine are very different in appearance. Specimens from
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Turlock, California, for example have the face marks bright yellowish, the
apical portion of the clypeus is strongly infused with reddish color and the
pronotal lobes and collar and the tegulae are conspicuously maculate. The
abdomen is smooth and shiny, with few or no distinct punctures. Such speci-
mens contrast sharply with others from the New England area in which the
maculae are dull yellowish-white, the clypeus is black and the thorax com-
pletely immaculate; such specimens may have the metasomal tergites with
obvious fine scattered punctures. The same is true if one compares the Turlock
specimens with samples from Revelstoke, British Columbia, or with others
from Sonora Pass in the Sierra Nevada of California. In all cases the two
extremes are connected, through the intervening portions of the range, by a
series of specimens of intermediate character. These series of intermediates
form, in their entirety, a well-defined dine for the characters noted.
Under these cricumstances continued recognition of H. pasadenae, de-
scribed from Pasadena, California, as a subspecies of H. cressoni, a status
accorded this form in the past, is untenable. This bee does not possess any of
the characteristics of a definable subspecies and I agree with Metz in placing
H. pasadenae in the synonymy of H. cressoni. The subsequent continued
recognition of H. pasadenae as a subspecies of H. cressoni seems to be trace-
able to Cockerell’s refusal to accept the synonymy published by Metz. How-
ever persistent such a refusal has been, there appears to be no published
indication why Metz’s proposal should not be adopted.
Lovell’s H. teleporus was proposed for H. cressoni-\i\iQ. males in which
the lateral face marks terminate acutely at the level of the lower margin of the
antennal sockets. These males seem to occur sporadically within the popula-
tions of H. cressoni in northern and montane areas. I have examined the
terminalia of males of this form and find no distinctive characters which will
serve to separate H. teleporus from H. cressoni. I believe that H. teleporus
should be placed in the synonymy of H. cressoni until conclusive evidence for
the specificity of this form can be advanced.
Cockerell and Sumner described H. repolitus from a single female taken
at Ogden, Utah. Dr. Rozen sent the type to me and I find nothing in any of its
characters to justify separating this bee from H. cressoni. The nearly smooth
basal zone of the propodeum is exactly like that of many females of H. cressoni
available to me which also show a strong reduction in the longitudinal rugulae
of the basal zone. This characteristic cannot be correlated with other features
nor is it geographically consistent.
Dr. Rozen has sent the type of H. laciniatus, and there is no doubt that
this name, too, is a synonym of H. cressoni cressoni. Before I had the oppor-
tunity to study the type, I was puzzled by the peculiar shape fo the seventh ven-
trite as figured by Cockerell and Sumner (1930:10, fig. 2). With the type
slide before me, it is clear that the apical lobes of this ventrite have been
broken off. The poor preparation of the slide, on which everything is badly
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No. 180
flattened and distorted, is also responsible for the seemingly expanded gono-
coxites, as these were illustrated by Cockerell and Sumner.
Cockerell and Sumner cite the type as follows:
“COLORADO— Boulder, May 24, 1913 (F. E. Lutz).” The labels on the
specimens clearly read “Boulder, Colo., M. D. Ellis, May 24, 1913,” and
“5u//x.” Since the specimen and slides are in accord with the original descrip-
tion and figures, and the type is so marked in Cockerell’s handwriting, I have
no reason to doubt that this is the true type.
Hylaeus (Hylaeus) cressoni mesillae (Cockerell)
Figures 3C; 5C; 6.
Prosopis subtilis Cockerell, 1895. Trans. Amer. Ent. Soc., 22:295. 9.
Preoccupied by Forster, 1871.
Prosopis mesillae Cockerell, 1896. Canad. Ent., 28:42. new name for
P. subtilis Cockerell, not H. subtilis Forster.
Prosopis magniclavis Swenk and Cockerell, 1910. Ent. News, 21:68.
5 9 . NEW SYNONYMY.
Prosopis cressoni, form mesillae, Metz, 1911. Trans. Amer. Ent. Soc.,
37:107 (in part).
Hylaeus cressoni, var. magniclavis, Meade-Waldo, 1923. IN Wytsman,
Gen. Insect., fasc. 181:30.
Hylaeus mesillae, Meade-Waldo, 1923. Op. cit. :30.
Hylaeus {Hylaeus) magniclavis. Linsley, 1951. IN Muesebeck, et al.,
Hymen. Amer. N. Mex., Synoptic Cat., Monogr. 2, U.S.D.A., p. 1051.
Hylaeus {Hylaeus) mesillae, Linsley, 1951. Op. cit. : 1051.
The status of H. cressoni mesillae has been confused. Proposed originally
as a full species, it was reduced by Metz to status as a “form” of H. cressoni.
Crawford, two years later, elevated the name to specific status based on differ-
ences in the shape of the eighth ventrite, and the bee has been recognized at
this level since then. Unfortunately, while Crawford’s conclusions were sound,
they were based on misidentified specimens. I have examined the genitalia and
associated ventrites of many males from New Mexico, Arizona, California
and Sonora, Mexico. With the exception of the California specimens all ex-
hibited characters typical of H. cressoni.
The California material, however, was represented by two very dis-
tinctly different types of terminalia. The first of these was typically H. cressoni,
identical to those specimens from New Mexico and Arizona. The second type
agreed closely with the figure published by Crawford under the name H.
mesillae. Crawford dissected the terminalia from at least three males, which
I have seen; all were from California. The type locality of H. mesillae is
Mesilla, New Mexico, and the terminalia of males from this area are con-
sistently of the H. cressoni type, and I am certain that the interpretation
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Studies on North American Bees
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offered here is the correct one: that H. mesillae is conspecific with H. cressoni,
and that Crawford’s specimens represent another species, described below
as new.
I have reduced H. mesillae to subspecific rank under H. cressoni. The
females of this subspecies are a little smaller, on the average, than those of
I
Figure 2. Hylaeus {H.) spp., male faces: A, H. fedorica; B, H. stevensi; C, H.
ellipticus; D, H. maritimns; E, H. granidatus; F, H. riidbeckiae; G, H. cressoni
cressoni', H, H. verticalis; I, H. sejunctiis; J, H. conspiciiiis. Figures by Ruth A.
DeNicola.
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No. 180
H. cressoni, and the integument of the body is less densely tesselate, hence
more shiny. The punctation, too, differs, being distinctly finer and sparser on
the mesoscutum and scutellum, as indicated in the key. Some western material,
such as the synonymous form of H. cressoni, H. pasadenae, are as extensively
maculate as H. c. mesillae but differ in that the maculae are bright yellow in
these, while in H. c. mesillae they are whitish. These highly maculate specimens
also possess large maculae on the pronotal collars, usually lacking in H. c.
mesillae. When such maculae are present in the desert subspecies, they are
small and inconspicuous.
Females from the desert areas commonly have the rugulae of the basal
zone of the propodeum absent or restricted to the extreme anterior margin. In
most material of H. cressoni cressoni the entire basal zone is distinctly longi-
tudinally rugulose, except in some samples from Colorado and Utah. These
appear to be intergrades between the two subspecies. This is somewhat puz-
zling, since such specimens do not seem to occur in other areas where the
ranges of these two forms overlap.
Another intermediate form occurs in western Nebraska, eastern Wyoming
and northeastern Colorado. This was described by Swenk and Cockerell as
H. magniclavis. I have examined most of the type series of this bee and find
no reason for its continued recognition. The males have the lateral face marks
slightly expanded at their upper ends, but are otherwise similar to that sex of
H. c. mesillae. I cannot find any differences to distinguish the females from
those of H. c. mesillae. The male terminalia do not differ significantly from
those of H. c. cressoni or H. c. mesillae, so there is no doubt that H. magni-
clavis is conspecific with these. Since the pale maculae and fine, sparse
thoracic punctation are in accord with those characters in H. c. mesillae, I
consider H. magnicalvis a synonym of this subspecies.
In figure 6 I have shown the results of a series of cephalic measurements
which reflect differences between H. c. cressoni and H. c. mesillae. In 6A two
phenomena are apparent. First that there are two concentration points on the
basis of head width. There is some overlap, but the existence of two centers of
concentration remains clear. A similar, but more obscure, pattern exists based
on head length. These measurements of head length and head width are use-
HL
fully converted into a Cephalic Index (Cl = Hw X 100) which demonstrates
their relationship as a ratio, figure 6B. A Cl value of 100, of course, indicates
HL and HW being equal, values lower than 100 indicate increasing relative
width. The Cl values of each form have been plotted in 4C as a frequency
indication. From this figure it is obvious that while both forms most commonly
possess Cl values ranging from 91 to 100, a much higher percentage (25%)
of H. c. mesillae have the head longer than broad than is true of the nominate
form (9% ).
The second noteworthy feature in figure 6A is the displacement of the
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Studies on North American Bees
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Figure 3. Hylaeiis (H.) spp., ventrite VIII of males: A, H. ellipticus; B, H. verticalis;
C,H. cressoni mesillae; D, H. sejiinctiis; E, H. scmiciilae; F, H. fedorica; G, H. mari-
timiis; H, H. stevensi.
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No. 180
Figure 4. Hylaeus {H.) spp., ventrites VIII and IX, respectively, of males: A, D,
H. granulatus; B, E, H. rudbeckiae; C, F, H. conspicuus.
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Studies on North American Bees
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Figure 5. Hylaeus (H.) spp., ventrite IX of males; A, H. ellipticus; B, H. verticalis;
C, H. cressoni mesillae; D, H. sejunctus; E, H. saniculae\ F, H. fedorica; G, H. mari-
timus; H, H. stevensi.
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Contributions in Science
No. 180
skew-line of H. c. mesillae to a level below that of H. c. cressoni. There is also
an apparent, though slight, reorientation of the skew-line angle to that of
H. c. cressoni. These suggest that perhaps H. c. mesillae might be best regarded
as a full species rather than as a subspecies. I am not inclined, at this time, to
do so since specimens of an apparently intermediate nature do exist. These
specimens come from those areas where they may be expected to occur if
H. c. mesillae is truly a subspecies, i.e., in the zones of sympatry. It is equally
possible that these are hybrids of two incompletely separate species. However,
since the male terminalia are, for practical purposes, so similar as to appear
identical, I prefer to consider the desert form a subspecies of H. cressoni. This
is also consistent with my treatment of other species.
This subspecies extends from western Nebraska, Kansas and Texas west-
ward to Nevada, California and Baja California, Mexico. Southward it ranges
into the Mexican States of Nuevo Leon, Coahuila, Chihuahua and Sonora.
Populations in Colorado and Utah are largely sympatric with those of the
nominate subspecies and hybridization occurs in these areas.
Hylaeus (Hylaeus) sejunctus Snelling, NEW SPECIES
Figures IH; 21; 3D; 5D.
Prosopis cressoni form mesillae, Metz, 1911, Trans. Amer. Ent. Soc.
37:107 (in part).
Prosopis mesillae, Crawford, 1913. Canad. Ent. 45 : 154-155. Not of
Cockerell, 1896.
This is the species which Crawford called P. mesillae when he raised that
name to specific level after Metz had reduced mesillae to a variety of H. cres-
soni (Cockerell) .
Diagnosis'. The male differs from that of H. c. mesillae in the less coarsely,
closely punctate abdomen and in the shape of ventrite IX. The female differs
in the much finer, sparser punctures of tergite II, the shorter, sparser pubes-
cence of the abdomen, and the smaller size.
Male. Very similar structually to that of H. c. mesillae but differs in hav-
ing the punctures of the mesoscutum finer, sparser, usually separated by a
puncture diameter or more; punctures of tergite I very fine, separated by two
or more times a puncture diameter; discal pubescence of tergite III shorter
than that of mesoscutum; apical lobes of ventrite IX broad, as long as, or
longer than, basal pedicle.
Female. Very similar structually to that of H. c. mesillae but differs as
follows : punctures of median area of upper face of tergite I extremely fine,
perceptible only under high magnification, very sparse; lateral areas of propo-
deum roughened, densely tesselate, with scattered, but distinct, punctures;
erect discal pubescence of tergite III little, if any, more than two-thirds an
ocellar diameter in length (a few widely scattered hairs may attain this length,
but are outnumbered by shorter hairs).
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Studies on North American Bees
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Length, both sexes, to apex of tergite II, 3. 2-3. 7 mm.
Holotype male, allotype female (Los Angeles County Museum of Natural
History), Winterhaven, Imperial Co., California, March 25, 1956 (R. R.
Snelling), Heliotropium curassavicum.
Paratypes. — CALIF. Imperial Co.: 13 $ , 10 $ $ , same data as holo-
type (LACM); 1 9 , 15.8 mi. NW Niland, April 11, 1963 (R. C. Dickson;
UCR), on Prosopis’, \ , 2 mi. S. Travertine Rock, March 29, 1936 (F. R.
Platt; UCR), on Prosopis) 1 9 , 5.2 mi. SE Kane Springs, April 15, 1949 (P. H.
Timberlake; UCR), on Tamarix gallica) 1^,59 9 , Coachella Canal, 20 mi.
SE Hwy 195, April 2, 1960 (R. C. Dickson, P. H. Timberlake; UCR), on
Prosopis) 299, U.S.D.A. Exp. Farm, May 31, 1912 (J. C. Bridwell;
USNM) ; 1 $ , 1 9 , same locality, June, 1912 (J. C. Bridwell; USNM) ; 1 $ ,
1 9 , El Centro, April, 1911 (J. C. Bridwell; USNM) ; 2 $ $ , 1 9 , Imperial
Valley, April, 1911 (J. C. Bridwell; USNM) . San Diego Co.: 1 $ , 1 9 ,
Borrego Valley, April 26, 1954 (J. G. Rozen; CIS) ; 1 $ , Palm Cyn., Borrego
Valley, March 29, 1936 (P. H. Timberlake; UCR), on Prosopis) 1 9 , Borrego
Valley, April 26, 1954 (M. Wasbauer; CIS) ; 1 9 , same locality and collector,
April 30, 1954 (CIS) , on Croton calif or nica) 11 ^ $ , 19 9 9 , Coyote Cr.,
Borrego Valley, April 5, 1963 (F. D. Parker, R. M. Bohart, M. E. Irwin, A.
Willink; UCD, IML); 2 $ $ , Borrego Springs, April 2, 1960 (M. Wasbauer;
CDA), on Phacelia) 2 $ $ , Split Mt., Anza Desert State Park, April 1, 1955
(W. R. Richards; CNC) ; 1 9, Borrego Valley, April 20, 1961 (Rozen and
Schrammel; AMNH), on Prosopis) 1 9 , Borrego Valley, April 11, 1962
(R. M. Bohart; UCD) . Riverside Co.: 10 $ $ , 5 ^ 9 , Andreas Cyn., March
26, 31, 1955 (W. R. M. Mason; CNC) ) \ $ , Willis Palms Oasis, April 9, 1955
(W. R. M. Mason; CNC) ; 2 $ $ , Indio, March 30, 1955 (W. R. Richards;
CNC) ; 1 9 , Andreas Cyn., April 11, 1955 (W. R. M. Mason; CNC) ; 1 $ ,
1000 Palms Cyn., April 3, 1944 (P. H. Timberlake; UCR) ; on Cryptantha
barbigera) 3 $ $ , SVi mi. NW Indio, April 9, 1936 (P. H. Timberlake; UCR) ,
on Prosopis) 3 ^ ^ , Andreas Cyn., April 2, 1955 (P. H. Timberlake; UCR),
on Hyptis enioryi) 1 9 , same locality and collector, April 1 1, 1936 (UCR), on
Prosopis) 299, 1000 Palms Cyn., April 10, 1937 (P. H. Timberlake; UCR) ,
on Prosopis) 19,5 mi. W Indio, April 30, 1949 (E. G. Linsley, J. W. Mac-
Swain, R. F. Smith; CIS) , on Melilotus. San Bernardino Co.: 10 $ $ , 1 2 9 ,
Cronise Valley, April 29, 1956 (J. Powell, M. Wasbauer; CIS) , on Prosopis)
5 $ $ , 1 9 , same locality, date, host (P. H. Timberlake; UCR), on Salix)
1 9 , 29 Palms, April 13, 1935 (P. H. Timberlake; UCR) , on Isomer is arbor ea)
7^^,1199, Needles, May 3, 1964 (P. Torchio & G. Bohart; USU), on
Tamarix and Prosopis) 1 $ , Needles, 20 mi. S., May 3, 1964 (P. Torchio &
G. Bohart; USU). Inyo Co.: 1 $ , Furnace Cr., Death Valley Natl. Mon.,
April 19, 1939 (E. G. Linsley; CIS), on Prosopis. NEVADA. 4 $ $ , Search-
light, 30 mi. S., Clark Co., April 21, 1966 (Torchio, Rust, Yousef; USU).
ARIZONA. 1 9 , Yuma, 21 mi. N., Yuma Co., April 4, 1963 (A. Willink;
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No. 180
IML) . MEXICO. 2 $ $ , 2 2 9 , Mexicali, 20 mi. W., Baja California del
Norte, April, 1939 (C. D. Michener; CAS), on Prosopis.
Hylaeus (Hylaeus) labiatifrons (Cockerell)
Prosopis labiatifrons Cockerell, 1896. Psyche, 7 (suppl.) : 437. $ .
Prosopis cressoni, Metz, 1911. Trans. Amer. Ent. Soc., 37 : 105 (in part) .
Hylaeus {Hylaeus) labiatifrons, Mitchell, 1960. No. Car. Agr. Exp. Sta.
Tech. Bull. 141:62,72. ^ .
The status of this form, described from a single male from Georgia, is
enigmatical. It is known only from the unique type, which I have not ex-
amined. Metz (191 1 ) considered it a synonym of H. cressoni, but Mitchell
(1960) has elevated it to species level and briefly redescribed the type. Un-
fortunately, Mitchell’s description contributes nothing new to permit the
recognition of H. labiatifrons. In all features it seems to agree quite closely
with males of H. cressoni cressoni, and seems to differ from that species only
in the entirely yellow tibiae.
This difference seems to me to be of little value. Another Nearctic
Hylaeus, H. illinoisensis (Robertson), is supposedly recognizable on the same
basis. In that species, at least, males with entirely yellow tibiae are exceptional.
Even those which have the tibiae wholly yellow show, under intense light, a
slightly darker basal two-thirds which corresponds to the black basal area of
most specimens. I suspect that the same is true in the case of H. labiatifrons,
which I regard as a probable synonym of H. cressoni cressoni. The solution
must wait until the type, especially ventrites eight and nine, can be examined
and compared with those of the related species.
Hylaeus {Hylaeus) fedorica (Cockerell)
Figures lA; 2A; 3F; 5F.
Prosopis digit at a var. fedorica Cockerell, 1909. Ann. Mag. Nat. Hist,
(ser. 8) 4:27. $ .
Prosopis grossicornis Swenk and Cockerell, 1910. Entomol. News, 21 :67.
S . Metz, 1911. Trans. Amer. Ent. Soc., 37:118. NEW SYNONYMY.
Hylaeus {Hylaeus) grossicornis, Linsley, 1951. IN Muesebeck, et al.,
U.S.D.A. Monogr. 2:1050; Mitchell, 1960. N. Car. Agr. Expt. Sta., Tech.
Bull., 141:71-72.
I have examined the type of Prosopis digit at a var. fedorica, now in the
American Museum of Natural History, and have compared it with homotypes
of H. grossicornis and there can be no doubt that these are conspecific, as first
suspected by Metz (1911). The type locality of H. fedorica is Fedor, Lee
County, Texas; H. fedorica has priority by one year over H. grossicornis.
This species ranges from Michigan to Minnesota, then southward to
Texas; an eastern extension exists through the southern United States to
North Carolina. The male may be readily distinguished by the characters
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Studies on North American Bees
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.5
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□□□□□□
□ □ □ □ □
□ □ □ □ □
.3
5
I
Q
Q
<
UJ
X
.2
.1
1.0
o #• • •
• <9 • •
o o o
o o O O o
O O § O o o
o o o o o
oo o o o o
o o o o o o
o o oo
o o
o
□ = rudbeckiae
• : cressoni
o= mesillae
— I 1 1 ^ 1 \ 1 r—
1.0 .1 .2 .3 .4
HEAD LENGTH (MM.)
Figure 6. A, scatter diagram plotting Head Width and Head Length of H. rudbeckiae
(squares), H. cressoni cressoni (solid circles) and H. cressoni mesillae (open
circles) ; B, Cephalic Index frequency for H. rudbeckiae (solid line), H. cressoni
cressoni (broken line) and H. cressoni mesillae (dot-dash line).
22
Contributions in Science
No. 180
given in the key. The female, similar to that of H. cressoni, is best recognized
by the shape of the head in full face view. In all other species of the H. cressoni
group, the inner margins of the eyes are strongly divergent above, with the
eyes rather strongly bulging. In these species the widest part of the head is
somewhat above the level of the antennal sockets. In the female of H. fedorica
the inner margins of the eyes are only slightly divergent above; the eyes are
much less strongly bulging above and the widest part of the head is at, or
below, the level of the antennal sockets. The head is always slightly longer
than broad.
Hylaeus (Hylaeus) saniculae (Robertson)
Figures 3E; 5E.
Prosopis saniculae Robertson, 1896. Can. Entomol., 28:137. $ $ .
Prosopis cressoni form saniculae, Metz, 1911. Trans. Amer. Ent. Soc.,
37:106, 107. ^ 2.
Hylaeus {Hylaeus) saniculae, Linsley, 1951. IN Muesebeck, et al,
U.S.D.A. Monogr. 2:1052. Mitchell, 1960. N. Car. Agr. Expt. Sta. Tech. Bull.,
141:73-74.
This rare species, largely northern in distribution, is very poorly known.
In the United States it extends from Minnesota to the New England states and
southward, through the Appalachian Mountains, to Tennessee and Georgia.
In Canada it is known from Ontario and Nova Scotia. Although the male may
be readily recognized by the peculiar oval depressions on the sides of the face,
the female appears to be indistinguishable from that of H. cressoni except for
its more restricted maculae. Attempts to characterize the female of H. sani-
culae are futile at this time because I have seen no females which I can confi-
dently consider to be this species. Reared series, including both sexes, would be
extremely useful in solving this problem.
Hylaeus {Hylaeus) rudbeckiae (Cockerell & Casad)
Figures IE; 2F; 3B; E; 6.
Prosopis rudbeckiae Cockerell and Casad, 1895. Trans. Amer. Ent. Soc.,
22:380. $ ; Metz, 1911. Trans. Amer. Ent. Soc., 37:111-114.
Prosapis bakeri Cockerell, 1896. Psyche, 7 (suppl.) : 26. $ .
Prosapis digit at us Cockerell, 1896. Op. cit., 30. $ .
Prosapis rudbeckiae ruidosensis Cockerell, 1896. Op. cit., 30. $ .
Prosapis rudbeckiae subdigitatus Cockerell, 1896. Op. cit., 31. ^ .
Prosopis rudbeckiae form digitatus, Metz, 1911. Trans. Amer. Ent. Soc.,
37:111, 112.
Prosopis rudbeckiae var. bakeri, Metz, 1911. Op. cit., Ill, 112.
Hylaeus rudbeckiae, Cockerell and Sumner, 1931. Amer. Mus. Novit.,
490:7, 15.
Hylaeus {H.) rudbeckiae, Linsley, 1951. IN Muesebeck, et al., U.S.D.A.
1970
Studies on North American Bees
23
Monogr. 2:1052; Mitchell, 1960. N. Car. Agr. Expt. Sta. Tech. Bull., 141:73.
Although it is a simple matter to separate the males of this species from
those of H. cressoni, I have found no reliable means to separate the females
of the two species. There is a difference in the overall size of the females of
these species which I have found most easily measured in terms of head width.
However, the difference is a minute one and requires precise measurements.
In figure 6A I have given the results of measurements of head width and head
length of females of H. rudbeckiae compared with females of H. cressoni
(represented by both subspecies). The range of head width is limited, varying
from 1.31 mm to 1.50 mm, with the majority of the specimens having a head
width in excess of 1.41 mm. In those specimens which have a head with less
than 1.40 mm, the head length is 1.24 mm or more. In H. cressoni the total
range of head width variation is much greater, extending from 1.12 mm to
1.37 mm. In those few specimens of H. cressoni which have a head width in
excess of 1.34 mm the head length does not exceed 1.23 mm. These differences
are slight but appear to be valid. They have been verified by checking against
associated males and in reared series.
Figure 6B contrasts the cephalic index of H. rudbeckiae with that of the
nominate form of H. cressoni. It may be noted that while both species most
frequently possess a Cl rating of 92, that is, with the head a little broader than
long, a distinct tendency exists in H. rudbeckiae for the head to be broader still,
and no specimens of this species seem to have a cephalic index over 100. While
such high ratings are not common in H. cressoni, they do occur. In H. rud-
beckiae, about 70% of the females examined had a cephalic index of 92 or
less; in females of H. cressoni, only 33% have such a low rating.
In other morphological features the females are very similar and I have
noted none that are sufficiently constant to be reliable. The two species are
almost completely sympatric, H. rudbeckiae absent only from those areas
inhabited by H. c. mesillae. In most areas H. rudbeckiae is slightly more
brightly maculate, the maculae as a rule both more extensive and of a brighter,
yellower tone. The integument of the mesoscutum and scutellum is more
densely tesselate and less shiny.
Hylaeus (Hylaeus) granulatus (Metz)
Figures 2F; 4A; D.
Prosopis rudbeckiae var. granulatus Metz, 1911. Trans. Amer. Fnt. Soc.,
37:114. $.
Although this closely resembles H. rudbeckiae, it may be recognized by
the features of the eighth and ninth ventrites (fig. 4A, D). External features
do not appear to be reliable since both species are so variable. The following
differences have been noted as tendencies which may be of value. The antennal
and thoracic maculae of H. granulatus are less extensive than is usually true of
H. rudbeckiae. In H. granulatus the scape may be black or with a small, basal
/
24 Contributions in Science No. 180
spot on the lower margin; the pronotal collar may be immaculate or with a
pair of minute yellowish spots at the extreme sides; the pronotal lobes are one-
half, or more, black; the tegulae may be either black or with a small median
spot. In H. rudbeckiae these areas are usually conspicuously maculate, except
in specimens from northern or high altitude sites. The metanotum of both
species is dull, but in H. granulatus it is rugosopunctate, while in H. rudbeckiae
it has fine sparse rugulae and a few distinct scattered punctures. In H. rud-
beckiae the basal zone of the propodeum is coarsely rugose, with the rugae in
the median portion basically parallel. The basal zone of H. granulatus may be
coarsely and irregularly rugulose, or with fine rugulae only on the anterior
half, the posterior half appearing granulose.
The status of this form is enigmatical. Metz described it from three males
from Ormsby County, Nevada. No type was ever designated and so I am
selecting a specimen from the original series, collected by Baker in July, as the
lectotype. This specimen has been deposited in the American Museum of
Natural History. Since the time of its description this bee has never been
regarded as anything other than a variety of H. rudbeckiae, of which it has
been considered a synonym. Its status here as a species is provisional, pending
the acquisition of more material. This form may ultimately prove to be a
hybrid resulting from a cross between H. rudbeckiae and H. cressoni. The
extremes in variation in the sculpturing of the basal zone of the propodeum
are suggestive of H. cressoni and the details of the eighth and ninth ventrites
are intermediate between H. cressoni and H. rudbeckiae. If H. granulatus is
a valid species, its female is currently unknown, but probably will be ex-
tremely similar to that of H. rudbeckiae.
In addition to the three males from Nevada I have seen others from
scattered localities in the Sierra Nevada of California, from Lassen County
to Tulare County, and from the San Gabriel Mountains in Los Angeles
County. I also have a few males from Colorado which seem referrable to this
form; these differ slightly, however, in the shape of the ninth ventrite which is
very similar to that of H. cressoni. In areas where these specimens were taken,
both H. cressoni and H. rudbeckiae are common.
Hylaeus (Hylaeus) maritimus Bridwell
Figures ID; 2D; 3G; 5G.
Hylaeus maritimus Bridwell, 1910. Proc. Haw. Ent. Soc. 4: 159. $ $ .
Hylaeus {Hylaeus) maritimus, Linsley, 1951. IN Musesebeck, et al.,
U.S.D.A. Monogr. 2:1051.
This rare species appears to be largely restricted to the coastal sand dunes
of central California. The two females taken by Timberlake at Mt. Pinos are
the only exceptions, and it is difficult to explain the presence of H. maritimus
in that area.
1970
Studies on North American Bees
25
Both sexes of this species, which is closely related to H. conspicuus, may
be readily recognized by the characters given in the key.
NEW RECORDS (all California) : Sonoma Co.\ 4 $ $ , Wright’s Beach,
August 22, 1950 (J. Linsley; CIS), on Ehogonum. Marin Co.: 1 5 , Pt. Reyes,
April 11, 1959 (G. I. Stage; LACM) ; 9 $ ^ , 2 $ 9 , same locality. May 16,
1958 (D. D. Linsdale; CIS) ; 2 $ $ , same locality and date (J. Powell; CIS) ;
1 9 , same locality. May 15, 1937 (R. C. Dickson; UCR) . San Francisco Co.:
1 $ , San Francisco (Land’s End), July, 1922 (F. X. Williams, UCR) ; 1 $ ,
San Francisco, April 16, 1960 (G. I. Stage and R. R. Snelling; LACM) ; 1 $ ,
San Francisco (Lobos Cr.), June 15, 1960 (G. I. Stage; GIS) ; 1 $ , San Fran-
cisco (Lone Mtn.), July 4, 1920 (F. X. Williams; CAS) ; 1^,49 9 , San
Francisco (sand dunes), April 9 to August 22, 1926 (C. L. Fox; CAS) ; 1 9 ,
Lake Merced, June 5, 1959 (T. R. Haig; UCD) . San Mateo Co.: 1 ^ , Salada
Beach, April 21, 1918 (E. P. Van Duzee; CAS) . Monterey Co.: 1 $ , Pacific
Grove, July 3 (T. D. A. Cockerell; UCR) ; 5 $ ^,109 9 , same locality. May
29, 31, 1954 (M. Wasbauer; CIS) ; 3 $ $ , same locality. May 16, 1959 (G. I.
Stage; GIS, LACM) ; 1 9 , Carmel, July 21, 1923 (L. S. Slevin; CAS) ;2$ $ ,
299, Asilomar, May 10, 16, 1959 (D. J. Burdick; GIS, LACM) ; 1 $ ,
Asilomar, 1 mi. S, May 17, 1959 (D. J. Burdick; LACM) ; 1 $ , Asilomar, 2
mi. S, September 26, 1959 (C. W. O’Brien; LACM) ; 1 5 , same locality. May
17, 1959 (G. I. Stage; GIS) . San Luis Obispo Co.: 1 $ <3,99 9 , Grover
City, July 4, 1956 (E. G. Linsley, P. D. Hurd; CIS), “dunes Assoc.” Santa
Barbara Co.: 1 $ , Refugio Cyn., July 4, 1959 (J. R. Russell: UCD) . Kern
Co.: 1 9 , Mt. Pinos, August 17, 1935 (P. H. Timberlake; UCR), on Hemi-
zonia corymbosa (var. bar clay i) ; 1 9 , same data, except, Grindelia (robusta)
platyphylla (UCR).
Hylaeus (Hylaeus) conspicuus (Metz)
Figures II; 2J; 4C; F.
Prosopis conspicua Metz, 1911. Trans. Amer. Ent. Soc. 37 : 1 14. S 9 .
Hylaeus conspicuus, Cockerell and Sumner, 1931. Amer. Mus. Nov.
490:9.
Cockerell and Sumner (1931) recorded a female from Jackson, Wy-
oming, July 15, 1920, presumably collected by F. E. Lutz. I have on hand
two females collected at Jackson on July 13-17, 1920, presumably by Lutz.
Although these authors were surprised at finding this species in Wyoming,
it is a widely distributed species in the northwestern states. In addition to the
considerable material from California, I have seen specimens of H. conspicuus
from the following states: Nevada (Ormsby Co., Gardnerville, Sparks, Para-
dise V.; AMNH, USNM, UCD, Oregon (Medford, UCD), Washington (Pull-
man; UCD), UC, CAS) , Idaho (Shoshone, Star, Murtaugh, Midvale, 5 mi.
NE Mackay, Parma, Dixie, Boise, Moscow Mts., Sweet, Emery Can., Melba,
8 mi. N Twin Falls, 18 mi. S Moscow; UI), and Wyoming (Jackson; AMNH,
CU).
26
Contributions in Science
No. 180
Hylaeus (Hylaeus) stevensi (Crawford)
Figures 5H.
Prosopis stevensi Crawford, 1913. Can. Ent., 45; 155. $ $ .
Hylaeus {Hylaeus) stevensi, Linsley, 1951. IN Muesbeck, et al, U.S.D.A.
Agr. Monogr. 2:1052. Mitchell, 1960. N. Car. Agr. Exp. Sta. Tech. Bui.
141:61,62, 74-75; figs. 11, 12, 13, 14.
This species is not closely related to any other in the Nearctic fauna, but
is very similar to the Palearctic H. bisinuatus Forster. The two are so closely
related that I know of no reliable external feautres which will separate them.
The male terminalia of the two are very similar to one another. More material
must be studied, however, to determine whether they are conspecific or not.
This species has recently been introduced into Hawaii (Beardsley, 1959),
and is evidently adventive in parts of its range where it is now quite common.
I believe that this species has extended its range appreciably within the past
50 years. The earliest known capture of this species is in 1912 at Fargo, North
Dakota; these specimens form the type series. The species has been collected
in North Dakota on many occasions since then, but remained rare in Hylaeus
samples taken until after 1925. After this date it became increasingly common,
until it is now the species most frequently captured.
The earliest records outside of North Dakota are 1915 (Colorado) and
1916 (Alberta, Canada). Earliest records for other states, proceeding west
from North Dakota to the Pacific coast, are: Wyoming and Idaho (1920),
Utah (1922), British Columbia and Washington (1925) and California
(1933). This species is found in eleven states east of the Mississippi River; the
earliest date of capture which I have seen is 1927 in Massachusetts. In none
of his numerous papers on eastern bees does Robertson mention this bee; this
species is so distinctly different from the remaining eastern Hylaeus that he
certainly would have recognized it as different from any previously described.
Within California the earliest record for this species is 1933 at Sacra-
mento. There are few records in the 1930’s and 1940’s. But, beginning with
1950 there are abundant records, largely from the Central Valley area. My
greatest field acquaintance with this species is in California, so the following
observation must be considered with this point in mind. All of the California
records are within, or on the periphery of, agricultural cultivated areas. I can
affirm, further, that the bulk of my records from most of the area west of the
Continental Divide come from similar situations, as nearly as I have been able
to determine.
This bee is of further interest in that it alone of all the species of Hylaeus,
s. str., exhibits a pronounced tendency toward oligolecty on flowers of the
family Leguminosae. Of 87 specimens bearing floral data, 56 were taken on
Melilotus species. These legumes are introduced from the Palearctic region.
Of the remaining 31 records, 13 are from introduced Palearctic plants in other
families (Amaranthaceae, Umbelliferae, Cruciferaceae, Tamaricaceae, Legu-
1970
Studies on North American Bees
27
minosae) and eight of these are records of males. The remaining 18 records
involve the families Polygonaceae, Asclepiadaccae, Salicaceae and Composi-
tae, all belonging to genera represented in the Palearctic region; six of these
records are of males.
The above data would seem to indicate the possibility that this species was
introduced into the area around Fargo, North Dakota sometime prior to 1912.
Once established there it began to extend its distribution both east and west
and by 1925 apparently reached both coasts; subsequent distribution has been
extended southward, so that this species now occurs in southern California,
Oklahoma and Georgia. It is now a well-established part of our fauna and
appears to be oligolectic on the legume genus Melilotus. If this is, in fact, an
introduced species, it may well prove to be a synonym of a described Palearc-
tic species. This, however, may not be decided for many years yet due to the
extremely chaotic status of the taxonomy of European Hylaeus.
It is possible that this species is a native component which was, prior to
the first quarter of the century, a very rare species. Subsequent to that time the
increased abundance of a suitable food-source plant (the introduced genus
Melilotus) and the conditions of an unbalanced ecology may have permitted
this species first to thrive in its native region and then to spread out to other,
similarly disturbed areas. Of numerous series of specimens reared from soda
straw traps by A. R. Gittins in Idaho, many are referrable to H. stevensi. This
species, then, is strongly apt to exploit artificial nesting possibilities, such as
exist in and around farm buildings.
Excluded Species
In 1966 I included H. gaigei (Cockerell) in this subgenus largely on the
conjectural statement of Mitchell (1960), who had seen the type, that it may
prove to be the female of H. rudbeckiae. Since then I have examined the type,
in the U.S. National Museum, and conclude that it was incorrectly associated
by me with the subgenus Hylaeus. The type, a female, is a small, obscurely
maculate individual. This species belongs to the subgenus Prosopis and may be
a synonym of H. modestus Say.
Paraprosopis
The subgenus Paraprosopis in America north of Mexico is exceeded in
number of species only by the subgenus Prosopis. This group is largely western
in distribution and seems to be centered in the southwest. Although Holarctic,
few species extend into Canada, and none of these very far. Only one species
may be considered truly boreal; species of Paraprosopis are most commonly
encountered in the Transition and Sonoran zones.
This subgenus is a difficult one with which to work and this is particularly
true of the females. The various species are very similar in appearance and
structure and several of the most widely distributed species are quite variable.
28
Contributions in Science
No. 180
Species groups are so poorly delimited that I see little point in attempting to
define them except in very general terms.
The largest group is that of H. wootoni (Cockerell) which includes the
following species: H. color ad ensis (Cockerell), H. cookii (Metz), H. fiori-
danus (Robertson) H. lunicraterius, n. sp., H. nevadensis (Cockerell), H.
personatellus (Cockerell), H. polifolii (Cockerell) and H. wootoni (Cock-
erell). There is considerable variation within the group, but all the species are
superficially very similar in the females. In these the first tergite is densely
and distinctly punctate (excepting H. fioridanus), the integument between
the punctures usually smooth and somewhat shining, the transverse propodeal
Carina is absent (except in H. cookii), the lateral and oblique propodeal cari-
nae often reduced or absent, the facial foveae ending one-third or less of the
distance between the eyes and ocelli, the mesopleurae densely and closely
punctate and the head as broad, or broader, than long. Although the clypeus
is typically immaculate, some females of one species {H. polifolii) may possess
a small preapical spot. The males are similar to their females except for the
usual sexual differences.
Three groups include but a single species each. The H. sonorensis group
contains only H. sonorensis Cockerell, a desert species recognizable by the
head being longer than broad, the elongate pale macula on the female clypeus,
the entire transverse macula on the pronotal collar and the extremely fine
punctures of the second tergite. Another species in which the pronotal macula
is entire is H. asininus (Cockerell and Casad), the sole member of the H.
asininus group. In this group the facial foveae of the female end much nearer
to the ocelli than the eyes, the transverse propodeal carina is present in both
sexes, and the female mandibles are truncate apically. The only other species
with an entire macula on the pronotal collar is H. megalot is. This unique
member of the H. megalot is group may be easily separated from the other
groups in the female by the elongate apical mandibular tooth, the absent or
incomplete oblique propodeal carina and the broadly translucent apical mar-
gins of the first four tergites. The male differs from that of most others by the
finely, sparsely punctate first tergite, the broadly translucent tergal margins
and the longer flagellar segments.
All three of these groups, while apparently related to the H. wootoni
group, seem not to be closely related to one another, although it is interesting
that all occur in arid and semi-arid regions. Of the three, that of H. sonorensis
seems farthest removed from the H. wootoni group as evidenced by the
elongate face and the male apical sternites.
The last group, that of H. calvus, includes four species: H. calvus (Metz),
H. georgicus (Cockerell) and H. timberlakei, n. sp., and the Palearctic species,
H. ater Saunders. In these species the facial foveae of the female end closer
to the ocelli than the eyes, the anterior coxae tend to be angulate or sub-
spinose laterobasally, the mesopleurae are shining and finely, sparsely punc-
1970
Studies on North American Bees
29
tate, the female genae are broader than the eyes in profile, the female clypeus
is usually distinctly maculate and the first tergite of the female is finely and
sparsely punctate.
The females of several of the most common western species are very
difficult to identify with certainty. The three most consistently difficult species
are H. coloradensis (Cockerell) , H. polifolii (Cockerell) and H. wootoni
(Cockerell) . Slight, but apparently consistent, differences in the shape of the
eyes have been utilized. In the full frontal aspect the Eye Length (EL) is
measured to the nearest tenth of a millimeter; the Frontal Eye Width (FEW)
is measured across the eye to the inner margin at the level of the antennal
sockets (fig. lOA) . Similar measurements are made in lateral view. Here,
however. Lateral Eye Width (LEW) is measured at its greatest point. Con-
siderable use has been made of the presence or absence of the oblique
propodeal carina as a supportive feature. This carina is best seen when the
dorsal surface of the propodeum is viewed obliquely from above and cephalad
of the propodeum.
The upper termination point of the facial foveae has also been frequently
used in the key to the females. The position of termination is determined by
viewing the top of the head from above. Measurement is made from the lateral
ocellus to the nearest point of the inner eye margin and the position of the
uppermost point of the fovea made on the basis of this line.
The method of determining the Facial Width (FW) used in previous
papers on this genus {i.e., the distance between the inner eye margins at the
level of the antennal sockets) was found to be nearly useless within this sub-
genus. A different method for determining facial width has been used below.
In full frontal view, the inner margins of the eyes are seen to be slightly sinuate
at a point a little above the level of the antennal sockets, at the lower end of
the foveae. This sinuation probably corresponds to the ocular sinuses of many
wasps. The measurement for facial width is made at the center of the slight
sinuation. The Facial Length (FL) is measured as in previous papers {i.e., the
distance from the anterior ocellus to the medio-apical clypeal margin). The
Upper Facial Distance (UFD) is the maximal distance between the eye
margins above, while the Lower Facial Distance (LED) is the minimal dis-
tance between the eyes below. The Clypeo-Ocular Distance (COD) is meas-
ured from the laterobasal clypeal angle to the nearest point on the eye margin;
the Basal Clypeal Width (BCW) is the maximal width of the clypeal base; the
Clypeo- Antennal Distance (CAD) is measured from the laterobasal clypeal
angle to the nearest point on the rim of the antennal sockets. The Supraclypeal
Quadrangel (SCQ) is bounded below by the clypeal base, on the sides by the
subantennal sutures and above by an imaginary line connecting the juncture of
the subantennal sutures with the rims of the antennal sockets.
At several places in the key to the females reference has been made to
punctures as being fine or coarse. The use of such poorly expressive terms has
30
Contributions in Science
No. 180
been relegated to the role of supportive features, since such vague expressions
are both subjective and relative. Punctures which appear to be fine on one bee
may appear relatively coarse on another bee of smaller size. In a rough way,
punctures which I refer to as “coarse” are those which exceed 0.025 mm in
diameter and are usually about 0.03 mm in diameter. Fine punctures are
usually less than 0.02 mm in diameter. Such measurements are nearly without
value, however, since the punctures of the mesopleura may vary as much as
50% in diameter measurements. Statements relative to puncture size are to be
interpreted as referring to the largest punctures on the pertinent sclerite. Ex-
amination of females of several different species will soon permit one to
adequately interpret the usage of such terms.
The figures of the eighth and ninth sternites of the males require some
comment. The eighth sternite consists of a complex arrangement of four
membranous lobes, conveniently termed the lateral (or basal) and apical
lobes, basad of which are the apodemal arms. The orientation on the plates
places the apical lobes uppermost and the apodemal arms below; this is the
same arrangement used in preceding papers and will be used in all subsequent
papers on this genus.
The lateral lobes of the eighth sternite are simple and usually possess a
number of simple setae along their lower margins and at the apices. Some-
times the apical portion of this lobe folds back upon itself and consequently
assumes a shape very different from that which is considered normal. The
apical lobes are much more apt to present difficulties because the membrane
is much more delicate and complex, hence subject to more pronounced dis-
tortion. These lobes may fold either along the inner margin or transversely at
the middle. A well defined crease line is evident from the apex of the lateral
arms to the outer margin and the transverse fold usually occurs at this point.
Such folds will, of course, greatly alter the contour of the apical lobes and,
because the membrane is so thin, this fact may easily pass unnoticed. An
example is the figure by Metz (1911; pi. 5, fig. 60) of the eighth sternite of
H. cookii; the distal half of the apical lobe has folded back upon itself, a fact
unsuspected by Metz.
The ninth sternite is a much simpler structure; it is elongate and apically
bifurcate. The apical lobes are thin and are partially sclerotized. The normal
plane of these lobes is more or less vertical, so that a true illustration of this
segment in a normal position would show two thin, apically divergent process-
es. However, since such a view would tend to conceal some characters of use
taxonomically, the figures are traditionally based on slide mounted, and some-
what flattened, specimens in which the apical lobes are even more divergent
than is normal and the plane is angularly distorted. The alternative to such a
procedure is to prepare two figures, one of the true ventral aspect and one of
the lateral aspect. I have here abided by the traditional method of figuring this
segment largely in order than a common ground for comparison between my
1970
Studies on North American Bees
31
figures and similar figures by other authors may be established. Thus, when
comparing these figures with specimens, important features to note are the
width of the basal shaft, the relative length of the lobes and the number and
arrangement of the plumose setae. The shape and degree of divergence of the
apical lobes should be viewed with caution.
The key below includes all North American species described by previous
authors, and recognized by me, as well as two new species described at this
time. There exist a number of undescribed species from the western United
States and Mexico but these are represented by too few specimens for me to
wish to describe them at this time. If and when suitable series become available
they will be described in a future part of this series, together with a new key.
KEY TO NEARCTIC SPECIES OF PARAPROSOPIS
1. Females; antennae twelve-segmented 2
Males; antennae thirteen-segmented 15
2. Gaster without evident punctures, first tergite shining, polished; basal
zone of propodeum as long as, or longer than, scutellum, much longer
than metanotum; facial foveae ending much nearer ocelli than eyes
georgicus (Cockerell)
Gaster with distinct punctures, at least on first tergite, the tergite pol-
ished or not; basal zone of propodeum shorter than scutellum, little, if
any, longer than metanotum; facial foveae variable 3
3. Apical mandibular tooth elongate, as long or longer than its basal
width, extending much beyond the well-defined inner tooth, apex of
mandible thus acute; oblique propodeal carina absent or incomplete;
facial foveae ending slightly nearer ocelli than eyes; first tergite polished,
shining, punctures fine, separated by a puncture diameter or more;
apical margins of tergites 1-4 broadly translucent yellowish
megalotis Swenk and Cockerell
Apical mandibular tooth much shorter than basal width, hardly ex-
tending beyond the poorly defined inner tooth, apex of mandible thus
truncate; oblique propodeal carina present or not; facial foveae vari-
able; tergal surface variable; tergites usually without pallid apical
margins 4
4. Facial length 1.4 times facial width, oblique propodeal carina absent,
lateral carina absent or obsolescent; facial foveae ending near mid-
point between eyes and ocelli; punctures of first tergite very fine, sep-
arated by about a puncture diameter; second to fifth tergites duller,
densely micropunctate and with fine appressed pubescence; clypeus
with longitudinal macula; pronotal collar with complete transverse
macula sonorensis Cockerell
Facial length 1.3 or less, times facial width; oblique and lateral propo-
deal carinae usually present; punctures of tergites variable, first tergite
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No. 180
frequently coarsely punctate; clypeus usually immaculate; pronotal
collar usually with macula, when present, interrupted in middle 5
5. Outer margin of fore coxae abruptly produced near base, forming a
triangular to subspinose process; facial foveae ending nearer ocelli than
eyes; mesopleura distinctly shining, with well-separated fine punctures;
first tergite finely, sparsely punctate; clypeus with longitudinal macula
...calvus (Metz)
Outer margin of fore coxae not abruptly produced near base; remain-
ing characters without the above combination 6
6. Basal zone of propodeum abruptly declivious behind, with distinct
transverse carina separating the more or less horizontal anterior face
from the vertical posterior face; metanotum densely tesselate, with
scattered distinct punctures; wings pale whitish, veins yellowish to
light brown 7
Without above combination of characters 8
7. Smaller species, facial length not over 1.2 mm.; second tergite sharply,
closely punctate; facial foveae ending nearer eyes than ocelli, or mid-
way between; macula on pronotal collar interrupted in middle
cookii (Metz)
Larger species, facial length 1,3 mm. or more; second tergite with very
fine obscure punctures or nearly impunctate; facial foveae ending nearer
ocelli than eyes; macula on pronotal collar usually entire
- asininus (Cockerell and Casad)
8. First tergite smooth, punctures extremely fine, sparse to scattered;
second tergite without obvious discernible punctures; mesopleura dull
to moderately shining, with fine punctures separated by a puncture
diameter or more; metanotum dull, without evident punctures
fioridanus (Robertson)
Without above combination of characters 9
9. Mesopleura moderately shining, with fine punctures separated by a
puncture diameter or more; propodeal disc with coarse oblique rugae
convergent toward mid-line, metanotum with coarse quadrate areolae at
extreme base; clypeus with longitudinal median stripe timberlakei Snelling
Mesopleura dull to slightly shining, punctures variable, usually sep-
arated by about one-half a puncture diameter; propodeal disc usually
without rugae, when rugae are present they are irregular; metanotum
either not areolate basally or areolae extremely fine, appearing as a
band of quadrate punctures; clypeus immaculate or with small pre-
apical spot 10
10. Facial length 1.3 times facial width; oblique propodeal carina present,
area above carina coarsely reticulate; transverse propodeal carina
present lunicraterius Snelling
Facial length no more than 1.2 times facial width; oblique propodeal
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Studies on North American Bees
33
Carina present or absent, area above usually not coarsely reticulate;
transverse propodeal carina usually absent 11
11. Small species, facial length not exceeding 1.2 mm., usually about 1.1
mm.; facial length 1.1 times facial width; facial foveae ending about
one-third of distance between eyes and ocelli; oblique propodeal carina
usually present and area above coarsely VQ\.\cu\?iiQ..nevadensis (Cockerell)
Larger species, facial length 1.3 mm. or more; facial length 1.2 times
facial width; facial foveae ending about midway between eyes and
ocelli; oblique propodeal carina and area above variable.... 12
12. Metanotum flattened, densely tesselate, but with distinct punctures
nearly as large as those of scutellum; scutellum distinctly flattened,
shinier than mesoscutum, more finely punctate, the punctures sep-
arated by a puncture diameter or slightly more; lateral face marks re-
duced, frequently absent; erect hairs of mesoscutum sparse, short, less
than half as long as those of scutellum personatellus (Cockerell)
Metanotum at least slightly convex, roughened, without evident punc-
tures; scutellum slightly convex, not much duller than mesoscutum,
punctures dense, nearly as large as those of mesoscutum; lateral face
marks usually filling most of area between eye and clypeus; erect hairs
of mesoscutum variable 13
13. Larger species, head width 1.6 mm. or more; oblique propodeal carina
evanescent or absent; lateral propodeal carina usually evanescent above;
clypeus always immaculate; EL 4. 5-4. 6 X ..coloradensis (Cockerell)
Smaller species, head width 1.5 mm. or less; oblique and lateral propo-
deal carina variable, but often present; clypeus often maculate; EL
3.3-4.5 X FEW 14
14. Oblique propodeal carina often absent, sometimes present but evanes-
cent; clypeus immaculate wootoni (Cockerell)
Oblique propodeal carina usually present and complete; clypeus often
with preapical spot in middle polifolii (Cockerell)
15. Basal zone of propodeum twice as long as metanotum, as long or longer
than scutellum; first tergite microtesselate, impunctate or with very fine
scattered punctures; scape 1.2-1. 3 times longer than greatest width
..georgicus (Cockerell)
Basal zone of propodeum usually less than twice as long as metanotum,
usually much shorter than scutellum; first tergite polished or tesselate,
as a rule with abundant punctures, scape variable 16
16. Head slightly longer than wide; EL 0.97 times HW, 1.17-1.20 times
EL, 1.46-1.50 times UFW; oblique propodeal carina absent, lateral
carina absent or evanescent; pronotal collar frequently with complete
stripe; scape half as broad as long sonorensis Cockerell
Head no longer than broad, usually broader than long; remaining
characters not in above combination 17
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No. 180
17. Eyes rather strongly converging below, UFW at least 1.85 x LFW...=....18
Eyes less strongly convergent below, UFW no more than 1.80 x LFW,
usually 1.75 or less 21
18. First two tergites with sparse, fine punctures, surface shining, with
strong bluish iridescence; FL 2.6, or more, times LFW
fioridanus (Robertson)
First two tergites with coarse, distinct punctures mostly separated by a
puncture diameter or less, surface variable, but never with bluish
iridescence; FL 2.5, or less, times LFW. 19
19. Transverse propodeal carina absent, oblique carina poorly developed,
basal zone narrowly rounded onto declivious face; second submarginal
cell on marginal cell longer than adjacent portion of first submarginal
cell; FL no more than 2.3 x LFW polifolii (Cockerell)
Transverse and oblique propodeal carinae well-developed, basal zone
abruptly declivious behind transverse carina; second submarginal cell
on marginal cell shorter than adjacent portion of first submarginal cell;
FL at least 2.50 x LFW 20
20. Smaller species, HW not exceeding 1.0 mm.; basal, horizontal portion
of propodeum nearly twice as long as metanotum, little shorter than
scutellum cookii (Metz)
Larger species HW 1.28 mm. or more; basal, horizontal portion of
propodeum little, if any, longer than metanotum, about half as long as
scutellum asininus (Cockerell and Casad)
21. First tergite finely, sparsely punctate, the punctures often obscured by
dense tesselation 22
First tergite coarsely, closely punctate, surface usually shining, punc-
tures always clearly defined ..23
22. First tergite densely tesselate, rather dull, the punctures very obscure;
lateral face marks ending abruptly at level of upper margin of antennal
sockets (fig. 8K) or narrowly extended along inner orbits; labrum
black or with median macula; flagellar segments beyond second slightly
longer than broad... calvus (Metz)
First tergite shining, the punctures distinct; lateral face marks slightly
expanded above, and ending well above level of upper margin of
antennal sockets (fig. 9F); labrum wholly yellowish; flagellar segment
beyond second at least as broad as long ..megalot is (Swenk and Cockerell)
23. COD 0.85 X BCW; supraclypeal quadrangle distinctly wider than long;
lateral face marks short, ending at or below upper margin of antennal
sockets; supraclypeal area usually immaculate, but sometimes with an
obscure, irregular macula nevadensis (Cockerell)
COD at least equal to BCW; supraclypeal quadrangle often as long as
wide; lateral face marks usually ending well above level of upper margin
of antennal sockets; supraclypeal area usually with a large, quadrate
1970
Studies on North American Bees
35
macula 24
24. COD 1.45 X BCW or less; HW less than 1.2 mm.; BCW 1.20-1.25 x
CAD wootoni (Cockerell)
COD : : BCW variable; HW more than 1.3 mm.; BCW : : CAD
variable 25
25. BCW less than CAD; COD more than BCW; sides of face usually
strongly depressed below (fig. 8F) coloradensis (Cockerell)
BCW equal to or exceeding CAD; COD : BCW; sides of face never
depressed below 26
26. CAD more than BCW; supraclypeal mark always present distinctly
quadrate; lateral face marks narrowed toward inner eye margin (fig.
8J) timberlakei Snelling
CAD = BCW; supraclypeal mark often reduced or absent; lateral face
marks ending broadly above (fig. 81) lunicraterius Snelling
Hylaeus (Paraprosopis) wootoni (Cockerell)
Figures 7A; 8A; lOB; HE.
Prosapis wootoni Cockerell, 1896. Psyche 7 (sup.): 26.
Prosapis divergens Cockerell, 1896. Op. cit.: 29. 5. NEW SYNONYM.
Prosopis wootoni, Metz, 1911. Tarns. Amer. Ent. Soc. 37:144.
Prosopis cookii Metz, 1911. Trans. Amer. Ent. Soc. 37:139-141. $, not
<$ .
Hylaeus wootoni, Cockerell and Sumner. Amer. Mus. Nov. 490:6.
Hylaeus divergens, Cockerell and Sumner. Op. cit.: 7-8.
Hylaeus perparvus Cockerell and Sumner, 1931. Op. cit.: 10-12. $.
NEW SYNONYM.
This is one of the more widely distributed species of the subgenus
Paraprosopis. Throughout its range it varies considerably in both size and
structure; one of these variants was named by Cockerell {H. divergens) . Metz
reduced the name to the synonymy of H. wootoni, but twenty years later
Cockerell and Sumner resurrected it on the basis of supposed genitalic differ-
ences as well as a slight difference in the shape of the lateral face marks. These
claimed differences will not hold up when one considers the range of variation
in a long series of specimens from many localities. Thus, one finds specimens
with the genitalic characters of H. wootoni and the face marks of H. diver-
gens, and vice verse. Furthermore, these varying combinations are usually
found to occur together in a single area. The differences noted are therefore
considered merely to be due to variation in the range exhibited by a single
species, H. wootoni. Unless there are other, more stable, characters which I
have not discovered, there seems to be no reason for the continued recognition
of H. divergens.
I have examined the types of H. cookii, which are in the collection of
36
Contributions in Science
No. 180
J K
Figure 7. Hylaeus {Paraprosopis) spp., female faces: A, H. wootoni; B, H. flori-
d anus', C, H. polifolii', D, H. nevadensis; E, H. cookii', F, H. coloradensis’, G, H. per-
sonatellus; H, H. sonorensis', I, H. lunicraterius; J, H. timberlakei; K, H. calvus.
Figures by Ruth A. DeNicola.
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Studies on North American Bees
37
J K L
Figure 8. Hylcieus (Paraprosopis) spp., male faces: A, H. wootoni; B, H. fioridaniis',
C, H. polifolii', D, H. nevadensis; E, H. cookii; F, H. coloradensis; G, H. personatel-
liis; H, H. sonorensis; I, H. liinicrateriiis; J, H. timberlakei; K, H. calvus; L, H. poli-
folii, melanic form. Figures by Ruth A. DeNicola.
38
Contributions in Science
No. 180
P. H. Timberlake, and find that the female allotype cannot be separated from
that sex of H. wootoni, although the type male represents a valid species (see
below) .
The holotype male of H. perparvus Cockerell and Sumner has been ex-
amined. This specimen is a typical male of H. wootoni. The supposed differ-
ences in the genitalia are due to distortion which resulted from extreme flat-
tening during the preparation of the slide. The distal portions of the lateral
wings of the eighth ventrite are curled upward on the mount; this segment is
at the edge of the balsam and difficult to see clearly. The ninth ventrite falls
well within the range of variation attributed to H. wootoni.
Specimens have been examined from the following localities: UNITED
STATES. Nebraska (Monroe Can. in Sioux Co.; UN), Colorado, New Mexi-
co, Arizona, Utah (Emigration Can. and Big Cottonwood Can. in Salt Lake
Co., Ogden; UCD) , Nevada, California, Oregon (Tumalo Res. and Sisters in
Deschutes Co.; UCD, CIS), Idaho (Craters of the Moon, Moscow Mts.,
Starkey, Ahsahke, 8 mi. E., Whitebird, 7 mi. N., Rock Cr., 12 mi. S., in Twin
Falls Co., Twin Spr., 2.5 mi. W., Emery Can. in Cassia Co.; UCD, UI),
Wyoming (Little Goose Can. near Sheridan; AMNH, CU) . MEXICO. Chi-
huahua (Madera, San Jose de Babicora; AMNH). CANADA. British Colum-
bia (Revelstoke; CU).
Hylaeiis (Paraprosopis) floridanus (Robertson)
Figures 7B; 8B; IOC; IIF.
Prosopis floridanus Robertson, 1893. Trans. Amer. Ent. Soc., 20:273. $ .
Prosopis eulophi Robertson, 1905. Canad. Ent., 37:236. $ $.
Prosopis modestus, Metz, 1911. Trans. Amer. Ent. Soc., 37:121. (in
part)
Hylaeus (Paraprosopis) packardi Mitchell, 1951. Jour. E. Mitchell Sci.
Soc., 67:244. Mitchell, 1960. N. Car. Agr. Exp. Sta. Tech. Bui., 141:62,
77. NEW SYNONYMY.
Hylaeus (Paraprosopis) floridanus, Mitchell, 1960. Op. cit., 61, 62, 77-
78; figs. 11, 12, 14. $ 9.
I have examined the paratype of H. packardi and find that, aside from
the immaculate pronotal collar and tegulae, it does not differ sufficiently from
H. floridanus to justify specific status. Since such maculations are highly
variable within this subgenus I do not feel that this form can be considered a
subspecies either. While I have seen no other specimens with the immaculate
pronotal collar, I have seen some with the tegular spot absent; it is commonly
greatly reduced.
This species is so far known from the eastern United States, from Maine
to Florida. A westward extension occurs along the Great Lakes as far as
Minnesota.
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Studies on North American Bees
39
Hylaeus (Paraprosopis) nevadensis (Cockerell)
Figures 7D; 8D; lOD; IIH.
Prosapis nevadensis Coc]f.QrQ\\, 1896. Psyche 7 (sup.): 32.
Prosopis nevadensis, Metz, 1911. Trans. Amer. Ent. Soc. 37 : 148.
Hylaeus oregonensis Bridwell, 1919. Proc. Haw. Ent. Soc. 4: 160-161.
$ $ . NEW SYNONYMY.
Bridwell based H. oregonensis on the lack of hairs on the eighth ventrite
of the male, which he stated was like that of H. calvus (Metz). I have exam-
ined the holotype specimen, which is in the United States National Museum,
and the paratype male. An examination of the eighth ventrites of these speci-
mens convinces me that there is no justification for continued recognition of
this form. The presence or absence of hairs on this segment appears to be a
variable feature in H. nevadensis and I have seen many degrees of expression
in specimens from widely scattered areas. In all other features H. oregonensis
agrees with material of a more typical nature. The paratype female is insep-
arable from H. nevadensis.
This species occurs at moderate elevations in California, Nevada, Oregon
(Mt. Jefferson, Grayback Camp in Josephine Co., Union Cr. in Jackson Co.;
USNM, UCD), Washington (Seattle; UC), Idaho (Troy, Craters of the Moon,
Whitebird, 7 mi. N., Athol, Granite, Nampa; UCD, UI), and British Colum-
bia (Lilleoet, Robson, Summerland; CU, CNC).
Hylaeus (Paraprosopis) coloradensis (Cockerell)
Figures 7F; 8F; lOF; IIJ.
Prosapis coloradensis Cockerell, 1896. Psyche 7 (sup.) : 7. 5 .
Prosopis coloradensis, Metz, 1911. Trans. Amer. Ent. Soc. 37 : 141. $ $ .
Hylaeus coloradensis, Cockerell and Sumner, 1931. Amer. Mus. Nov.
490:6. $ (in part).
Specimens examined are from the following states and Canadian prov-
inces: UNITED STATES. Wyoming (Yellowstone Natl. Park; AMNH),
Colorado, Utah, New Mexico, Arizona, Nevada, California, Idaho (White-
bird, 7 mi. N., Leadore; UI) , Washington (Mt. Constitution, Pullman, Colfax;
CAS, UC) . CANADA. British Columbia (Revelstoke; CU) .
Hylaeus (Paraprosopis) personatellus (Cockerell)
Figures 7G;8G; 101; 1 IK.
Prosopis personatella Cockerell, 1915. Annals and Magazine of Natural
History (ser. 8) 16:485. 9.
This species was described from a single female from Corona, Colorado,
a locality which I have been unable to locate. Cockerell remarked, however,
that the collection was made above timber-line which suggests a high altitude.
The species has remained virtually unknown since that time although (Linsley,
1951) added California to the distribution. During this study I have seen few
40
Contributions in Science
No. 180
specimens which I can assign to this species, the type of which I have ex-
amined at the U.S. National Museum.
The females which I assign to this species all agree more or less closely
with Cockerell’s original description and with the type. The lateral face marks
are commonly absent; when present they are greatly reduced, restricted to an
irregular stripe or blotch along the lower inner orbits. Since females of other
species of Paraprosopis taken in the same area frequently also have reduced
face marks, this character alone is of little value. The pronotal collar seems
always to be immaculate; the pronotal lobes and tegulae are maculate or not.
As a rule, only the hind tibiae have a yellow basal spot, although some have
such a mark on the fore tibiae.
The females most closely resemble those of H. coloradensis, to which
this species appears to be related. They may be separated from that species
by the distinctly flattened scutellum which is distinctly more shining than the
mesoscutum and with fine, well-separated punctures. The metanotum, too, is
flattened. As in H. coloradensis, it is dull and densely tesselate, but has distinct,
well-separated punctures about equal to those of the scutellum. Females from
areas where this species occurs together with H. coloradensis differ from that
species also in having the erect hairs of the mesoscutum very sparse and
short, so that they are hardly distinguishable from the reclinate hairs. These
erect hairs are much shorter, usually no more than one-third as long, than
those of the scutellum. In the sympatric females of H. coloradensis the erect
mesoscutal hairs are abundant, and are usually as long, or nearly so, as those
of the scutellum.
The presumed male of this species is similar to that of H. coloradensis
but the sides of the face are not depressed as in that species. There are tangible
differences in head shape as may be seen upon comparing figures 8F and 8G.
Although the eyes are apparently more strongly convergent below in H.
personatellus this is not actually the case, as the ratios are very nearly identical
in the two species. The greater eye length of H. personatellus may be used as
a convenient factor. In this species the eye is 3.4 times its greatest width but
twice as long as the lower interorbital distance. In H. coloradensis the eye is
3.8 times longer than its greatest width, but only about 1.7 times longer than
the lower interorbital width. The few males of H. personatellus which I have
seen are all more densely pubescent on the thoracic dorsum than is true of
H. coloradensis. The difference, however, is a matter of degree, and I do not
attribute much significance to this character. Otherwise, the males are very
similar externally and no reliable differences between them have been dis-
covered.
In addition to the type I have seen the following additional specimens:
COLORADO. 1 9, Ouray, 7500-8000', 1-15 July 1897 (H. F. Wickham;
LACM). CALIFORNIA. 1 9, Strawberry Valley, El Dorado Co., 16 Aug.
1912 (E. C. Van Dyke; CAS) ; 1 9 , Sonora Pass, Tuolumne Co., 4 Aug. 1948
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Studies on North American Bees
41
(P. D. Hurd & J. W. MacSwain; CIS); 1 $, Blue Canyon, Sonora Pass,
Tuolumne Co., 10 Aug. 1960 (J. W. MacSwain; CIS); 1 2, same locality, 30
Aug. 1960 (P. D. Hurd; CIS) ;9 S S , Tioga Pass, 7 mi. E., Mono Co., 15 July
1961 (H. V. Daly, G. I. Stage; CIS) ; I S , Tom’s Place, 1 mi. W., Mono Co.,
13 Aug. 1957 (D. D. Linsdale; CIS), on Eriogonum elatum; A $ ^ \ 2 , near
Mono Pass, 12,000', Inyo Co., 15 Aug. 1957 (C. D. MacNeill; CAS) ; 5 $ 2 ,
Mono Pass, Inyo Co., 13 Aug. 1957 (J. Powell, D. D. Linsdale; CIS), one on
Pentstemon newberryi.
Hylaeus (Paraprosopis) polifolii (Cockerell)
Figures 1C; 8C; 8L; lOG; 1 IG.
Prosopis polifolii Cockerell, 1901. Canad. Ent., 33:281. $ . Metz, 1911.
Trans. Amer. Ent. Soc., 37 : 146-148.
Hylaeus polifolii catalinensis Cockerell, 1938. Annals and Magazine of
Natural History, ser. 11, 2:150. $ 2. NEW SYNONYMY.
This common California species is closely related to H. wootoni and
females are apt to be very difficult to separate in the absence of males. The
two species are largely allopatric with overlaps occurring only in some areas
of southern California and along the western portions of the Sierra Nevada.
Females of H. polifolii which possess the preapical clypeal macula are easily
recognized; I have never seen specimens of H. wootoni so maculated. The
population of H. polifolii present on Santa Catalina Island, Los Angeles Co.,
Calif., was given a varietal name by Cockerell; in the females of this popula-
tion, the clypeal macula seems always to be present. Populations from main-
land California may or may not have a maculate clypeus, but this feature is
subject to much variation within a given sample and is, moreover, not cor-
related with distribution. I see nothing to be gained by recognition of this
insular form and have synonymized it under the nominate form.
The facial maculae of the males, too, are subject to considerable varia-
tion. In what I consider to be the basic or “typical” pattern, the clypeus is
entirely yellow, the supraclypeal area is black, and the lateral face marks
terminate above at a point about equal to a socket diameter above the antennal
sockets. One variant form, very rare, exhibits an obscure maculation of vari-
able size on the supraclypeal area. In the second variant form, which is fairly
common, the clypeus becomes darkened along the lateral margins; this in-
fuscation frequently encroaches so extensively onto the clypeus that the
maculation is reduced to an erect median stripe.
Hylaeus {Paraprosopis) cookii (Metz)
Figures 7E; 8E; lOE; 111.
Prosopis cookii Metz, 1910. Trans. Amer. Ent. Soc. 37:139-141.
not 2 .
As indicated above, the female assigned by Metz to this species is H.
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Contributions in Science
No. 180
wootoni; the male type, now in the collection of the University of California
at Riverside, is from Filmore Canyon, New Mexico, collected on Aug. 27 by
T. D. A. Cockerell. Another male, with identical data is in the collections of
the Los Angeles County Museum.
Until the present study was undertaken H. cookii was known only from
the type. It is doubtful if this species could have been recognized from the
original description, or the figures of the genitalia and hidden ventrites as
illustrated by Metz. When I was able to examine the type, I found it to be
identical with a species which P. H. Timberlake and I had independently
assumed to be undescribed. Because of the inadequacies of the original de-
scription, the species is redescribed and the hidden ventrites illustrated.
Diagnosis. — Male: Small species, head width not exceeding 1 mm.; scape
less than half as wide as long; lateral face marks ending well above level of
antennal sockets; frontal punctures coarse, distinct; mesoscutal punctures
distinctly finer than those of frons; punctures of tergite I as coarse as those of
mesoscutum, those of tergite II finer, dense on both segments. Female: Small,
head width not exceeding 1.3 mm.; frontal punctures coarse, distinct; meso-
scutal punctures equal to those of frons; metanotum not rugulose, with
distinct punctures; abdominal punctation similar to that of male, but punc-
tures finer, tergite III distinctly, though finely, punctate.
Male. — Integument black, except the following pale creamy-white: cly-
peus; supraclypeal mark; lateral face marks, ending acutely on eye margin
well above level of antennal sockets; medially interrupted stripe on pronotal
collar; pronotal lobe; spot on anterior half of tegula; small apical spot on all
femora; stripe on anterior tibiae; elongate basal and small apical spots on
middle and hind tibiae; basitarsal and mediotarsal segments. Apical tarsal
segments light ferruginous; tibial spurs whitish. Wings whitish-hyaline, veins
and stigma light ferruginous. Erect pubescence whitish.
Head: UFD 0.76 x FL; LFD 0.52 x UFD; OCD 1.06-1.21 x TFD; basal
width of clypeus 0.77 x distance from laterobasal angle to eye margin; distance
between antennal sockets about equal to a socket diameter, distance from
sockets to eye margin about 0.85 x a socket diameter. Maximum scape width
about 0.42 x maximum length; minimum length of first flagellar segment
about one-half length of second. Maculate portions of face slightly shining,
finely punctate, those of clypeus separated by one-half to a full puncture
diameter, those of lateral part of face somewhat elongate, linearly arranged,
spacing variable; frons coarsely, contiguously punctate, the punctures distinct,
the frons somewhat bulging on either side of midline. Vertex sharply raised
above tops of eyes, coarsely punctate, the punctures crowded and less sharply
defined than on frons.
Thorax: Mesoscutum dull, closely coarsely punctate, the punctures
slightly larger than those of frons, separated by one-third to one-half a punc-
ture diameter; mesoscutellum slightly shining, punctures about equal to those
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Studies on North American Bees
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of mesoscutum, separated by one-half to a full puncture diameter; metanotum
dull, densely tesselate, with punctures finer than those of mesoscutum, sep-
arated by about a puncture diameter; mesopleurae dull, tesselate, punctures
about equal to those of mesoscutum, separated by about one-third to one-half
a puncture diameter. Basal area of propodeum areolate; lateral areas coarsely
punctate, the punctures poorly defined; transverse carina poorly developed,
oblique carina absent; disc densely tesselate, with coarse punctures.
Abdomen: Tergite I shining, smooth, densely punctate, the punctures
smaller than those of mesoscutum, separated by one-half to a full puncture
diameter; tergite II more finely closely punctate, apical one-third (approxi-
mately) , depressed, with scattered micropunctures, apical margin slightly
reflexed; tergite III more finely, sparsely punctate; apical ventrites as
illustrated.
Length, to apex of second tergite, 3. 0-3. 4 mm.; of forewings, 2.45-2.9
mm.
Female. — Similar to male in essential features, except as follows: clypeus
and supraclypeal area dark, lateral face marks more rounded above; UFD
0.72-0.80 X FL; LFD 0.55-0.61 x UFD; OCD 1.02-1.07 x TFD; basal width
of clypeus equal to distance from laterobasal angle to eye margin; distance
between antennal sockets about 1.5 x a socket diameter; distance from sockets
to eye margin slightly exceeding a socket diameter.
Length, to apex of second tergite, 4. 0-4. 3 mm.; of forewings, 3. 2-3. 5 mm.
New records. — Arizona: 1 $ , Rincon Mtn., IX-16-1937 (E. D. Ball) ;
$ $ , Stockton Peak, Pinaleno Mts., V-20-1954 (F. Werner), on Fallugia;
6 $ $ , Madera Can., Sta. Rita Mts., IX-28-1956 (F. Werner), swept from
Aplopappus and Eriogonum; 2 $ $ , Sta. Catalina Mts., V-15-1955 (G. D.
Butler) ; 1 9 , Ramsey Can., Huachuca Mts., IX-8-1955 (F. G. Werner and
G. D. Butler) , on Petalostemum candidum\ 1 9 , west slope Patagonia Mts.,
V-21-1955 (G. D. Butler) , on Prosopis; 3 $ $ , 15 9 9 , same locality, IX-7-
1955 (G. D. Butler and F. G. Werner; all UA) , on Erigeron canadensis;
7 9 9 , Sycamore Can., Santa Cruz Co., IX-7, 8-1961 (G. I. Stage) ; 5
9 9 , Southwest Research Station, IX-3, 4-1959 (G. I. Stage; all LACM) ;
1 $ , same locality, V-14-1956, 1 9 , same locality, VI-14-1956, 1 9 , same
locality, VII-19-1956 (all M. Statham) ; 699, Paradise, 4 mi. W., IX-9-
1950 (W. Gertsch and M. Cazier; all AMNH) ; 1 9 , 7 mi. W. Onion Saddle,
Chiricahua Mts., Cochise Co., IV-10-1954, on Haplopappus gracilis; 299,
Cave Cr., 5400 ft., Chiricahua Mts., Cochise Co., IX-8-1958, on Solidago;
499, Prescott, VI-27-1932, on Asclepias tuberosa; 1 $ , 5 9 9 , same lo-
cality, VIII-30-1930, on Eriogonum; 1 $ , 1 9 , same locality and host, VII-
31-1930 (all P. H. Timberlake; all UCR) ; 1 $ , Painted Desert, Apache Co.,
VIII-18-1958 (G. I. Stage; UCR), on Eriogonum aureum. Nevada: 1 $ ,
Eastgate, 1 mi. W., Churchill Co., VIII-1 1-1958 (E. G. Linsley; CIS), at
honeydew on Chrysothamnus sp. California: 12 $ $ , 2 9 9 , Paradise Camp,
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Contributions in Science
No. 180
Mono Co., V-7-1960 (A. E. Menke; UCD); 1 ^ , 1 $ , same locality and
date (F. D. Parker; UCD); 1 $ , Cottonwood Spr., Riverside Co., IV- 12-
1950 (W. F. Barr; UI) ; 10 ^ ^ , Apple Valley, San Bernardino Co., V-10,
12-1955 (W. R. M. Mason; CNC); 5 $$,322, Ivanpah, 12 mi. SE., San
Bernardino Co., V-1-1956 (M. Wasbauer) ; 1 $ , same data, on Salvia; 1 $ ,
same locality and date (B. J. Adelson), on Salvia; A $ $ , same locality and
date (P. D. Hurd; all CIS), 2 on Sphaeralcea; 2 $ $ , same locality and date,
on Rhus trilobata; 1 $ , same locality and date, on Salvia vitosa; 2 $ $ , same
locality and date, on Sphaeralcea ambigua; 1 $ , 3 2 2 , Cactus Flat, 6000 ft.,
San Bernardino Mts., IX-1-1936, on Chrysothamnus stenophyllus; 1 2 ,
Morongo Valley, San Bernardino Co., IX-29-1944, on Eriogonum plumatella;
3 $ 9 , same locality, X-5-1934, on Gutierrezia lucida; 1 9 , same locality,
X-14-1934, on G. lucida; 2 2 2, Yucca Valley, San Bernardino Co., IX-28-
1944, on G. lucida; 1 9 , same locality and host, IX-30-1944; 1 9 , Mt. Home
Cr., San Bernardino Mts., 5000 ft., VIII-14-1934, on Phacelia ramosissima;
299, Andreas Cyn., Riverside Co., X-27-1934, on Solidago calif or nica;
1 9 , Pinon Flat, San Jacinto Mts., VI-4-1934, on Nolina perry i; 1 9 , same
data except, on Eriogonum fasciculatum; 1 $ , Whitewater, Riverside Co.,
IV-22-1928, on E. fasciculatum; 1 2 2, Whitewater Cyn., Riverside Co.,
X-14-1951, on Lepidospartum; 899, Lone Pine Cyn., 6000 ft., San Gabriel
Mts., VII-21-1936, on Tetradymia canescens; 1 9 , same locality and date, on
E. fasciculatum; 1 $ , Surprise Cyn., Panamint Mts., Inyo Co., IV-25-1957
(all P. H. Timberlake; all UCR), on Eriogonum inflatum.
Hylaeus (Paraprosopis) sonorensis Cockerell
Figures 7H; 8H; lOK; IIL.
Hylaeus sonorensis Cockerell, 1924. Proc. Calif. Acad. Sci. (4) 12:530-
531. 2.
Hylaeus sonorensis var. melanorhinus Cockerell, 1924. Op. cit. :531. 2.
NEW SYNONYMY.
Diagnosis: Although the superficial appearance of this species is quite
similar to that of H. as minus (Cockerell & Casad), the shape of ventrites VIII
and IX of the male ally it to H. wootoni (Cockerell) . Both sexes differ from
H. asininus in the more elongate face, denser and finer thoracic punctures and
microscopic punctures of tergite II. From H. wootoni this species differs in
the longer, narrower face, more finely punctate abdomen and more extensive
maculae.
Male. — Integument black, except the following pale creamy-white;
elongate spot at base of mandibles; labrum, clypeus; elongate supraclypeal
mark; paraocular areas and along inner orbits, ending acutely on level mid-
way between anterior ocellus and antennal sockets; entire transverse stripe on
pronotal collar; pronotal tubercules; anterior half of tegulae; apical spot on all
femora; entire fore tibiae except irregular blotch on inner surface; basal half
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Studies on North American Bees
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of middle tibiae; basal three-fifths of hind tibiae; all basitarsi and first hind
mediotarsal segment. Underside of flagellum yellowish-ferruginous. Medio-
and distitarsal segments brownish; wings clear hyaline, veins and stigma
brownish. Pubescence all pale whitish, on abdomen suberect and obscure.
Head: UFD 0.64-0.65 x FL; LFD 0.59-0.60 x UFD; OCD 1.19-1.21 x
TFD; clypeus distinctly narrower basally than distance from latero-basal angle
to eye margin; distance between antennal sockets less than socket diameter,
distance from sockets to eye margins equal to socket diameter; clypeus, supra-
clypeal area and maculate facial area dull, densely tesselate, with scattered
obscure punctures; nonmaculate areas with large, subcontiguous punctures,
the densely tesselate interstices slightly shining; genae shining, punctures
small, separated; minimum length of first flagellar segment about one-half
length of second.
Thorax: Outer margin of fore coxae slightly convex; mesopleural punc-
tures about equal to those of frons, mostly separated by about one-half a
puncture diameter, the tesselate interstices more shining than frons; mesoscu-
tal punctures equal to those of frons, subcontiguous, interstices slightly shin-
ing; mesoscutellar punctures larger than mesoscutal, spacing variable, but
denser medially; metanotum not at all shining, very densely tesselate, with a
few, coarse, deep punctures. Basal area of propodeum with a few irregular
striae, the lateral ones transverse; latero-basal area roughened, shining, im-
punctate; lateral area slightly shining, with coarse, close punctures; oblique
Carina absent; lateral carina feebly developed, absent above lower half of disc.
Abdomen: Tergite I micro-tesselate, slightly shining, with fine punctures
mostly separated by about a puncture diameter; tergite II more distinctly tesse-
late, punctures very fine, obscure, separated by about a puncture diameter;
remaining tergites tesselate, slightly shining, with a few, scattered, obscure
punctures; ventrites VIII and IX as in figures lOK and 1 IL.
Measurements: Body length (front of vertex to apex of second tergite),
3.5 to 3.8 mm.; forewing length, 2.9 to 3.1 mm.
Female. — Similar to male in sculpture and color, except as follows :
mandibles and labrum entirely dark; clypeus with _L-shaped macula; supra-
clypeal area with small irregular mark; lateral face marks more rounded above;
facial foveae ending about one-third of distance between eyes and lateral
ocelli; UFD 0.69-0.70 x FL; LFD 0.68-0.70 x UFD; OCD 0.94-0.96 x TFD;
distance between latero-basal clypeal angle and inner eye margin 0.75 times
basal width of clypeus; distance between antennal sockets equal to a socket
diameter and to distance between sockets and eyes. Fore basitarsus con-
colorous with medio- and distitarsi, hind basitarsus dark at apex. Punctures of
tergite I a little finer and more separated than in male.
Measurements: Body length (front of vertex to apex of second tergite),
4.5 to 4.7 mm.; forewing length, 3.4 to 3.7 mm.
Both forms of H. sonorensis were described by Cockerell from single
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No. 180
females from Guaymas, Sonora, Mexico; I have examined the types of both
at the California Academy of Sciences and can affirm that they are nonspecific.
Although Cockerell claimed this species to be related to those of tropical
Mexico, this is not true. On the basis of currently available specimens H.
sonorensis ranges from the Death Valley region of California and Nevada
into southern Arizona, thence south into Sonora, Mexico at least as far as
Guaymas. Another extension intrudes into Baja California at least as far as
Bahia de Los Angeles.
The variety melanorhinus was distinguished by the lack of a supraclypeal
mark and the reduction of the clypeal macula to a transverse apical band with
a median upward extension. In the “typical” form the apical third of the
clypeus was pale yellowish with a broad, cuneate upward extension. Such
variants are common in the series of females available and there seems to be
no good reason for the recognition of these variant forms, since the variations
are not correlated with distribution.
Distribution. — MEXICO. Baja California del Norte : 2 $ $ , Bahia de
los Angeles, May 14, 15, 1949 (R. C. Dickson; UCR), one on Prosopis
glandulosa. Sonora: 1 $ , Guaymas, April 8, 1921 (E. P. Van Duzee; CAS,
type of H. s. melanorhinus) ; 1 $ , Guaymas, April 10, 1921 (E. P. VanDuzee;
CAS, type of H. sonorensis) . UNITED STATES. Arizona: 4 $ $ , 2 2 $ ,
Alamo Cyn., Organ Pipe Cactus Natl. Mon., April 15, 1955 (G. D. Butler and
F. G. Werner; UA), on mesquite; 1 2 , same data, except on Encelia (UA) ;
1 2 , Sabino Cyn., Santa Catalina Mts., May 26, 1962 (F. D. Parker and L. A.
Stange; UCD). Nevada: 1 $ , Aurora, 7500', Mineral Co., August 9, 1939
(W. D. Pierce; LACM) , on Scrophularia californica. California : 3 $ $ ,
6 2 $ 5 Surprise Cyn., Panamint Mts., Inyo Co., April 22, 1957, on Erio-
gonum inflatum; 2 $ $ , 9 2 $ , same locality and host, April 23, 1957;
1 $ , same locality, April 23, 1957, on Stanleya pinnata; 3 $ $ , 16 2 2,
same locality, April 25, 1957, on E. inflatum; 1 $ , same locality, April 25,
1957, on Prosopis; 2 $ $ , same locality, April 25, 1957, on S. pinnata;
2 $ $ same locality, April 28, 1957, on S. pinnata; 2 $ $ , same locality,
April 29, 1957, on S. pinnata {all P. H. Timberlake; UCR) ; 14 2 2,9 $ $ ,
same locality, April 24, 1957 (P. D. Hurd and G. I. Stage) , on E. inflatum;
2 $ $ , same locality, April 28, 1953 (P. D. Hurd) ; 3 $ $ , same locality and
date (G. A. Marsh), one on Eucnide urens, two on Eriogonum sp.; 1 2,
Furnace Cr., Death Valley, Inyo Co., March 3, 1953 (J. W. MacSwain) ;
1 2 , Cronise Valley, San Bernardino Co., April 29, 1956 (M. Wasbauer), on
Prosopis; 2 $ $ , Plaster City, 15 mi. N., Imperial Co., March 27, 1961
(C. A. Toschi;^// CIS).
Hylaeus (Paraprosopis) lunicraterius, new species
Figures 71; 81; lOH; 12A.
This species is very closely related to the following species. The males
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Studies on North American Bees
47
are especially similar in details of the hidden ventrites and genitalia.
Diagnosis’. Female with head as broad as long; clypeus immaculate;
facial foveae ending about one-third of distance between eyes and ocelli;
mesopleural punctures separated by less than one-third a puncture diameter,
uniform in size; propodeal disc roughened and with scattered oblique rugulae;
punctures of first tergite dense, coarser than those of second. Male with head
as broad as long; lateral face marks ending abruptly a little above level of
antennal sockets; supraclypeal mark present; lower part of face coarsely
punctate; apical portion of tergites II-VI broadly impressed, margins slightly
reflexed.
Male. — Integument black, with pure white maculae as follows: clypeus;
quadrate supraclypeal mark; lateral marks to a point ending about an antennal
socket diameter above upper margin of antennal sockets; medially interrupted
stripe on pronotal collar; pronotal lobe; anterior tegular spot; minute apical
spot on all femora; complete stripe on anterior tibiae; basal and apical spots
on mid and hind tibiae; all basitarsi, except small ferruginous apical spot;
tibial spurs. Wings hyaline, veins and stigma brown. Scape entirely dark,
flagellum light ferruginous beneath. Erect pubescence whitish.
Head’. UFD 0.71-0.79 x FL; LFD 0.58-0.61 x UFD; OCD 1.05-1.17 x
TFD. Distance from laterobasal clypeal angle to eye margin 1.3 times basal
clypeal width; distance between antennal sockets slightly less than a socket
diameter, distance from sockets to eye margins about equal to a socket dia-
meter. Facial integument somewhat shining, maculate areas moderately
coarsely punctate, punctures separated by one-third to three-fourths a punc-
ture diameter on sides of face, the punctures somewhat elongate, linearly
arranged; clypeal punctures more rounded, separated by about one-half a
puncture diameter; frontal area slightly swollen, coarsely, closely punctate, the
punctures for the most part clearly defined; vertex coarsely punctate laterally,
the punctures a little larger than those of frons; ocellar triangle and occiput in
middle dull, densely tesselate, contiguously punctate. Scape distinctly flattened,
greatest width about one-half its length, median flagellar segments about as
long as broad.
Thorax’. Mesoscutal punctures a little smaller than those of clypeus, sep-
arated by half a puncture diameter or less; mesoscutellar punctures a little
coarser, separated by about half a puncture diameter; metanotum densely
tesselate and roughened, with a few obscure punctures; mesopleural punctures
about equal to those of mesoscutellum, separated by about one-fourth a punc-
ture diameter. Basal area of propodeum coarsely areolate, the areolae finer
laterally; transverse and oblique carinae obsolescent; lateral area roughened,
with some indistinct punctures, a few fine rugae below at the sides.
Abdomen’. Tergite I shining, coarsely punctate, punctures separated by
about a puncture diameter, with scattered micropunctures; tergite II more
finely punctate, duller, punctures separated by one-half to three-fourths a
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No. 180
puncture diameter; apical one-third depressed, very finely and sparsely punc-
tate, apical margin slightly reflexed; ventrites VIII and IX as illustrated
(figures lOH and 12A) .
Length, to apex of second tergite, 3. 8-4. 3 mm.; of forewings, 3.15-3.50
mm.
Female. — Integument black, except yellowish-white maculae as follows:
lateral face marks, ending slightly above level of antennal sockets; inter-
rupted stripe on pronotal collar; pronotal lobe; spot anteriorly on tegulae;
minute apical spot on femora; basal spot on all tibiae; tibial spurs. Underside
of flagellum light ferruginous. Erect pubescence whitish. Wings hyaline, veins
and stigma brown. A very minute ferruginous to yellowish preapical spot
sometimes present on clypeus.
Head: UFD 0.71-0.78 x FL; LFD 0.62-0.69 x UFD; OCD 0.81-0.92 x
TFD. Basal margin of clypeus about 1.8 times as wide as distance from
laterobasal angle to eye margin; distance between antennal sockets about 1.4
times a socket diameter; distance between sockets and eye margin slightly
exceeding a socket diameter. Clypeus reticulate-lineolate, with fine punctures
separated by a puncture diameter or more; supraclypeal area lineolate, with
punctures separated by about a puncture diameter; maculate portions of face
finely punctate, punctures somewhat elongate, linearly arranged; frontal punc-
tures coarse, contiguous, becoming finer toward midline; occiput and ocellar
triangle densely tesselate between coarse, close punctures; genae striatopunc-
tate. Median flagellar segments broader than long.
Thorax: Mesoscutum dull, finely tesselate with punctures subequal to
those of clypeus, separated by one-third a puncture diameter or less; meso-
scutellar punctures about equal to those of mesoscutum, separated by about
one-half a puncture diameter; metanotum somewhat granulose, punctures
subequal to those of mesoscutum, separated by a puncture diameter or more;
mesopleural punctures finer than those of clypeus, separated by about one-
half a puncture diameter; propodeal basal area coarsely areolate, the areolae
finer laterad; transverse carina present by poorly indicated, oblique carina
barely evident; lateral areas strongly roughened; disc obliquely striolate.
Abdomen: Tergite I smooth, shining, finely punctate, punctures sep-
arated by a puncture diameter or more; tergite II smooth, shining, very finely
punctate, punctures separated by one and one-half to two times a puncture
diameter.
Length, to apex of second tergite, 3.95-4.35 mm., of forewings, 3.70-
3.95 mm.
Holotype male, allotype female (Los Angeles County Museum of Natural
History) ; Craters of the Moon National Monument, Idaho, Aug. 21, 1964
(D. S. Horning, Jr.) , on Chrysothamnus naiiseosus.
Paratypes, all from the type locality, collected by D. S. Horning, Jr., dates
ranging from July 3 to Sept. 5: 23 $ $ , 12 $ $ . Host associations: Chryso-
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Studies on North American Bees
49
Figure 9. Hylaeus (Paraprosopis) spp., faces: A, H. peorgiciis, female; B, same,
male; C, H, asininiis, female; D, same, male; E, H. megalotis, female; F, same, male.
Figures by Ruth A. DeNicola.
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No. 180
thamnus viscidiflorus, C. nauseosus, Chamaebatiaria millefolium, Senecio
serra, Eriogonum ovalifolium, Phacelia leucophylla and Populus tremuloides;
two males taken on Aug. 17, 1965 are marked “prey of Ph-2.” Paratypes in
United States National Museum, University of Idaho and the Los Angeles
County Museum of Natural History.
The specific name for this species is derived from the type locality.
Hylaeus (Paraprosopis) timberlakei Snelling, NEW SPECIES
Figures 7J; 8J; lOJ; 12B.
This species bears a strong resemblance to H. calvus in most features.
The male differs from that of H. calvus externally in having the punctures on
the dorsal surface of the first tergite sharply defined, the interspaces smooth
and shining, while in H. calvus the punctures are obscured due to the strongly
tesselated and roughened interspaces. The female of H. timberlakei has the
declivity of the propodeum with coarse, oblique rugulae (absent or poorly
defined in H. calvus) and the punctures of the first tergite are moderately
dense and distinctly larger than those of the second tergite (sparse, finer than
those of second tergite in H. calvus).
Diagnosis'. Female with head broader than long; clypeus maculate; facial
foveae ending slightly less than halfway between eyes and ocelli; punctures of
mesopleura separated by more than a puncture diameter, uniform in size;
rugulae of propodeal declivity coarse, obliquely directed toward mid-line;
punctures of first tergite dense, distinctly larger than those of second tergite.
Male with head slightly broader than long; lateral face marks ending rather
broadly on eye margin about midway between level of antennae sockets and
median ocellus; supraclypeal mark present; punctures on lower part of face
coarse; apical margin of second tergite broadly impressed, very finely and
sparsely punctate.
Male. — Integument black, with creamy-white maculae as follows: cly-
peus; triangular supraclypeal mark; lateral face marks, ending broadly on
inner eye margins, about midway between level of antennal sockets and
middle ocellus; pronotal collar, interrupted medially; spot on pronotal lobe;
anterior spot on tegulae; small apical spot on all femora; complete outer
stripe on anterior tibiae; basal and apical spots on middle and hind tibiae; all
basitarsi (usually with distinct apical infuscation) . Anterior medio- and
distitarsi ferruginous, middle and hind medio- and distitarsi fuscous to black.
Tibial spurs yellowish. Wings, hyaline, veins and stigma brownish. Scape all
black; pedicel with reddish-yellow apical spot beneath; flagellum yellowish
beneath, reddish-brown above. Erect pubescence whitish.
Head'. UFD 0.70-0.77 x FL; LED 0.50-0.61 x UFD; OCD 1.00-1.11 x
TED. Distance from latero-basal clypeal angle to eye margin 1.3 times basal
clypeal width; distance between antennal sockets slightly more than a socket
diameter, distance from sockets to eyes slightly more than a socket diameter.
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Studies on North American Bees
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Facial integument dull, maculate areas coarsely punctate, punctures sep-
arated by a puncture diameter or less; frontal area slightly swollen, coarsely
rugosopunctate; vertex and genae coarsely continguously punctate. Scape
somewhat distinctly flattened, greatest width about one-half its length; median
flagellar segments about as long as broad, or slightly longer.
Thorax: Mesoscutal punctures finer than those of vertex, separated by
half a puncture diameter or less, interspaces tesselate and dull; punctures of
mesoscutellum coarser than those of mesoscutum, about equal to those of
vertex, separated by about half a puncture diameter on disc, apical one-fifth
of the segment dull, densely rugosopunctate; metanotum dull, rugosopunctate;
mesopleural punctures about equal to those of mesoscutellum, separated by
one-fourth a puncture diameter or less. Basal area of propodeum areolate, the
ridges sharply defined by very irregular; lateral area rugulose; disc rugulose
laterally, with a few poorly defined oblique rugae medially; transverse carina
present across basal area, oblique carina poorly defined. Outer margin of fore
coxa subangulate basally to evenly convex.
Abdomen: Tergite I shining, densely punctate, punctures a little smaller
than those of mesoscutum, separated by one-half to three-fourths a puncture
diameter; punctures of tergite II a little finer, separated by about one-half a
puncture diameter or less; apical one-third to one-fourth of tergite II de-
pressed, brownish, transversely lineolate, with scattered very fine punctures;
ventrites VIII and IX as illustrated (figures lOJ and 12B).
Length, to apex of second tergite, 3.8 to 4.3 mm.; of forewings, 2.6 to
3.0 mm.
Female. — Integument black, except for yellow maculae as follows:
median clypeal macula, varying from small preapical spot to broad vertical
stripe, broad lateral face marks, ending slightly above antennal sockets;
medially interrupted stripe on pronotal collar; large spot on pronotal lobe;
triangular spot anteriorly on tegulae; basal spot on all tibiae. Underside of
flagellum light reddish. Erect pubescence whitish. Wings hyaline, veins and
stigma brownish.
Head: UFD 0.77-0.84 x FL; LFD 0.61-0.66 x UFD; OCD 0.81-0.91 x
TFD. Basal margin of clypeus 1.5 times as wide as distance from laterobasal
angle to inner margin; distance between antennal sockets 1.5 times a socket
diameter; distance from socket to eye margin 1.3 times a socket diameter.
Clypeus finely reticulate, with small punctures over most of the disc separated
by about a puncture diameter, becoming denser, coarser, more elongate toward
apical margin; supraclypeal area lineolate, somewhat roughened above, with
scattered obscure punctures; maculate portions of sides of face lineolate, with
fine punctures separated by one-half to a full puncture diameter; frons coarse-
ly, contiguously punctate, becoming finely, but very densely punctate on
vertex, especially in ocellar triangle and behind; genal punctures obscure, a
little larger than those of clypeus. Flagellar segments broader than long.
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Thorax: Mesoscutum dull, finely tesselate with punctures finer than those
of frons, separated by about one-half a puncture diameter on disc; mesoscutel-
lum a little shinier, punctures variable, but mostly about same size as those of
mesoscutum, some a little larger, mostly separated by about a puncture dia-
meter; metanotum dull, rugulose; mesopleura bipunctate, i.e., with punctures
of two sizes, the larger ones equal to those of mesoscutum, punctures variably
spaced, but most separated by about a puncture diameter. Basal zone of
propodeum coarsely areolate; transverse, oblique and lateral carinae distinct;
disc finely rugulose above, the rugulae becoming coarser, oblique, below.
Outer margin of fore coxa evenly convex, without basal angulation or spine.
Abdomen: Tergite I shining, smooth, finely punctate, the punctures sep-
arated by a puncture diameter or more; tergite II shining, very finely reticulate,
punctures finer than those of tergite I, separated by from one to one and one-
half times a puncture diameter; remaining tergites increasingly more distinctly
reticulate, with scattered micropunctures.
Length, to apex of second tergite, 4.5-5. 1 mm.; of forewings, 3.5-
4.0 mm.
Holotype: Male (Los Angeles County Museum of Natural History) ;
Forest Hill, 12 mi. E., Placer Co., Calif., June 24, 1962 (R. R. Snelling), on
Ceanothus. Allotype : Female (University of California, Riverside) ; Tyler,
4.5 mi. W., Nevada Co., Calif., Sept. 27, 1956 (P. H. Timberlake), on
Haplopappus.
Paratypes (all California) : 8 $ $ , same data as allotype (UCR) ; 1 $,
Goodyear’s Bar, Sierra Co., Calif., Aug. 11, 1963 (E. J. Montgomery; UCD) ;
1 $ , Buckhorn, Amador Co., July 25, 1955 (E. I. Schlinger; UCD) ; S $ $ ,
Boca, Nevada Co., June 25, 1961 (F. D. Parker; UCD) ; 1 $ , same locality,
June 26, 1960 (F. D. Parker; UCD) ; 1 $ , same locality, June 26, 1960
(A. S. Menke; UCD) ; 1 $ , Pollock Pines, Eldorado Co., Aug. 9, 1957 (E. I.
Schlinger; UCD) ; 1 ^ , Long Barn, 2.5 mi. W., Tuolumne Co., July 29, 1960
(G. I. Stage; GIS) ; 1 $ , Camp Bob MacBride, 5400', near Pinecrest, Tuo-
lumne Co., Aug. 1, 1952 (R. R. Snelling; LACM), on Solidago californica;
10 ^ ^ , Shasta Co., May 2, 1913 (F. W. Nunenmacher; CIS, LACM) ; 1 $ ,
1 $ , Eagle Creek, Trinity Co., Aug. 12, 1952 (H. L. Hansen; CIS) .
This species is dedicated to Professor P. H. Timberlake, in recognition
of his eminence in apoid systematics, and of the generous assistance he has
given to me over many years. Professor Timberlake had recognized this
species as new and with characteristic generosity permitted me to describe it.
VARIATION : In the females, the clypeal maculation varies a great deal.
Some have the spot so minute as to be barely perceptible and other have it
so developed as to occupy as much as 75% of the total area. The Eagle Creek
female has the rugulae of the propodeal declivity confined to the lower one-
third. The males of this species are remarkably uniform structurally. They do
possess a melanic form which strongly resembles that of such species as
1970
Studies on North American Bees
53
Figure 10. Hylaeiis (Paraprosopis) spp. A, female face, illustrating methods of
obtaining measurements used in text: BCW = Basal Clypeal Width; CAD=:Clypeo-
Antennal Distance; COD = Clypeo-Ocular Distance; EL = Eye Length; EL — Facial
Length; LW — Facial Width; FEW = Frontal Eye Width; LEW = Lower Facial
Width; SCQ = Supra-clypeal Quadrangle; UFW = Upper Facial Width. B-K, sternite
VIII of males: B, H. wootoni; C, H. floridaniis; D, H. nevadensis; E, H. cookii; F, H.
coloradensis; G, H. polifolii; H, H. limicrateriiis; I, H. personatellus; J, H. timber-
lakei; K, H. sonoreusis.
54
Contributions in Science
No. 180
Figure 11. Hylaeiis (Paraprosopis) spp., sternite VIII of males: A, H. megalotis;
B, H. asininiis; C, H. calviis', D, H. georgiciis. E-L, sternite IX of males: E, H.
wootoni; F, H. fioridaniis', G, H. polifolii', H, H. nevadensis; I, H. cookii; J, H. color-
adensis; K, H. personatellus; L, H. sonorensis.
1970
Studies on North American Bees
55
Figure 12. Hylaeiis (Paraprosopis) spp., sternite IX of males: A, H. liinicraterius',
B, H. timberlakei', C, H. megalotis; D, H. asiuinus', E, H. calviis; F, H. georgiciis.
56
Contributions in Science
No. 180
H. calvus and H. polifolii, but may be separated from these by the characters
in the key.
Hylaeus (Paraprosopis) calvus (Metz)
Figures 7K; 8K; IIC; 12E.
Prosopis rudbeckiae, Metz, 1911. Trans. Amer. Ent. Soc., 37:111-114.
9 (in part).
Prosopis calvus Metz, 1911. Op. cit. : 143-144. 5.
This common species ranges from southern Oregon to northern Baja
California. A disjunct population exists in north central Arizona. In California
H. calvus occurs through much of the Upper Sonoran area, along the coast
ranges and the eastern foothills of the Sierra Nevada. Although in flight from
early April until the middle of July it is most abundant during May and June.
The females are often taken in large numbers at flowers of various chaparral
shrubs such as Rhamnus, Eriodictyon, Rhus and Ceanothus.
The following are the marginal localities known to me for H. calvus:
MEXICO. Baja California del Norte: 1 2 , Encinas, 4 mi. S., Sierra San Pedro
Martir, VI-2-1958 (J. Powell; CIS). UNITED STATES. Oregon: 2 $ $,
Griffin Creek, Jackson Co., VI-6-1957 (C. Fitch; UCD). Nevada: 1 $ , 1 $ ,
Verdi, Washoe Co., VI-25-1961 (F. D. Parker; UCD) . Arizona: 1 $ , Parker
Creek, Sierra Ancha, V-4-1947 (H. and M. Townes) ; 4 I $ , Oak
Creek Canyon, Coconino Co., V-16-1947 (H. and M. Townes) ; 1 $ , same
locality and collectors, V-19-1947; 1 $ , 2 $ $ , same locality and collectors,
V-20-1947; 1 9, same locality, VI-26-1950 (L. D. Beamer; all UK).
This is one of the least variable of our species of Paraprosopis and one of
the most easily recognized. In the female the combination of maculate clypeus,
angularly produced laterobasal angle of the anterior coxae, shining, sparsely
punctate mesopleurae and sparsely punctate abdomen are diagnostic. Typical
males may be recognized by the lateral face marks truncate at the level of the
antennal sockets, the finely, sparsely punctate shining mesopleurae and dull,
densely tesselate abdomen. Males of this species, as well as those of H. neva-
densis and H. polifolii, may have the clypeal borders broadly infuscated and
the lateral face marks strongly narrowed above, ending acutely on the eye
margin. Such males of this species may be recognized by the shining meso-
pleurae and dull abdomen with the punctures obscured by the dense tes-
selation.
Hylaeus (Paraprosopis) georgicus (Cockerell)
Figures 9A; 9B; 1 ID; 12F.
Prosopis georgica Cockerell, 1898. Psyche, 7 (suppl.) : 438. $ .
Prosopis georgica var. leeana Cockerell, 1909. Annals and Magazine of
Natural History, (ser. 8) 4:27. 5.
Hylaeus (Metziella) hydrangeae Mitchell, 1951. Jour. E. Mitchell Sci.
1970
Studies on North American Bees
57
Soc., 67:244. 9; Mitchell, 1960. N. Car. Agr. Exp. Sta. Tech. Bui. 141:60,
79; Snelling, 1968. L. A. Co. Mus. Nat. Hist., Contrib. Sci., 144:3, 4. NEW
SYNONYMY.
A careful reappraisal of the characters of H. {Metziella) hydrangeae has
convinced me that this is actually the previously unrecognized female of H.
georgicus. Both sexes were associated by me on the basis of a series of speci-
mens from Texas before I realized that the females were conspecific with
H. hydrangeae. In the Texas material two characteristics are outstanding: the
greatly elongated basal zone of the propodeum and the extremely fine abdomi-
nal punctation. The angulate laterobasal angle of the fore coxa is much more
pronounced in the female than in the male, but this is true also of these sexes
of H. calvus. The propodeal structure of the female, while aberrant for a
Paraprosopis is much more similar to that of other Paraprosopis than it is to
that of the subgenus Metziella. In my paper on that subgenus (Snelling; 1968)
I included H. hydrangeae because at that time there seemed no good reason
not to do so. This is now obviously incorrect inasmuch as the male of this
species is, aside from propodeal structure, a perfectly typical Paraprosopis.
The distribution of this species is limited. From New York it extends
south to Florida and west to south central Texas. Mitchell (1960) recorded
males from Ithaca, New York and Raleigh and Wake Co., North Carolina.
The female was recorded (as H. hydrangeae) from Cruso, North Carolina.
Snelling (1968) recorded a female (as H. hydrangeae) from Ft. Gordon,
Georgia. Cockerell described H. georgicus leeana from males taken by Birk-
mann in Fee County, Texas. Additional specimens, including both sexes, have
been examined which were collected in Fee County by Birkmann. In addition
to the above I have examined the following: 1 $ , Augusta, Richmond Co.,
Georgia, 1 l-IV-1959 (R. R. Snelling; FACM) , on Crataegus’, 1 9, Gadsden
Co., Florida, 1 l-IV-1960 (H. V. Weems; FPB) , on Crataegus’, 1 9 , Hwy.
269, 8 mi. S. Chattahoochee, Gadsden Co., 6-IV-1955 (D. C. L. Gosling &
party; MSU) ; 1 9 , Torreya State Park, Fiberty Co., Florida, 12-IV-1960
(H. A. Denmark; FPB), on Crataegus’, 1 S, Lexington, Lee Co., Texas, 24-
III- 1951 (C. D. Michener; KU), on Salix’, 2 $ s. Tilling, Caldwell Co.,
Texas, 30-III-1951 (L. D. Beamer; KU), on Salix.
Hylaeus {Paraprosopis) megalotis (Swenk and Cockerell)
Figures 9E; F; 11 A; 12C.
Prosopis megalotis Swenk and Cockerell, 1910. Ent. News, 21:69-70.
$ 9.
This very distinctive species was originally described from a series of
specimens from Glen, Sioux County, NEBRASKA and a single female from
Fort Garland, COLORADO. The types were taken on Cleome serrulata. In
addition to the types I have seen specimens of both sexes taken at the type
locality on Aug. 28, 1959 (W. E. LaBerge and O. W. Isakson; UN) on
58
Contributions in Science
No. 180
Solidago. The following are new records for this species. MONTANA: 13
$ 0, Whitehall, Aug. 13, 1931 (J. Nottingham; KU). COLORADO: 1 2,
Blue Mtn., Motfatt Co., Aug. 14, 1966 (G. F. Knowlton; USU). UTAH: 1 $ ,
Castle Dale, Emery Co., Aug. 10, 1957 (C. D. Michener; KU) , on Cleome
serrulata; 2 $ $ , 1 2 9 , Clear Creek Canyon, Box Elder Co., June 11, 1954
(G. F. Knowlton; KU); 1 5, Hanksville, Wayne Co., Sept. 11, 1938 (G. F.
Knowlton, F. C. Harmston; USU) ; 1 2, Eureka, Juab Co., Aug. 6, 1958
(G. E. Bohart; USU), on Chrysothamnus; 8 $ $ , Manila, Daggett Co., Aug.
11, 1962 (G. E. Bohart; USU) . ARIZONA: 1 2 , Painted Desert, Apache
Co., Aug. 18, 1958 (G. I. Stage; LACM), on Eriogonum aureus.
The female of this species is easily recognized by its complete pronotal
macula and the elongate apical mandibular tooth. This tooth is subject to wear,
however, and in older individuals may not be noticeably elongate. The male is
very similar to that of H. asininus but may be separated by the finely and
sparsely punctate first tergite, the head about as broad as long (broader than
long in H. asininus) and by the gradually, rather than abruptly, depressed
apico-lateral portion of the second tergite.
Species of Uncertain Position
Hylaeus (Paraprosopis) suffusus (Cockerell)
Prosapis suffusus Cockerell, 1896. Psyche, 7 (supl.) : 32 $ .
This name is based on a single male from an unknown locality in Nevada.
Although the type should be in the collections of the Academy of Natural
Sciences, Philadelphia, it is apparently lost or destroyed. If the type cannot
be found, it seems best to drop the name altogether since the species cannot be
determined from the original description alone. The present assignment to
the subgenus Paraprosopis is also tentative.
Hylaeus {Paraprosopis) tuertonis (Cockerell)
Prosopis tuertonis Cockerell, 1906. Bui. Amer. Mus. Nat. Hist., 22:
423-424. 2.
This species was described from two females, one from Tuerto Moun-
tain, near Santa Fe, New Mexico, and one from Florissant, Colorado. I have
not seen these specimens, and the original description offers little information
of use in identifying the species. It is probably correctly assigned to Para-
prosoips and seems nearest to H. wootoni, of which it may be a synonym. The
description applies equally well, however, to H. cookii. Until the types can be
studied, the status of this name must remain uncertain.
Acknowledgments
The material used during the preparation of this paper has been made
available to me through the kindness of many individuals who have con-
tributed specimens from the collections in their care. I wish to thank each of
1970
Studies on North American Bees
59
the following, not only for the original loan of specimens, but for their patience
and cooperation during the course of this study: G. E. Bohart, Utah State
University (USU); G. D. Butler and F. G. Werner, University of Arizona
(UA) ; R. L. Fischer, Michigan State University (MSU) ; A. R. Gittins, Uni-
versity of Idaho (UI) ; P. D. Hurd, California Insect Survey, University of
California, Berkeley (CIS) ; K. V. Krombein and G. I. Stage, United States
National Museum (USNM) ; U. N. Lanham, University of Colorado (UC) ;
H. B. Leech and G. I. Stage, California Academy of Sciences (CAS) ; A. T.
McClay and R. O. Schuster, University of California, Davis (UCD) ; C. D.
Michener, University of Kansas (KU) ; H. E. Milliron, Canadian National
Collections (CNC) ; J. G. Rozen, American Museum of Natural History
(AMNH) ; G. I. Stage, personal collection (GIS) ; P. H. Timberlake, Univer-
sity of California, Riverside (UCR) ; M. S. Wasbauer, California Department
of Agriculture (CDA) ; A. Willink, Institute Miguel Lillo, Tucuman, Argen-
tina (IMF) . The facial figures of the various species were prepared by Ruth
A. DeNicola.
Literature Cited
Beardsley, J. W. 1959. Proc. Haw. Ent. Soc., 17:23 (note presented at meeting,
Oct. 13, 1958).
Bridwell, j. C. 1919. Miscellaneous notes on Hymenoptera, with descriptions of
new genera and species. 22. North American Hylaeiis. Hawaiian Entomol.
Soc., Proc., 4:156-162.
Cockerell, T. D. A. 1937. Bees of Alberta. V. Canad. Entomol., 69:126-127.
Cockerell, T. D. A. and R. Sumner. 1931. Rocky Mountain bees. III. The genus
Prosopis. Amer. Mus. Nov., 490: 1-15.
Crawford, J. C. 1913. Notes on some species of the genus Prosopis. Canad.
Entomol., 45; 154-156.
Linsley, E. C. 1951. In Muesebeck, et al., Hymenoptera of America North of
Mexico, Synoptic Catalog. Monog. 2, U. S. Dept. Agric., 1-1420.
Metz, C. W. 1911. A revision of the genus Prosopis in North America. Amer.
Entomol. Soc., Trans., 37: 85-156.
Mitchell, T. B. 1960. Bees of the Eastern United States. N. Car. Agric. Exp. Sta.,
Tech. Bui. 141, 538 pp.
Snelling, R. R. 1966a. Studies on North American bees of the genus Hylaeiis.
1. Distribution of the western species of the subgenus Prosopis with descrip-
tions of new forms. Los Angeles Co. Mus., Contrib. Sci., 98: 1-18.
1966b. Studies on North American bees of the genus Hylaeiis. 2. Description
of a new subgenus and species. Biol. Soc. Wash., Proc., 79: 139-143.
1966c. Studies on North American bees of the genus Hylaeiis. 3. The Nearctic
subgenera. So. Calif. Acad. Sci., Bull., 65: 164-175.
1968. Studies on North American bees of the genus Hylaeiis. 4. The subgenera
Cephalylaeiis, Metziella and Hylaeana. Los Angeles Co. Mus. Nat Hist., Con-
trib. Sci., 144: 1-6.
SwENK, M. H. AND T. D. Cockerell. 1910. New bees of the genus Prosopis.
Entomol. News, 21 : 67-71.
Accepted for publication December 15, 1969.
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* ANGELES
COUNTY
MUSEUM
CONTRIBUTIONS
iTzg. IN SCIENCE
juMBER 181 February 27, 1970
TYPE SPECIMENS OF FOSSIL INVERTEBRATA IN THE
LOS ANGELES COUNTY MUSEUM OF NATURAL HISTORY,
EXCLUSIVE OF PALEOENTOMOLOGY
By Edward C. Wilson and Donald E. Bing
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 sep-
arately, and numbers run consecutively regardless of subject matter. Number 1 was
issued January 23, 1957. The series is available to scientific institutions and scien-
tists on an exchange basis. Copies may also be purchased at a nominal price. Inquiries
should be directed to Virginia D. Miller, Los Angeles County Museum of Natural
History, 900 Exposition Boulevard, Los Angeles, California 90007.
Robert J. Lavenberg
Managing Editor
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. Al-
though priority will be given to manuscripts by staff members, or to papers dealing
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to those specifications listed below and will be examined for suitability by an Edi-
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side the Museum.
Authors proposing new taxa in a CONTRIBUTIONS IN SCIENCE must
indicate that the primary type has become the property of a scientific institution of
their choice and cited by name.
MANUSCRIPT FORM.— (1) The 1964 AIBS Style Manual for Biological
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Virginia D. Miller
Editor
TYPE SPECIMENS OF FOSSIL INVERTEBRATA IN THE
LOS ANGELES COUNTY MUSEUM OF NATURAL HISTORY,
EXCLUSIVE OF PALEOENTOMOLOGY
By Edward C. Wilson' and Donald E. Bing-
Abstract: Type specimens of fossil Porifera, Coelemerata,
Brachiopoda, Bryozoa, Molliisca, Arthropoda, and Echinoder-
mata in the Los Angeles County Museum of Natural History
are listed by the original published names. Paleoentomological
types are not included. The original reference, the museum type
number, the age, the formation, and the locality are given for each
specimen. Figured specimens not of the holotype or paratype
groups are included.
Introduction
This catalogue is published in compliance with Recommendation 72D
of the International Commission on Zoologieal Nomenclature (1964, p. 75).
It includes all the type and figured specimens, as defined below, of fossil
invertebrates in the Los Angeles County Museum of Natural History, except
for the paleoentomological types of W. D. Pierce and Georg Statz. A list of
these will be published separately.
The catalogue is arranged systematically by phyla and classes. Within
these, the genus and species group names, as originally published, are listed
alphabetically. Each entry also contains the author and bibliographic reference
for the lowest species-group name, the type category and number, the age
assignment, the formation (if known), and the locality. We assign a separate
number to every type specimen, unless it is not feasible to distinguish indi-
viduals, as in a rock containing many specimens, in which case the number
refers to a lot. Figure references are included only if our specimen was figured.
A question mark after the figure citation indicates some doubt that the figure
was made from the specimen in our collections. “Missing” means that the
specimen cannot now be found.
Definitions of type categories by Schenk, McMasters, Keen, and Muller
(1956: 5-9) and the International Commission on Zoological Nomenclature
(1964) are followed. In this list, hypotype refers to a specimen figured later
than the original description of the species to which it is referred, or figured
without a species designation. Casts of type specimens are classified like the
originals from which they were made, with the prefix “plasto.” Figured refer-
ences for these refer to the original specimens.
^Curator of Invertebrate Paleontology, Los Angeles County Museum of Natural
History.
-Student Professional Worker, Los Angeles County Museum of Natural History.
1
SMITHSONIAN
INSTITUTION
MR 0 4 1970
Contributions in Science
No. 181
Catalogue
PORIFERA: HYALOSPONGEA
Chancelloria eros Walcott, 1920
MASON, 1935, p. 100, pi. 15, fig. 1.
Hypotype 1290; Cambrian, Cadiz Formation; Marble Mountains, San
Bernardino County, California.
COELENTERATA: ANTHOZOA
Astrangia cf. A. insignifica Nomland, 1916
HERTLEIN and GRANT, 1960, p. 79, pi. 19, figs. 1-4, 7.
Hypotypes 1731 (figs. 1, 7), 1732 (figs. 2-4); Pliocene, San Diego For-
mation; hills south of Tia Juana River, San Diego County, California.
Balanophyllia elegans Verrill, 1864
HERTLEIN and GRANT, 1960, p. 84, pi. 19, figs. 14, 19-21; pi. 24, figs.
4, 5,8, 13, 15-17.
Hypotypes 1738 (fig. 14), 1739 (fig. 19), 1740 (fig. 20), 1741 (fig. 21),
1742 (fig. 4), 1743 (fig. 5), 1744 (fig. 8), 1745 (fig. 13), 1746 (fig. 15),
1747 (fig. 16), 1748 (fig. 17) ; Pliocene, San Diego Formation; hills
south of Tia Juana River, San Diego County, California.
Bergaueria sp.
ARAI and McGUGAN, 1969, p. 93, pi. 18.
Hypotype 1232 (fig. 1), 1233 (figs. 2, 7), 1234 (figs. 3, 6), 1235 (fig. 4),
1236 (fig. 5) ; Cambrian; near Goldfield, Esmeralda County, Nevada.
Dendrophyllia cf. D. oldroydi Faustino, in Oldroyd, 1925
HERTLEIN and GRANT, 1960, p. 82, pi. 19, figs. 5, 6, 15.
Hypotype 1737; Pliocene, San Diego Formation; hills south of Tia Juana
River, San Diego County, California.
Paracyathus stearnsii Verrill, 1869
HERTLEIN and GRANT, 1960, p. 80, pi. 19, figs. 8-13.
Hypotypes 1733 (figs. 8, 9), 1734 (figs. 10, 11), 1735 (fig. 12), 1736
(fig. 13) ; Pliocene, San Diego Formation; hills south of Tia Juana River,
San Diego County, California.
BRACHIOPODA: INARTICULATA
Glottidia albida (Hinds, 1844)
HERTLEIN and GRANT, 1960, p. 89, pi. 19, figs. 18, 24, 25.
Hypotypes 1749 (fig. 18), 1750 (fig. 24), 1751 (fig. 25) ; Pliocene, San
Diego Formation; hills south of Tia Juana River (1749) ; quarry at end of
Arroyo Drive, San Diego (1750, 1751 ), all San Diego County, California.
1970
Type Specimens of Fossil Invertebrata
3
BRACHIOPODA: ARTICULATA
Laqueus californianus (Koch), in Kuster, 1844
HERTLEIN and GRANT, 1960, p. 96, pL 20, figs. 1-3, 5-7.
Hypotypes 1753 (figs. 1-3), 1754 (figs. 5-7); Pliocene, San Diego For-
mation; quarry at end of Arroyo Drive, San Diego, San Diego County,
California.
Laqueus vancouveriensis cliegensis Hertlein and Grant
HERTLEIN and GRANT, 1960, p. 97, pi. 20, figs. 4, 8, 12.
Paratypes 1755 (figs. 4, 8, 12), 1756; Pliocene, San Diego Eormation;
end of Coring Street (1755), 0.2 miles north of Harbor Boulevard and
Tourmaline Street (1756), both San Diego, San Diego County, California.
Or this decipiens Phleger
PHLEGER, 1933, p. 17, pi. 1, fig. 2.
Holotype 1258; Ordovician, Barrel Spring Formation; Barrel Spring
Canyon, Inyo County, California.
Or this minusculus Phleger
PHLEGER, 1933, p. 7, pi. 2, figs. 6, 7.
Syntypes 1 239 (fig. 6?) , 1240 (fig. 7) ; Ordovician, Mazourka Eorma-
tion; Mazourka Canyon, Inyo County, California.
Plectambonites angulatus Phleger
PHLEGER, 1933, p. 18, pi. 1, fig. 1.
Syntypes 1259 (fig. 1), 1260; Ordovician Barrel Spring Formation;
Barrel Spring Canyon, Inyo County, California.
Plectorthis mazourkaensis Phleger
PHLEGER, 1933, p. 9, pi. 2, figs. 3-5.
Syntypes 1244 (fig. 3), 1245 (fig. 5) , 1246, 1247 (fig. 4) , 1 248 ; Ordovi-
cian, Mazourka Eormation; Mazourka Canyon, Inyo County, California.
Plectorthis patulus Phleger
PHLEGER, 1933, p. 10, pi. 2, figs. 1, 2.
Syntypes 1241 (fig. 1), 1242 (fig. 2) , 1 243 ; Ordovician, Mazourka Eor-
mation; Mazourka Canyon, Inyo County, California.
Terehratalia hemphilli Dali, 1902
HERTLEIN and GRANT, 1960, p. 93, pi. 19, fig. 26.
Hypotype 1752; Pliocene, San Diego Formation; end of Coring Street,
San Diego, San Diego County, California.
BRYOZOA: GYMNOLAEMATA
Ceriocava eastoni Woollacott
WOOLLACOTT, 1966, p. 226, figs, la-lc.
Syntypes 1148; Cretaceous, Ladd Formation; Williams Canyon Road,
0.5 mile east of Santiago Canyon Road, Santa Ana Mountains, Orange
County, California.
4
Contributions in Science
No. 181
MOLLUSCA: PELECYPODA
Anatina (?) sp.
PACKARD, 1922, p. 423, pi. 31, fig. 5.
Plastohypotype 2425; Cretaceous; west side of Baker Creek, Orange
County, California.
B.[arbatia] Morsel Gabb
GABB, 1864, p. 216, pi. 32, fig. 286.
Plastoholotype 2421; Cretaceous [Eocene]; San Diego, San Diego
County, California.
Bornia cooki Willett
WILLETT, 1937b, p. 389, pi. 25, figs. 2, 3.
Holotype 1047; Pleistocene; Baldwin Hills, Los Angeles, Los Angeles
County, California.
Cardita hilli Willett
WILLETT, 1944, p. 19, pi. 7.
Holotype 1067; Pleistocene, Palos Verdes Sand; Newport Bay mesa.
Orange County, California.
Cardium {Protocardium) marysvillensis Dickerson, 1916
CLARK and WOODFORD, 1927, p. 94, pi. 15, fig. 12.
Plastoparatype 2433; Eocene, Meganos Formation; north side of Mt.
Diablo, Contra Costa County, California.
Cardium sorrentoensis Hanna
HANNA, 1927, p. 285, pi. 41, fig. 10.
Plastoholotype 2430; Eocene, La Jolla Formation; east side of Rose
Canyon, 2 miles east of La Jolla, San Diego County, California.
Chione picta Willett
WILLETT, 1944, p. 21, pi. 8, figs. B.
Paratypes 1068 (pi. 8, figs. B, destroyed, see SPHON, 1962, p. 57) ,
1954-1975, 2313-2414; Pleistocene, Palos Verdes Sand; Newport Bay
mesa. Orange County, California.
Exputens Uajasensis Clark
CLARK, 1934, p. 270, pi. 37, fig. 14.
Plastoparatype 2432; Eocene; branch of Llajas Canyon, Ventura County,
California.
Leda rosa Hanna
HANNA, 1927, p. 271, pi. 25, fig. 9.
Plastoholotype 2428; Eocene, La Jolla Formation; Rose Creek, San
Diego County, California.
Lucina packi Dickerson
DICKERSON, 1916, p. 484, pi. 36, fig. 12.
Plastoholotype 2422; Eocene; Parson’s Peak, Fresno County, California.
1970
Type Specimens of Fossil Invertebrata
5
Liicina packi Dickerson, 1916
TURNER, 1938, p. 52, pi. 9, fig. 11.
Plastohypotype 2423; Eocene; Simi Valley, Ventura County, California.
Nucula (Acila) semirostrata Grant and Gale
GRANT and GALE, 1931, p. 113.
Paratype 1976; Pliocene; \ Va miles west of San Martinez Grande Can-
yon, boundary of Los Angeles and Ventura Counties, California.
Perna goniglensis Hanna
HANNA, 1927, p. 275, pi. 27, fig. 13 (?).
Plastoholotype 2429; Eocene, La Jolla Formation; sec. 12, La Jolla
Quadrangle, San Diego County, California.
Rochefortia reyana Willett
WILLETT, 1937b, p. 388, pi. 25, figs. 1.
Syntypes 1046 (fig. 1, left) , 1703 (fig. 1, right) , 1704-1728; Pleistocene;
Baldwin Hills, Los Angeles, Los Angeles County, California.
T.[ellina] ooides Gabb
GABB, 1864, p. 157, pi. 22, fig. 135.
Plastolectotype 2420; Cretaceous; west of Martinez, Contra Costa
County, California.
Tellina santana Packard
PACKARD, 1922, p. 426, pi. 33, fig. 4.
Plastoholotype 2424; Cretaceous; crest of Santiago-Aliso divide. Orange
County, California.
MOLLUSC A: GASTROPODA
Acmaea mitchelli Lipps
LIPPS, 1963, p. 3, figs. 2a-d, 3a-z, 3aa-kk, 4a-z, 4aa-kk.
Holotype 1126 (figs. 2a-d, 3a, 4a) , paratypes 1 127 (figs. 3b, 4b) , 1781
(figs. 3c, 4c), 1782 (figs. 3d, 4d), 1783 (figs. 3e, 4e), 1784 (figs. 3f, 4f),
1785 (figs. 3g, 4g), 1786 (figs. 3h, 4h), 1787 (figs. 3i, 4i), 1788 (figs. 3j,
4j), 1789 (figs. 3k, 4k), 1790 (figs. 31, 41), 1791 (figs. 3m, 4m), 1792
(figs. 3n, 4n), 1793 (figs. 3o, 4o), 1794 (figs. 3p, 4p), 1795 (figs. 3q,
4q), 1796 (figs. 3r, 4r), 1797 (figs. 3s, 4s), 1798 (figs. 3t, 4t), 1799
(figs. 3u, 4u), 1800 (figs. 3v, 4v), 1801 (figs. 3w, 4w), 1802 (figs. 3x,
4x), 1803 (figs. 3y, 4y), 1804 (figs. 3z, 4z), 1805 (figs. 3aa, 4aa), 1806
(figs. 3bb, 4bb), 1807 (figs. 3cc, 4cc), 1808 (figs. 3dd, 4dd), 1809 (figs.
3ee, 4ee), 1810 (figs. 3ff, 4ff), 1811 (figs. 3gg, 4gg), 1812 (figs. 3hh,
4hh), 1813 (figs. 3ii, 4ii), 1814 (figs. 3jj, 4jj), 1815 (figs. 3kk, 4kk),
1816-1861 ; Pleistocene; San Nicolas Island, Ventura County, California.
A cteocina anomala Willett
WILLETT, 1945, p. 29.
Holotype 1082, paratypes 1083, 2179-2181 ; Pleistocene (?) ; near shore
of Salton Sea, Imperial County, California.
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Alabina effiae V^xWeii
WILLETT, 1939, p. 202, pi. 54.
Holotype 1061 (pi. 54), paratypes 1729, 1730; Pleistocene, San Pedro
Formation; Hilltop Quarry, San Pedro (holotype); Pleistocene, Timms
Point Formation; Timms Point (paratypes), both Los Angeles County,
California.
Architectonica llajasensis Sutherland
SUTHERLAND, 1966, p. 1, figs. 1, 2.
Holotype 1140; Eocene, Llajas Formation; Las Llajas Canyon, Santa
Susana Mountains, Ventura County, California.
Balds {Balds) tersa Berry
BERRY, 1954, p. 261.
Paratype 2182; Pleistocene, Lomita Formation; Hilltop Quarry, San
Pedro, Los Angeles County, California.
Balds (Vitreolina) ebriconus Berry
BERRY, 1954, p. 265.
Paratype 2188; Pleistocene, Lomita Formation; Hilltop Quarry, San
Pedro, Los Angeles County, California.
Balds {Vitreolina) incallida Berry
BERRY, 1954, p. 264.
Paratypes 2186, 2187; Pleistocene, Lomita Formation; Hilltop Quarry,
San Pedro, Los Angeles County, California.
Balds {Vitreolina) obstipa Berry
BERRY, 1954, p. 262.
Paratypes 2183-2185; Pleistocene, Lomita Formation; Hilltop Quarry,
San Pedro, Los Angeles County, California.
Boetica hertleini Kanakoff
KANAKOFF, 1966, p. 3, fig. 1.
Holotype 1 145 (fig. 1), paratypes 1 146, 2176-2178; Pliocene, Pico For-
mation; mile south of Humphreys railroad station, Los Angeles
County, California.
Calyptraea calabasasensis Nelson
NELSON, 1925, p. 419, pi. 54, figs. 8a, 8b.
Plastoholotype 2437; Eocene, Martinez Formation; Simi Hills, Ventura
County, California.
Caly[p]traea filosa (Gabb, 1866)
CLARK, 1915, p. 560, pi. 65, figs. 23, 24.
Plastohypotype 2438; Miocene, San Pablo Group; Mt. Diablo quad-
rangle, Contra Costa County, California.
Chedvillia steward Clark
CLARK, 1942, p. 117, pi. 19, fig. 8.
Plastoparatype 2439; Eocene, Llajas Formation; Ventura County,
California.
1970
Type Specimens of Fossil Invertebrata
7
Diodora constantiae Kanakoff
KANAKOFF, 1953, p. 67, pis. 12, 13.
Holotype 1089 (pi. 12), paratypes 1094 (pi. 13, missing?), 2191-2312;
Pleistocene, Palos Verdes Sand; Vermont Avenue, south of Sepulveda
Boulevard, Wilmington, Los Angeles County, California.
Exilia taliaferroi Vokes
VOKES, 1939, p. 125, pi. 17, fig. 21.
Plastoholotype 2435; Eocene, Arroyo Hondo Formation; north side of
Domengine Creek, Fresno County, California.
Fusinus {Exilia) lincolnensis (Weaver, 1916)
CLARK, 1918, p. 178, pi. 23, fig. 10.
Plastohypotype 2440; Oligocene, Kreyenhagen Shale; 15 miles north of
Coalinga, Fresno County, California.
Fusinus ucalius Vokes
VOKES, 1939, p. 137, pi. 18, fig. 15.
Plastoholotype 2436; Eocene, Domengine Formation; first large canyon
south of Big Tar Canyon, Kings County, California.
Homalopoma berryi McLean
McLEAN, 1964, p. 132, pi. 24, figs. 7-8.
Holotype 1138 (figs. 7-8), paratypes 1139, 1657-1702; Pleistocene,
Timms Point Formation; Timms Point, Los Angeles County, California.
Kelletia vladimiri Kanakoff
KANAKOFF, 1954, p. 114, pis. 29-31.
Holotype 1097 (pi. 29), paratypes 1098 (pi. 30), 1099 (pi. 31); Plio-
cene, Pico Formation; Vi mile south of Humphreys railroad station, Los
Angeles County, California.
Leptothyra suhobsoleta Willett
WILLETT, 1937a, p. 63, pi. 25.
Holotype 1054; Pleistocene, Timms Point Silt; San Pedro, Los Angeles
County, California.
Macrarene diegensis McLean
McLEAN, 1964, p. 131, pi. 24, figs. 9-14.
Holotype 1136 (figs. 9, 10), paratypes 1137 (fig. 11), 1300 (figs. 12-
14) — 1656; Pliocene, San Diego Formation; hills south of Tia Juana
River, San Diego County, California.
Marginella jewettii nanella Oldroyd
OLDROYD, 1925, p. 24.
Paratypes 2152-2154; Pleistocene, San Pedro Formation; Nob Hill, San
Pedro, Los Angeles County, California.
Mediargo mediocris (Dali, 1909)
TERRY, 1968, pi. 4, fig. 2.
Hypotype 1190; reworked Pliocene in Pleistocene Lomita Marl; Western
Avenue cut, Palos Verdes Hills, Los Angeles County, California.
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Megistostoma caminoensis Hanna
HANNA, 1927, p. 330, pi. 57, figs. 9, 10.
Plastoholotype 2431; Eocene, La Jolla Formation; 0.4 mile south of
Sorrento Station, San Diego County, California.
Micrarionta (Xerarionta) agnesae Kanakoff
KANAKOFF, 1950, p. 85, pi. 27.
Holotype 1088 (pi. 27), paratypes 1864-1953; Pleistocene (?); China
Point, San Clemente Island, Los Angeles County, California.
Mitromorpha galeana Berry
BERRY, 1941, p. 12.
Paratypes 2189, 2190; Pleistocene, Lomita Formation; Hilltop Quarry,
San Pedro, Los Angeles County, California.
Nassarius (Caesia) fossatus forma coiloterus (Woodring), in Woodring,
Bramlette, and Kew, 1946
ADDICOTT, 1965, p. 8, pi. 2, figs. 30, 31.
Hypotypes 1129 (fig. 30), 1130 (fig. 31); Pleistocene, Palos Verdes
Sand; Playa del Rey, Los Angeles County, California.
Nassarius (Catilon) hildegardae Kanakoff, 1956
ADDICOTT, 1965, p. 16, pi. 3, figs. 11, 12.
Hypotypes 1131 (fig. 11), 1132 (fig. 12) ; Pliocene, Pico Formation
(Towsley Formation) ; Vi mile south of Humphreys railroad station, Los
Angeles County, California.
Nassarius hildegardae Kanakoff
KANAKOFF, 1956, p. 113, pi. 31.
Holotype 1 110 (pi. 31 ), paratypes 2097-2151 ; Pliocene, Pico Forma-
tion; Vi mile south of Humphreys railroad station, Los Angeles County,
California.
Nassarius stocki Kanakoff
KANAKOFF, 1956, p. 110, pi. 30A-C.
Holotype 1109 (pi. 30A, C), paratypes 1977 (pi. 30B)— 2096; Plio-
cene, Pico Formation; Vi mile south of Humphreys railroad station, Los
Angeles County, California.
Neadmete schwartzi Mount
MOUNT, 1970, p. 3, fig. 1.
Holotype 1779 (fig. 1), paratype 1780; Pliocene, Fernando Formation;
Newport Bay mesa. Orange County, California.
Neadmete siitherlandi Kanakoff and McLean
KANAKOFF and M3LEAN, 1966, p. 4, figs. 1, 2.
Holotype 1150 (figs. 1, 2), paratypes 1151, 1862, 1863; Pleistocene,
Lomita Marl; San Pedro, Los Angeles County, California.
Odostomia (Amaura) menzola Oldroyd
OLDROYD, 1925, p. 33.
1970
Type Specimens of Fossil Invertebrata
9
Paratype 2173; Pleistocene, San Pedro Formation; Nob Hill, San Pedro,
Los Angeles County, California.
Odostomia (Amaura) timessa Oldroyd
OLDROYD, 1925, p. 35.
Paratypes 2174, 2175; Pleistocene, San Pedro Formation; Nob Hill, San
Pedro, Los Angeles County, California.
Odostomia (Chrysallida) elsiae Willett
WILLETT, 1948, p. 19, pi. 4, fig. 3.
Holotype 1071 (fig. 3), paratype 2415; Pleistocene, Palos Verdes Sand;
Newport Bay mesa. Orange County, California.
Odostomia (Evalea) manca Oldroyd
OLDROYD, 1925, p. 32.
Paratypes 2170-2172; Pleistocene, San Pedro Formation; Nob Hill, San
Pedro, Los Angeles County, California.
Odostomia (Menestho) effiae Willett
WILLETT, 1948, p. 19, pi. 4, fig. 4.
Holotype 1070 (fig. 4), paratypes 2416-2419; Pleistocene; Palos Verdes
Sand; Newport Bay mesa. Orange County, California.
Surcula (Surculites) andersoni Dickerson
DICKERSON, 1914, p. 149, pi. 16, fig. 11.
Plastoholotype 2434; Eocene, Martinez Group; southeast of Muir Sta-
tion, Contra Costa County, California.
Triphora fossilis Willett
WILLETT, 1937a, p. 62, pi. 24.
Holotype 1053; Pleistocene, Timms Point Silt; San Pedro, Los Angeles
County, California.
Triphora kanakoffi Willett
WILLETT, 1948, p. 20, pi. 4, fig. 2.
Holotype 1072; Pleistocene, Palos Verdes Sand; Newport Bay mesa.
Orange County, California.
Turbonilla (Mormula) epiphanea Oldroyd
OLDROYD, 1925, p. 28.
Paratype 2169; Pleistocene, San Pedro Eormation; Nob Hill, San Pedro,
Los Angeles County, California.
Turbonilla (Pyrgolampros) collisella Oldroyd
OLDROYD, 1925, p. 25.
Paratypes 2157, 2158; Pleistocene, San Pedro Formation; Nob Hill, San
Pedro, Los Angeles County, California.
Turbonilla {Pyrgolampros) idae Oldroyd
OLDROYD, 1925, p. 26.
Paratypes 2159-2168; Pleistocene, San Pedro Formation; Nob Hill, San
Pedro, Los Angeles County, California.
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Turbonilla (Strioturbonilla) pecora Oldroyd
OLDROYD, 1925, p. 24.
Paratypes 2155, 2156; Pleistocene, San Pedro Formation; Nob Hill, San
Pedro, Los Angeles County, California.
Turbonilla {Turbonilla) grouardi
WILLETT, 1948, p. 17, pi. 4, fig. 1.
Holotype 1069; Pleistocene, Palos Verdes Sand; Newport Bay mesa.
Orange County, California.
Turritella andersoni subsp. lawsoni Dickerson, 1916
MERRIAM, 1941, p. 77, pi. 12, fig. 4.
Plastohypotypes 2427; Eocene, Llajas Formation; Las Llajas Canyon,
Simi Valley, Ventura County, California.
Turritella schencki Merriam
MERRIAM, 1941, p. 81, pi. 10, fig. 10.
Plastoholotype 2425 (fig. 10) , plastoparatype 2426; Eocene, Tejon For-
mation; Teeny a Creek, Kern County, California.
ARTHROPOD A: TRILOBITA
Acrocephalites ? trifossatus Mason
MASON, 1935, p. 102, pi. 15, figs. 2, 3.
Syntypes 1279 (fig. 2), 1280 (fig. 3) ; Cambrian, Cadiz Formation;
Marble Mountains, San Bernardino County, California.
Alokistocare linnarssoni (Walcott, 1884)
MASON, 1935, p. 104, pi. 15, fig. 4.
Hypotype 1281 ; Cambrian, Cadiz Formation; Marble Mountains, San
Bernardino County, California.
Amecephalus piochensis (Walcott, 1886)
MASON, 1935, p. 105, pi. 15, figs. 5-7.
Hypotypes 1282 (fig. 5), 1283 (fig. 6), 1284 (fig. 7?) ; Cambrian, Cadiz
Formation; Marble Mountains, San Bernardino County, California.
Amecephalus strangulatus Mason
MASON, 1935, p. 106, pi. 15, figs. 8, 9.
Holotype 1285 (fig. 8), paratype 1286 (fig. 9) ; Cambrian, Cadiz For-
mation; Marble Mountains, San Bernardino County, California.
Anomocarella ? spatha Mason
MASON, 1935, p. 107, pi. 15, fig. 10.
Holotype 1287; Cambrian, Cadiz Formation; Marble Mountains, San
Bernardino County, California.
A noria lodensis (Clark, 1921 )
MASON, 1935, p. 109, pi. 15, fig. 11.
Hypotype 1288 (fig. 11), 2442 (fig. 12, missing) ; Cambrian, Cadiz
Formation; Marble Mountains, San Bernardino County, California.
1970
Type Specimens of Fossil Invertebrata
11
Bathyiiriscus howelli Walcott, 1886
MASON, 1935, p. 110, pi. 15, fig. 13.
Flypotype 1289; Cambrian, Cadiz Formation; Marble Mountains, San
Bernardino County, California.
Bathyiiriscus maximiis Mason
MASON, 1935, p. 1 1 1, pi. 15, figs. 14-16.
Syntypes 1291 (fig. 14), 1292 (fig. 15), 1293 (fig. 16) ; Cambrian, Cadiz
Formation; Marble Mountains, San Bernardino County, California.
Biimastus billingsi Raymond and Narraway
RAYMOND and NARRAWAY, 1908, p. 250, pi. 62, figs. 1, 2.
Plastoholotype 2441; Ordovician, Trenton Limestone; Hull, Quebec,
Canada.
Ceraurus infrequens Phleger
PHLEGER, 1933, p. 15, pi. 2, fig. 12.
Holotype 1255; Ordovician, Mazourka Eormation; Mazourka Canyon,
Inyo County, California.
Corynexochiis ? sp.
MASON, 1935, p. 113, pi. 15, fig. 17.
Hypotype 1294; Cambrian, Cadiz Formation; Marble Mountains, San
Bernardino County, California.
Cybeloides calliteles Phleger
PHLEGER, 1933, p. 14, pi. 2, fig. 8.
Holotype 1254; Ordovician, Mazourka Eormation; Mazourka Canyon,
Inyo County, California.
Dorypyge quadriceps (Hall and Whitfield, 1877)
MASON, 1935, p. 1 16, pi. 15, figs. 18-22.
Hypotypes 1295 (fig. 18), 1296 (fig. 19), 1297 (fig. 20), 1298 (fig. 21),
1299 (fig. 22); Cambrian, Cadiz Formation; Marble Mountains, San
Bernardino County, California.
Encrinurus hastula Phleger
PHLEGER, 1933, p. 12, pi. 2, figs. 13, 14.
Syntypes 1251 (fig. 13), 1252 (fig. 14); Ordovician, Mazourka Eorma-
tion; Mazourka Canyon, Inyo County, California.
Encrinurus octonarius Phleger
PHLEGER, 1933, p. 13, pi. 2, fig. 9.
Holotype 1253; Ordovician, Mazourka Eormation; Mazourka Canyon,
Inyo County, California.
Isotelus gigas Dekay, 1824
PHLEGER, 1933, p. 3, pi. 1, figs. 5, 6.
Hypotypes 1277 (fig. 5), 1278 (fig. 6); Ordovician, Barrel Spring Eor-
mation; Barrel Spring Canyon, Inyo County, California.
Isotelus spurius Phleger
PHLEGER, 1933, p. 20, pi. 1, fig. 7.
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Syntypes 1275 (fig. 7), 1276; Ordovician, Barrel Spring Formation;
Barrel Spring Canyon, Inyo County, California.
Lloydia obsoletus Phleger
PHLEGER, 1933, p. 11, pi. 2, fig. 15.
Syntypes 1249 (fig. 15, left), 1250 (fig. 15, right); Ordovician, Mazour-
ka Formation; Mazourka Canyon, Inyo County, California.
PUomerops barrandei (Billings, 1865)
PHLEGER, 1933, p. 16, pi. 2, figs. 10, 11.
Hypotypes 1256 (fig. 10), 1257 (fig. 11); Ordovician, Mazourka Eor-
mation; Mazourka Canyon, Inyo County, California.
Remopleurides occidens Phleger
PHLEGER, 1933, p. 18, pi. 1, figs. 3, 4.
Syntypes 1261 (fig. 3?), 1262 (fig. 4), 1263-1274; Ordovician, Barrel
Spring Formation; Barrel Spring Canyon, Inyo County, California.
ARTHROPODA: CRUSTACEA
Balaniis gregarius (Conrad, 1856)
ZULLO, 1969, p. 6, figs. 3-7, 45.
Hypotypes 1197 (fig. 3, missing), 1198 (fig. 4, missing), 1199 (fig. 5),
1200 (fig. 6), 1201 (fig. 7, missing), 1202 (fig. 45) ; Pliocene, San Diego
Formation; Market Street, east of Euclid Avenue, San Diego, San Diego
County, California.
Balanus kanakoffi. Zullo
ZULLO, 1969, p. 7, figs. 11-12, 15-17, 19-20, 22.
Holotype 1209 (fig. 19), paratypes 1205 (fig. 11), 1206 (fig. 12), 1207
(fig. 15), 1208 (figs. 16-17), 1210 (fig. 20), 1211 (fig. 22); Pliocene,
San Diego Eormation; hills south of Tia Juana River, San Diego County,
California.
Balanus nubilus Darwin, 1854
ZULLO, 1969, p. 8, figs. 8-10.
Hypotype 1 203 (figs. 8-9) , 1204 (fig. 10) ; Pliocene, San Diego Forma-
tion; hills south of Tia Juana River, San Diego County, California.
Balanus proinus Woodring, in Woodring and Bramlette, 1950
ZULLO, 1969, p. 16^figs. 26-28, 30-32.
Hypotypes 1214 (figs. 26, 27), 1215 (fig. 28), 1216 (figs. 30-32); Plio-
cene, San Diego Formation; hills south of Tia Juana River, San Diego
County, California.
Balanus sp., cf. B. pacificus Pilsbry, 1916
ZULLO, 1969, p. 10, figs. 23, 25.
Hypotypes 1212 (fig. 23 ) , 1213 (fig. 25) ; Pliocene, San Diego Forma-
tion; hills south of Tia Juana River, San Diego County, California.
1970
Type Specimens of Fossil Invertebrata
13
Balanus wilsoni Zullo
ZULLO, 1969, p. 10, figs. 33-34, 36, 38-39, 43-45.
Holotype 1224 (fig. 45), paratypes 1217 (fig. 33), 1218 (fig. 34), 1219
(fig. 36), 1220 (fig. 38), 1221 (fig. 39), 1222 (fig. 43), 1223 (fig. 44);
Pleistocene (?), end of Loring Street (1222); Pliocene, San Diego For-
mation; Market Street, east of Euclid Avenue (1217-1221, 1223, 1224),
both San Diego, San Diego County, California.
Cancer branneri Rathbun, 1 926
MENZIES, 1951, p. 170, figs. 2a-2d.
Hypotypes 2448 (figs. 2a, 2b), 2449 (figs. 2c, 2d); Pleistocene, Timm’s
Point Formation; Timm’s Point, Los Angeles County, California.
Cancer jordani Rathbun, 1 900
MENZIES, 1951, p. 169, figs. 5a-5d.
Hypotypes 2446 (figs. 5a, 5b), 2447 (figs. 5c, 5d); Pleistocene; San
Pedro, Los Angeles County, California.
Cancer productus Randall, 1839
MENZIES, 1951, p. 168, figs. 4a-4d.
Hypotypes 2444 (figs. 4a, 4b, missing), 2445 (figs. 4c, 4d, missing);
Pleistocene, San Pedro Formation; San Pedro, Los Angeles County,
California.
Cetolepas hertleini Zullo
ZULLO, 1969, p. 17, figs. 47, 49-50, 53, 55, 58-59, 61-63, 66, 68, 71-72.
Holotype 1230 (figs. 49, 61-62), paratypes 1225 (figs. 47, 59), 1226
(figs. 58, 71-72), 1227 (figs. 50, 63), 1228 (figs. 53, 66), 1229 (figs. 55,
68); Pliocene, San Diego Formation; hills south of Tia Juana River, San
Diego County, California.
Coronula barbara Darwin, 1854 (?)
ZULLO, 1969, p. 21, figs. 73-75.
Hypotype 1231 ; Pliocene, San Diego Formation; hills south of Tia Juana
River, San Diego County, California.
Lepas sp.
ZULLO, 1969, p. 4, figs. 1,2.
Hypotype 1 196; Pliocene, San Diego Formation; hills south of Tia Juana
River, San Diego County, California.
ECHINODERMATA: STELLEROIDEA
Inyoaster bradleyi Phleger
PHLEGER, 1936, p. 82.
Paratypes 1237, 1238; Ordovician, Barrel Spring Formation; Barrel
Spring Canyon, Inyo County, California.
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No. 181
ECHINODERMATA: ECHINOIDEA
Briaster townsendi woynari Hertlein and Grant
HERTLEIN and GRANT, 1960, p. 132, pi. 25, fig. 5; pi. 26, figs. 1-3.
Holotype 1778 (figs. 1-3), paratype 1777 (fig. 5); Pliocene, San Diego
Formation; quarry at end of Arroyo Drive, San Diego, San Diego County,
California.
Dendr aster ashleyi (Arnold), in Arnold and Anderson, 1907
HERTLEIN and GRANT, 1960, p. 117, pi. 21, figs. 2, 6; pi. 22, fig. 1;
pi. 26, fig. 9.
Hypotypes 1762 (fig. 2), 1763 (figs. 6, 9), 1764 (fig. 1); Euclid Avenue
between Federal Boulevard and Home Avenue (1762, 1764); 2400
block of Euclid Avenue (1763), both San Diego, San Diego County,
California.
Dendraster ashleyi ynezensis Kew, 1919
HERTLEIN and GRANT, 1960, p. 118, pi. 21, figs. 4, 5; pi. 22, fig. 2;
pi. 26, fig. 7.
Hypotypes 1765 (fig. 4), 1766 (figs. 5, 7), 1767 (fig. 2); Pliocene, San
Diego Formation; quarry at end of Arroyo Drive (1765) ; Market Street,
14 mile south of Euclid Avenue (1766); 3550 Dove Street (1767), all
San Diego, San Diego County, California.
Dendraster casseli Grant and Hertlein, 1938
HERTLEIN and GRANT, 1960, p. 120, pi. 22, fig. 7.
Hypotype 1768; Pliocene, San Diego Formation; quarry at end of Arroyo
Drive, San Diego, San Diego County, California.
Encope tenuis Kew, 1914
HERTLEIN and GRANT, 1960, p. 126, pi. 23, figs. 7, 13; pi. 24, fig. 11.
Hypotype 1775; Pliocene, San Diego Formation; quarry at end of Arroyo
Drive, San Diego, San Diego County, California.
Eucidaris cf. E. thouarsii (Valenciennes) , in Agassiz and Desor, 1846
HERTLEIN and GRANT, 1960, p. 103, pi. 24, figs. 19-22.
Hypotypes 1757 (fig. 19), 1758 (fig. 20), 1759 (fig. 21), 1760 (fig. 22);
Pliocene, San Diego Formation; hills south of Tia Juana River, San Diego
County, California.
Lovenia hemphilli Israelsky, 1923
HERTLEIN and GRANT, 1960, p. 130, pi. 24, fig. 18.
Hypotype 1776; Pliocene, San Diego Formation; end of Loring Street,
San Diego, San Diego County, California.
Mellita kanakoffi Durham
DURHAM, 1961, p. 5, pi. 2, fig. 2; text-fig. ID.
Holotype 1121 (fig. 2, text-fig. ID), paratype 1122 (missing) ; Pleisto-
cene, Palos Verdes Sand; Newport Bay mesa. Orange County, California
(1121); Lomita Boulevard and Main Street, Wilmington, Los Angeles
County, California (1122).
1970
Type Specimens of Fossil Invertebrata
15
Merriamaster cf. M. israelskyi (Jordan and Hertlein, 1926)
HERTLEIN and GRANT, 1960, p. 122, pi. 23, figs. 12, 14.
Hypotypes 1769 (fig. 12), 1770 (fig. 14); Pliocene, San Diego Forma-
tion; under Snyder Continuation School, San Diego, San Diego County,
California.
Merriamaster pacificus (Kew, 1920)
HERTLEIN and GRANT, 1960, p. 124, pi. 23, figs. 2-4, 6, 11; pi. 25,
fig. 6; pi. 26, fig. 11.
Hypotypes 1771 (pi. 23, figs. 2, 3, 11), 1772 (pi. 23, fig. 4), 1773 (pi.
23, fig. 6) , 1774 (pi. 25, fig. 6; pi. 26, fig. 11); Pliocene, San Diego For-
mation; 0.2 mile north of Harbor Boulevard and Tourmaline Street
(1771, 1772); end of Coring Street (1773, 1774), both San Diego, San
Diego County, California.
Strongylocentrotus purpiiratus (Stimpson, 1857)
HERTLEIN and GRANT, 1960, p. 112, pi. 19, fig. 29.
Hypotype 1761; Pliocene, San Diego Eormation; Pacific Beach, San
Diego, San Diego County, California.
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1970
Type Specimens of Fossil Invertebrata
19
Species Index
agnesae, Micrarionta (Xerarionta), p. 8
albida, Glottidia, p. 2
andersoni, Surcula (Surculites), p. 9
angulatus, Plectambonites, p. 3
anomala, Acteocina, p. 5
ashleyi, Dendraster, p. 14
barbara, Coronula, p. 13
barrandei, Pliomerops, p. 12
branneri, Cancer, p. 13
berryi, Homalopoma, p. 7
billingsi, Bumastus, p. 1 1
bradleyi, Inyoaster, p. 13
calabasasensis, Calyptraea, p. 6
californianus, Laqueus, p. 3
calliteles, Cybeloides, p. 1 1
caminoensis, Megistostoma, p. 8
casseli, Dendraster, p. 14
coiloterus, Nassarius (Caesia) fossatus,
p. 8
collisella, Turbonilla (Pyrgolampros),
p. 9
constantiae, Diodora, p. 7
cooki, Bornia, p. 4
decipiens, Orthis, p. 3
diegensis, Laqueus vancouveriensis,
p. 3
diegensis, Macrarene, p. 7
eastoni, Ceriocava, p, 3
ebriconus, Balcis (Vitreolina), p. 6
effiae, Alabina, p. 6
effiae, Odostomia (Menestho), p. 9
elegans, Balanophyllia, p. 2
elsiae, Odostomia (Chrysallida), p. 9
epiphanea, Turbonilla (Mormula), p. 9
eros, Chancelloria, p. 2
filosa, Calyptraea, p. 6
fossilis, Triphora, p. 9
galeana, Mitromorpha, p. 8
gigas, Isotelus, p. 1 1
goniglensis, Perna, p. 5
gregarius, Balanus, p. 12
grouardi, Turbonilla (Turbonilla),
p. 10
hastula, Encrinurus, p. 1 1
hemphilli, Lovenia, p. 14
hemphilli, Terebratalia, p. 3
hertleini, Boetica, p. 6
hertleini, Cetolepas, p. 13
hildegardae, Nassarius, p. 8
hilli, Cardita, p. 4
howelli, Bathyuriscus, p. 1 1
idae, Turbonilla (Pyrgolampros), p. 9
incallida, Balcis (Vitreolina), p. 6
infrequens, Ceraurus, p, 1 1
insignifica, Astrangia, p. 2
israelskyi, Merriamaster, p. 15
jordani. Cancer, p. 13
kanakoffi, Balanus, p. 12
kanakoffi, Mellita, p. 14
kanakoffi, Triphora, p. 9
lawsoni, Turritella andersoni, p. !0
lincolnensis, Fusinus (Exilia), p. 7
linnarssoni, Alokistocare, p. 10
llajasensis, Architectonica, p. 6
llajasensis, Exputens, p. 4
lodensis, Anoria, p, 10
manca, Odostomia (Evalea), p. 9
marysvillensis, Cardium
(Protocardium), p. 4
maximus, Bathyuriscus, p. 1 1
mazourkaensis, Plectorthis, p. 3
mediocris, Mediargo, p. 7
menzola, Odostomia (Amaura), p. 8
minusculus, Orthis, p. 3
mitchelli, Acmaea, p. 5
morsei, Barbatia, p. 4
nanella, Marginella jewettii, p. 7
nubilus, Balanus, p. 12
obsoletus, Lloydia, p. 12
obstipa, Balcis (Vitreolina), p. 6
occidens, Remopleurides, p. 1 2
octonarius, Encrinurus, p. 1 1
oldroydi, Dendrophyllia, p. 2
ooides, Tellina, p. 5
pacificus, Balanus, p. 12
pacificus, Merriamaster, p. 15
packi, Lucina, p, 4
patulus, Plectorthis, p. 3
pecora, Turbonilla (Strioturbonilla),
p. 10
picta, Chione, p. 4
piochensis, Amecephalus, p. 10
productus. Cancer, p. 13
proinus, Balanus, p. 12
purpuratus, Strongylocentrotus, p. 15
quadriceps, Dorypyge, p. 1 1
reyana, Rochefortia, p. 5
rosa, Leda, p, 4
santana, Tellina, p, 5
schencki, Turritella, p. 10
schwartzi, Neadmete, p. 8
semirostrata, Nucula (Acila), p. 5
20
Contributions in Science
No. 181
sorrentoensis, Cardium, p. 4
spatha, Anomocarella, p. 10
spurius, Isotelus, p. 1 1
stearnsii, Paracyathus, p. 2
stewarti, Chedvillia, p. 6
stocki, Nassarius, p. 8
strangulatus, Amecephalus, p. 10
subobsoleta, Leptothyra, p. 7
Sutherland!, Neadmete, p. 8
taliaferroi, Exilia, p. 7
tenuis, Encope, p. 14
tersa. Balds (Balds), p. 6
thouarsii, Euddaris, p. 14
timessa, Odostomia (Amaura), p. 9
trifossatus, Acrocephalites, p. 10
ucalius, Fusinus, p. 7
vladimiri, Kelletia, p. 7
wilsoni, Balanus, p. 13
woynari, Briaster townsendi, p. 14
ynezensis, Dendraster ashleyi, p. 14
Accepted for publication November 18, 1969.
LOS
ANGELES
COUNTY
MUSEUM
CONTRIBUTIONS
IN SCIENCE
fUMBER 182
April 17, 1970
AQUATIC ADAPTATIONS OF THE WATER MOUSE,
RHEOMYS UNDERWOODI
By Andrew Starrett and George F. Fisler
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
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AQUATIC ADAPTATIONS OF THE WATER MOUSE,
RHEOMYS UNDERWOODI
By Andrew Starrett and George F. Fisler^
Abstract: An adult male Rheomys imderwoodi caught alive
along the Rio Poasito, at 2000 m on Volcan Poas, Alajuela Prov-
ince, Costa Rica, in April, 1966, was maintained in captivity for a
short time. Limited observations on maintenance behavior and
locomotion of this animal are presented. In addition, morphologi-
cal features (body form, tail shape and foot structure) related to
the high degree of specialization for aquatic locomotion shown by
this rodent are described, and some ideas concerning the evolution
of the semi-aquatic habit are discussed in relation to Rheomys
and other small mammals.
Introduction
Information available on the biology of the genus Rheomys is quite
limited. These animals live primarily in and around swift flowing streams of
the Neotropics. Howell (1930) described certain aquatic features of Rheomys
but did not give a full account of the high degree of adaptation to a semi-
aquatic existence which the species R. underwoodi, in particular, exhibits.
Hooper (1968) briefly mentioned some of the aquatic characteristics of
Rheomys in a comparison of R. underwoodi and R. hartmanni, the two species
which occur in Costa Rica, and also included ecological information and
stomach analyses for both. Our capture of a living animal has prompted us
to record some observations on the behavior and morphology of the water
mouse with additional comments on aquatic adaptations in this and other
species.
On the basis of information and materials provided for us by Emmet T.
Hooper, we set 31 traps (18 museum specials, 13 National single door live
traps) along the banks of the Rio Poasito, 2000 m, Volcan Poas, in Alajuela
Province, Costa Rica, on the evening of 9 April 1966. This same area was
trapped on 6 and 7 April 1966 by Hooper and James H. Brown and is described
by Hooper (1968) . Instead of using the method of trapping Rheomys de-
scribed by Stirton (1944), we set seven live traps along relatively quiet eddy
pools just outside white-water areas. Our Rheomys was taken near such a
pool in a trap which was placed on rocks about 7.5 cm above the water line.
The mouth of the trap was about 20 cm from the pool, one meter from any
white water, and 15-20 meters below a small waterfall. Bait used was peanut
butter with sardine meat and oil spread over it, placed on the treadle inside
of the trap. Some sardine oil was smeared on the trap entrance but no bait
was placed outside of the trap. No other mammals were taken in the live traps
■■Both authors: Research Associates, Los Angeles County Museum of Natural
History; and Department of Biology, San Fernando Valley State College, North-
ridge, California 91324.
1
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No. 182
but three Reithrodontomys creper and one Peromyscus nudipes were caught
in the snap traps. About half of the snap traps were placed next to the water.
The water mouse was an adult male having abdominal testes (10 x 18 mm)
with epididymides well developed and caudae protruding into the inguinal
passages. The standard measurements are 300-158-39-7 (mm).
General Behavior
The actions of the Rheomys when first observed were slow and deliberate,
as if it had been in the trap for some time and was weakened, but later it
became more active. It darted about the trap hurriedly, in the manner of a
shrew, yet when quiet, it sat mostly hunched over like a muskrat {Ondatra
zibet hicus) . Stirton (1944) described it as a miniature model of a beaver
( Castor canadensis) . When seated, its hind feet were spread out at an angle
of some 30-40 degrees from the body with its fore feet usually tucked along
its chest. A striking, and probably adaptively significant reaction, was its
tendency to bite at objects with which it came into contact. Even while being
carried in the trap, it nipped at fingers whenever it encountered them at the
edges of the trap as it dashed about. When handled, however, it was docile
and did not bite, and attempts to induce it to nibble fingers were successful
only if the animal blundered into them, as it did not seem to seek out such
prey. The biting reaction on contact with moving objects was also later shown
when the animal was observed swimming. In the capture of prey under water,
where vision and olfaction may be of minimal use, it would be of great ad-
vantage for an animal to be adapted to biting at any moving object with which
it came in contact. The eyes of this rodent are very small and presumably are
of little use, particularly under water, and especially in fast moving, turbulent
streams. The face is rather well endowed with vibrissae, probably advan-
tageous in hunting and orientation, but it was difficult to tell if they were
used as sensors.
When the trap was first observed, all of the bait had been eaten, or, less
likely, washed away. Our captive animal was fed small pieces of sardines
which it ate from the container rather than manipulating the food while sitting
up as do many of the more familiar rodents. It occasionally manipulated larger
chunks of sardine (10-12 mm) with its fore feet, so perhaps the size and con-
sistency of the food determines the manner of feeding. It probably handles
insect larvae with its fore feet. In general, the animal would eat for a minute
or so, pause for 15-20 seconds, and then resume eating. It was first fed about
thirty minutes after the trap was picked up and it ate roughly a small thimbleful
of sardines in the succeeding hour. Attempts at feeding it green vegetation
(leaves and herbs from its habitat) while it was still hungry were unsuccessful.
Stirton (1944) found no evidence that Rheomys was piscivorous but thought
that it was omnivorous and even cannibalistic. Goldman (1920) suggested
that one species may feed on snails. Hooper’s (1968) analysis of contents
from six stomachs of Rheomys from Costa Rica and Guatemala (one stomach
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Adaptations of a Water Mouse
3
empty) shows a diet of primarily aquatic insect larvae. Fish are present in the
Rio Poasito but are not native. Our captive was observed drinking sparingly. It
lapped with its tongue in the familiar manner of many small rodents, directly
off the water surface.
When first seen in the trap, the Rheomys was dry except for its tail, un-
derparts, feet and muzzle. After removal from the stream area, it dried
quickly. Grooming was accomplished in the manner of many rodents, except
that the enlarged hind feet serve as very effective combs for the body fur.
When wet or grooming, or both, as grooming is evidently an important part of
the drying process, the animal moved the skin of the back in a peculiar fashion,
appearing to erect the guard hairs, a grooming as well as a drying aid. This
movement appeared to pass in waves over the hunched back antero-posteriorly
and the reverse. Ail of the time the animal sat much like a muskrat. After
leaving the water, the animal groomed extensively.
Limited swimming observations of the captive Rheomys were made in a
small 27 x 36 cm wash basin. The water was about 8 cm deep and for part of
the time the trap was used as an island although the basin could barely ac-
commodate it. The small size of the container restricted the movement of the
animal. It tried to climb the sides, violently treading water with its hind feet,
maintaining its body about half to two-thirds out of the water. The large hind
feet made tremendous splashes as it attempted to escape, very strikingly
pointing out their propulsive power. Observations of diving were limited but
the process was extremely quick. While swimming on the surface, the mouse
had its head out of the water, the rump under, while the tail was curved con-
cavely with the tip at the water surface. Most of the swimming motion was
with the hind feet with little body undulation except when sculling (noted only
2 or 3 times). These latter motions seemed to give the animal added impetus
for quick acceleration or turning. The fore feet could not be seen while
swimming, indicating that they are not used other than to hold on to
objects when landing, or perhaps to fend off objects while swimming. In all
swimming motions, the animal was extremely quick, presumably an adaptive
feature for feeding under water as well as for swimming in swift moving
streams. Hooper (1968) has reported on the swiftness in movement and
swimming of an animal, undoubtedly Rheomys underwoodi, observed in the
wild.
While in captivity, the animal lived in the trap. Soft tissue and news-
paper given to the Rheomys was slightly shredded but no real nest building
occurred. Excelsior was ignored. Rather, the animal slept mostly on its side
rolled in the typical rodent ball, under the paper. Stirton (1944) mentioned
that water mice probably nest in dead brush and logs lodged in the stream
or under large boulders or other rocks.
After the swimming session on the evening of capture, the animal
appeared to be drying rather slowly. However, by the following morning ( 1 1
April) it was dry and appeared to be in good health. It ate readily (sardines) .
It was placed in a light cloth sack for transport to Los Angeles, California,
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No. 182
but it died in transit. The specimen was preserved as a skin and skull with
the body preserved in fluid and is now in the collection of the Los Angeles
County Museum of Natural History (LACM 28260) .
Morphology
Because of the absence of such information from the literature, and in
order to lay the background for a discussion of the adaptations of this animal
to a semi-aquatic existence, a detailed account of pertinent morphological
characters is here provided. The body is streamlined to a high degree with
essentially no projections other than the hind feet and tail when in the water.
The head is somewhat depressed and the mouth opening is small. The eyes
are also reduced in size. The snout is truncated in dorsal view and the
rhinarium is prominent. The nostrils are situated laterally and open posteri-
orly behind flaplike valves. The numerous vibrissae are situated dorsally
and laterally on the snout and cheeks posterior to the rhinarium and around
the mouth. They reach maximum lengths of 32 to 34 mm, although most
are 14 to 26 mm long. The ear pinnae are reduced essentially to small,
pointed posterior lobes, about 4 mm in length, which do not extend above
the surface of the fur.
The pelage is generally dense and soft, the type characteristically asso-
ciated with the smaller aquatic mammals. In the mid-dorsal region the
underfur is dense with fine hairs which range from 5 to 7 mm in length.
Abundant but less dense and grosser hairs, about 12 to 14 mm long, form
an outer layer. The ventral pelage is similar to that of the dorsum, but the
underfur is shorter.
The fore limbs are relatively short and do not project much from the
body, even when walking. The proximal portions of the hind limbs blend
similarly into the body contours, leaving the remarkable hind feet as the only
limb segments which project obviously from the furred outline of the body.
The tail also is flared into the body outline over the proximal 15 mm of its
length. The front feet are much smaller than the hind feet with long, curved,
sharply pointed claws. On the hind feet, digit I is the shortest, then in order
of increasing length, digits V, II, III, and IV. The greatest differences in
length are between digits I and II and, to a lesser degree, between digits
V and IV. The first and, to a lesser extent, the fifth digits appear to have the
greatest amount of independent movement.
The hind feet have laterally compressed digits interconnected by webbing.
The sides of the feet and toes are lined with single rows of closely spaced stiff
hairs which reach maximum lengths of three and four mm on the medial and
lateral sides of the feet, respectively, and two to three mm on the toes and
webbing, with the shortest hairs being those near the middle of each web. The
claws are deep and essentially as broad as the tip of the terminal phalanx, that
on digit I being the deepest and the claw on digit II the broadest. The tips are
pointed in lateral profile, but somewhat rounded and scooplike in ventral
aspect. The massive appearance of the hind claws is very striking when they
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Adaptations of a Water Mouse
5
are compared with those of other rodents of similar size such as Neotoma,
Nelsonia, Xenomys, Tylomys, Nyctomys, and Rattus.
When the digits are abducted, the webbing reaches the level of the
pennultimate interphalangeal joint between digits II, III, IV, and V, but is less
extensive between digits I and II. Each web originates at the base of the claw
and extends, as a widening flap, to the levels just mentioned. There the margin
turns and a similar but narrower flap continues to about the same claw level on
the next toe. On the lateral (post-axial) side of each digit, the webbing arises
along the dorsolateral surface, whereas on the medial side of the succeeding
digit it connects along the medial surface. When the foot is extended, the digits
rotate so that their ventral surfaces turn mediad. The metatarsophalangeal
joints are somewhat loose, apparently allowing the digits to rotate even more
than do the metatarsals. The metatarsals (except III and IV) and free digits
abduct and, since they are rotated, the latter also extend at the metatarso-
phalangeal joints in a more or less horizontal plane. These actions spread the
foot and toes while turning the digits so that they present their broad com-
pressed surfaces and spread the webbing to its greatest extent. Part of the
rotation of the digits may be passively effected by the tension of the webbing.
When fully expanded, the hind foot of our specimen measured 14 mm
across the plantar surface at the level of the distal ends of the metatarsals, and
the expanded hair fringes added another six mm at that level, resulting in
greatest dimensions of 20 x 39 mm. Planimetric measurements of the best of
a number of inkpad prints of the right hind foot, including hair fringes, showed
areas of 3.8 to 4.0 cm^. The two hind feet, then, form impressive propulsive
surfaces during extension in the backward stroke while swimming. In the re-
covery stroke, it is probable that the foot is flexed and the digits flexed, ab-
ducted, and possibly rotated somewhat laterad again so that the compressed
toes are curled slightly and lie close together in a staggered sequence (cross
sectional), with digit I leading, and the hair fringes flattened against the sides
of the foot and toes. Thus the pes becomes narrowed with the digit alignment
presenting a laterally compressed, rather than an anteroposteriorly flattened
unit, approaching the characteristics of a hydrofoil, and offering minimum
resistance to the water.
The free tail, from the point at which it leaves the furred base which
flares into the body, measures 143 mm to the tip. The proximal 30 mm is
circular in cross section and approximately 7 mm in diameter. At the end of
this basal portion, there is a constriction to about 6.5 mm diameter and the tail
gradually tapers and becomes compressed. In the last 15 mm the tail narrows
more rapidly to almost a point in dorsal view and, in lateral aspect, reaches its
shallowest dimension (a little less than 5 mm), then expands slightly ventrally
and ends in a somewhat rounded tip which gives the appearance of turning a
little ventrad. Along the ventral surface of the free tail, particularly from the
level of the basal constriction where the cross section of the tail begins to
change shape, the hairs arising from the ventral surface of the tail, represent-
ing approximately the tail circumference arc, become abruptly longer (to
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No. 182
about 5 mm) and more dense, forming a kind of broad fringe extending
along the undersurface of the tail to the point where it expands ventrally near
the tip. The hairs on the rest of the free tail are shorter and relatively sparsely
but evenly distributed. Internally the most notable feature of the tail structure
is the presence of numerous heavy tendons attaching firmly to the vertebrae,
fascia and skin, distal to the basal constriction. The extraction of the tail beyond
this point during skinning was difficult. (James H. Brown, personal communi-
cation, recalled the same difficulty with the specimens of R. underwoodi he
prepared from the same locality.) This clearly suggests well developed mus-
cular control of the tail, further evidence of which was obtained by selective
pulling of the tendons, while the tail was partially extracted, which caused
bending in various directions and at different points in the more distal portions
of the tail.
The streamlining provided by reduction and blending of projecting parts
of the body and the generally fusiform shape and flared tail base make
Rheomys a hydrodynamically efficient body. The large hind feet provide the
main propulsive force and in recovery strokes tend to give minimum resistance
by the combined characters of compressed digits and the foot action mech-
anism. The tail apparently serves a stabilizing and a supplementary propulsive
function, being adapted for both by compression, by effective control through
the tendon arrangement, and probably also by the paddle-shaped tip. Dorsal-
ventral bending of the tail was noted above, during surface swimming. The
slightly ventrally oriented distal portion of the tail (the paddle) probably is
significant when the tail is thus curved in such a way that the tip approaches,
or is at, the water surface while the rest of the tail is below. The tip then would
be more nearly horizontally oriented. This curving might also make the ventral
hair fringe stand out more and so contribute to the compressed tail shape. This
fringe may serve other functions since the hairs are not particularly stiff.
Possibly it protects the ventral surface of the tail from abrasion by streamside
rocks and gravel, or it may serve as a drip line for rapid water runoff.
The thoracic vertebrae of the skinned carcass were capable of rather
acute kyphotic curvature. The panniculus carnosus is a rather broad, thin
sheet posteriorly (lumbosacral region), but gives rise to a number of long,
narrow extensions running up over the back to the head and shoulder region.
Contraction of this muscle pulls the skin back and forth over the thoracic
region. This action separates the hairs of the back when the thoracic region is
curved during grooming.
Discussion
In attempting to understand aquatic adaptation of Rheomys underwoodi,
we surveyed certain literature on semi-aquatic mammals. (Anthony, 1921,
1923, 1929; Bauchot and Stephan, 1968; Conaway, 1960; Dickey, 1928;
Enders, 1938; Goldman, 1912, 1920; Goodwin, 1959; Guth et al., 1959;
Handley and Mondolfi, 1963; Hershkovitz, 1944, 1955; Hooper, 1947, 1968;
Howell, 1930; Lonnberg, 1921; Malzy, 1965; Peyre, 1956; Stirton, 1944;
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Adaptations of a Water Mouse
7
Svihla, 1934; Thomas, 1893, 1897, 1906a, 1906b, 1924a, 1924b; De Winton,
1896). Most of our comparative information was taken from Walker ( 1968).
Although the survey was not exhaustive, we found that relatively little is
known about semi-aquatic small mammals, especially the more highly special-
ized forms. However, limited discussion of the position which R. iinderwoodi
occupies in the evolution of specializations in these mammals is possible.
Rheomys iinderwoodi is among the most highly hydrodynamically spe-
cialized mammals, particularly when compared with species of comparable
size and habits, the smaller semi-aquatic rodents and insectivores, and the
marsupial Chironectes. On the basis of what we could determine, it is quite
likely that most aquatic features of R. iinderwoodi are not unique, but a
comparable total combination of specific features is apparently limited to
members of only eight other genera: three of rodents (in addition to certain
other Rheomys species), Anotomys, Ichthyomys (both closely related to
Rheomys) , and the hydromyine Crossomys; five of insectivores, Limnogale,
and Micropotamogale (M. ruwenzori) (Tenrecidae) , Desmana and Galemys
(Talpidae) and Nectogale (Soricidae). As might be expected, the details of
specialization are most similar among the rodents. Limnogale and Nectogale
seem to be the most closely comparable of the insectivores.
Davis (1964:323) considered two factors to be “primarily responsible
for adaptive modifications in the morphology of mammals: locomotion, and
foods and feeding.” In the case of forms which are more highly adapted for
aquatic living, the locomotor requirements strongly influence the morphologi-
cal modification related to obtaining food. In those particularly adapted for
rapid swimming, morphological feeding specializations not in accord with
the hydrodynamic requirements imposed by locomotor specializations are not
to be found. Although this applies to semi-aquatic, or amphibious mammals,
as well as to the more completely aquatic, the degree of overall modification in
the former is limited by the requirements of terrestrial locomotion, and a
structural-functional balance is struck according to the relative significances
of aquatic and terrestrial locomotion in the lives of these animals. Further-
more, not all aquatic adaptations contribute directly to aquatic locomotion,
and the ability to swim does not necessarily involve much in the way of mor-
phological modification (Howell, 1930). Rheomys iinderwoodi must be con-
sidered a semi-aquatic rodent with a high degree of morphological specializa-
tion reflecting primarily selection for effective, rapid aquatic locomotion as
the major influence.
Davis (1964:323) further stated that “In mammals the major forces
impinging on locomotion are escape from enemies, pursuit of prey and wan-
dering in search of food or water.” These categories can be modified to include
as potentially significant functions (forces or factors) : 1) simple translocation
or unforced movement (wandering for whatever reason); 2) escape from
predators; 3) escape in the event of changing environmental conditions;
4) locating and feeding on plants (or mollusks; i.e. stationary food); 5) pur-
suing and capturing other animals (mobile food). All of these can play a part
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No. 182
in determining the degree of adaptation for aquatic locomotion in the evolu-
tion of semi-aquatic forms. One other important factor remains which has a
basic influence on the selective requirements related to the five functions just
listed: namely, the nature of the aquatic environment. Ability to swim in still
water requires less in the way of morphological specialization than does the
ability to swim in a strong current (function 1 or 4 above), and if forced
locomotion (functions 2 and 5) is required, then increased specialization
becomes a necessity under both types of aquatic conditions. Escape by swim-
ming from predators that do not swim puts more stress on the ability to get
into the water rapidly than it does on swimming. The high degree of hydro-
dynamic specialization in Rheomys underwoodi appears to have evolved as an
adaptation of this semi-aquatic rodent for unforced movement, escape from
predators and, probably most significantly, for obtaining moving food in the
sometimes torrential streams along which it lives. The adaptive features re-
lated to these factors doubtless have survival value, as well, when the streams
rise rapidly and become swifter and more turbulent during heavy rains
(function 3).
The types of food (discussed in more detail by Bauchot and Stephan,
1968) and the type of aquatic environment appear to be the most important
factors in the evolution of the semi-aquatic radiation among small mammals.
Most semi-aquatic herbivorous mammals occur in level or lowland and coastal
regions in which graminoid and low herbaceous vegetation forms a major
source of food and where marshes, ponds and river overflows are to be found
but rapid streams are unusual. In such situations, water edge species probably
took to water, mainly, in order to take advantage of the additional (similar)
plant food thus made available and also to survive during periods of flooding.
In this category apparently fall most of the semi-aquatic rodents, of which
many are little specialized aquatically; others show varying degrees of spe-
cialization, but all are still limited by the importance of terrestrial locomotion
as a means of dispersal and of relocation during times of change in their
aquatic habitats.
Among those showing little or no specific adaptations for aquatic loco-
motion, but which more or less routinely enter water and swim, can be men-
tioned species of Thryonomys (Thryonomyidae) , Bandicota, Dasymys, Hy-
bomys (Muridae), Microtus, Oryzomys and Reithrodontomys (Cricetidae) ,
to name only a few in just these three families (see Walker, 1968; Fisler, 1961,
1965). Species of other genera have evolved definite aquatic locomotor spe-
cializations and apparently utilize more of the food occurring in the water: the
oryzomine Nectomys, the sigmodont Holochilus, and the microtine Arvicola
are obvious examples. Those genera (such as Hydrochoerus, Myocastor,
Ondatra, and Neofiber), which have become more dependent on semi-aquatic
or aquatic plant food, have evolved more aquatic locomotor specializations
and are capable of swimming in relatively slow and steady currents present in
small streams and rivers. It seems significant that these last mentioned genera
are among the larger rodents, suggesting that there is a relationship between
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Adaptations of a Water Mouse
9
the size of the animal and the ability to enter larger bodies of water and
stronger currents where more sustained swimming is required. As must be the
case with all aquatic and semi-aquatic mammals, their respective degrees of
specialization have evolved in relation to the most severe aquatic conditions
(currents, distances to be covered in water) in the bodies of water which they
frequent or inhabit. Many of the rodents mentioned may be found in a variety
of aquatic situations, but few, if any, characteristically occur in water with
rapid currents. None feeds to any extent on active aquatic food although some
do utilize inactive aquatic animals at least occasionally. The water opossum,
Chironectes, seems to be comparable with this group of rodents in terms of
degree of aquatic specialization, but it also fits in with the next group of small
mammals in that it both feeds on aquatic animals and probably evolved in a
forest environment.
Apparently the semi-aquatic insectivores and the more highly specialized
semi-aquatic rodents have evolved, for the most part, in hilly or mountainous
regions. Here forest is more prevalent than open grassland or other vegeta-
tional types which provide much in the way of leafy food, and moving water
characterizes the aquatic habitats (streams and small rivers) . Some of these
conditions, and/or other coincident factors, have apparently been favorable
to insectivore radiations, and the presence of good sources of food along the
water edge and in the water itself has provided the setting for the evolution of
the semi-aquatic members of the order. The more highly specialized insecti-
vores already mentioned are characteristically found associated with streams
or small rivers, usually in forested areas. The other known semi-aquatic in-
sectivores show somewhat wider and more variable ecological characteristics
but probably have evolved under similar conditions. Potamogale, one of the
largest insectivores, is unique among semi-aquatic members of the order in
having a highly specialized tail as its sole locomotor organ, a feature which
probably makes it one of the most efficient swimmers of the group (more
nearly comparable to the otters) . This African tenrecid is found in all types
of water, from lakes to torrential mountain streams. The remaining semi-
aquatic members of this order are not highly specialized for aquatic locomo-
tion showing only relatively slight modifications of the terrestrial locomotor
apparatus. They do not possess swimming abilities comparable to those of the
insectivores already mentioned. The relatively large hind feet, long tail and
more streamlined body of the star-nosed mole, Condylura, (as compared to
typical moles) are of positive value in aquatic locomotion, if not direct spe-
cializations for this function, as well may be the less fossorially modified fore
limb of this animal. The water shrews belonging to the soricid genera Neomys
and Chimarrogale and species Sorex palustris and S. bendihi probably can be
considered ecological equivalents (roughly) in their respective separate ranges
of distribution, having approximately the same morphological adaptations
(fringed feet and tail) . They appear to be most similar in aquatic adaptations
to the less specialized species of the Neotropical rodent genera Holochilus and
Nectomys. Finally, Micropotamogale lamottei, characteristically found in
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No. 182
slower waters, seems to be morphologically adapted for aquatic locomotion
to about the same extent as the water shrews although it is larger and shows
more aquatic specializations.
The insectivores mentioned previously as being adaptively comparable
to Rheomys underwoodi are more highly specialized for aquatic locomotion
and are, in fact, more aquatic than those just discussed, with the exception of
Potamogale, and they characteristically live in rivers and rapid to torrential
streams. Their adaptations have evolved in response to the requirements im-
posed by more or less active pursuit of food in the currents and turbulence of
the aquatic environment.
In comparison with this last group of insectivores, rodents living in regions
characterized by forested streams and rivers have less potential food available
in and around the water if they depend on plant sources. Those entering the
streams have had to assume a diet of moving animal food (most likely before
taking to the water), the capturing of which, under the aquatic conditions
presented by the active waters, has been a significant factor in the evolution
of their locomotor specializations. Two groups of rodents have shown some-
what similar radiations in such situations: the Neotropical ichthyomyine
cricetids and the New Guinea- Australian hydromyine murids.
The ten genera of hydromyines (with some 16 species) are diversified
and include small mouse and shrew-like forms (Microhydromys, Parahydro-
mys) and other apparently terrestrial herbivores, as well as three genera which
have taken to the water and include animals in their diets. These three,
Xeromys, Hydromys and Crossomys, have followed different lines of speciali-
zation representing the extremes of the spectrum of aquatic modifications
found in other groups of rodents (Lidicker, 1968; Mahoney, 1968; Tate,
1951). Xeromys, the only endemic Australian genus of hydromyines, feeds on
aquatic vegetation and inactive aquatic mollusks in swampy situations and has
little, if any, apparent morphological adaptations for aquatic locomotion. The
more widely distributed genus, Hydromys, represents a more muskrat-like
type of semi-aquatic specialization, with the widespread H. chrysogaster being
associated with a wide range of aquatic environments and feeding on a variety
of animal and plant foods. Finally, the monotypic New Guinean genus
Crossomys, found along mountain streams and very likely feeding on aquatic
animals, appears to be most similar to Rheomys underwoodi in ecological
characteristics and in both degree and detail of morphological adaptation for
aquatic locomotion.
The five genera of ichthyomyines are less diversified than the hydro-
myines, rather they show differing degrees of specialization for life along
mountain streams. Hershkovitz (1962) proposes that the ichthyomyines are
derived from pastoral forms, primarily on the basis of dentition; we believe
that, in any case, the ancestral forms most likely had to enter the forest to be
exposed to the stream habitat, since gallery forest, at least, is almost always
associated with permanent streams in the New World Tropics. The monotypic
genera Neusticomys and Daptomys exhibit only slight morphological modifi-
1970
Adaptations of a Water Mouse
11
cation for aquatic habits, whereas the various species (16 named forms; eight
to 14 species, depending on the authority consulted) of Rheomys and Ichthy-
omys carry aquatic locomotor modifications to varying extents of specialization
(Cabrera, 1961; Goodwin, 1959; Handley and Mondolfi, 1963; Hooper,
1968). Anotomys (monotypic), Rheomys imderwoodi, probably R. mexicanus
(according to the informative description provided by Goodwin, 1959) and
several other species of Rheomys (Hooper, 1968) and of Ichthyomys (An-
thony, 1929) appear to have attained more or less similar high levels of aquatic
locomotor adaptation, along several parallel evolutionary pathways. As already
pointed out by Hooper (1968), the other ichthyomyine species found in Costa
Rica, Rheomys hartmanni, shows less specialization for aquatic locomotion
than does R. imderwoodi. The LACM specimen of R. hartmanni (LACM
25418) from north of San Isidro del General, in the Cordillera de Talamanca
(Hooper’s 5200-ft. station, Rio Buena Vista tributaries, Prov. San Jose:
Hooper, 1968) shows the lesser degree of specialization of the fur, ears, hind
feet and tail indicated by Hooper (1968) and, furthermore, shows much less
(negligible) webbing of the hind feet, and a tail which appears to be round
and to lack the ventral hair fringe. This specimen also lacked the specialized
tail muscle-tendon complex described above in R. imderwoodi.
Differences such as those between Rheomys imderwoodi and R. hart-
manni, which have been outlined, are not always readily explainable. A more
thorough interpretation of the evolution of the adaptations of small mammals
to the semi-aquatic habit awaits a detailed knowledge of these adaptations.
Most references we have seen have been taxonomic in nature and have dis-
cussed mainly diagnostic or comparative features of the mammals described,
presenting little useful information on other aspects of their morphology. More
ecological data, such as kinds of food consumed and detailed characteristics
of habitats where the animals are found, along with information on behavioral
adaptations, will provide bases for interpreting the significance of morphologi-
cal specializations.
Acknowledgments
The LACM specimens and original field data reported in this paper were
collected in 1963 and 1966 during work carried on in Costa Rica as part of
the Los Angeles County Museum of Natural History mammalian ectoparasite
project (LACM-USACR; see Starrett and Casebeer, 1968) under U.S. Army
Medical Research and Development Command grants No. DA-MD-49-183-
62-G54 and DA-MD-49-193-63-G94 (F. S. Truxal and C. A. McLaughlin,
Jr., principal investigators). It should also be mentioned that both LACM
specimens of Rheomys were previously recorded in relation to this project by
Geest and Loomis (1968). In addition, in 1966, facilities and various services
were provided by The Organization for Tropical Studies (O.T.S.) and the
Universidad de Costa Rica, San Jose. The authors wish to thank, in particular,
Norman J. Scott, Jr., O.T.S. , for his help. We also here express our apprecia-
tion to Emmet T. Hooper and James H. Brown for their invaluable courtesy in
loaning traps to us and providing information and advice on several occasions.
12
Contributions in Science
No. 182
Resumen
Un macho adulto de Rheomys underwoodi fue capturado vivo, en Abril,
1966, en la orilla del Rio Poasito, a 2000 m en el Volcan Poas, Provincia de
Alajuela, Costa Rica. En captividad, este animal comio sardinas de lata
vorazmente, pero reuso varios tipos de plantas de la region de captura que
le fueron ofrecidas. En general, sus acciones al moverse en la trampa, como
limpiandose y comiendo, fueron rapidas y parecidas a las acciones de musa-
ranas. Basado en observaciones limitadas de este roedor semi-acuatico en el
agua de un recipiente restringido, su comportamiento mostro tambien acciones
rapidas al entrar y salir del agua, nadando en la superficie o zambullendose.
La propulsion era efectuada por movimientos alternados de las anchas patas
traseras.
Entre las especializaciones morfologicas adaptadas a la locomocion acu-
atica, que este raton presenta, las siguientes ban sido notadas: un cuerpo
hidrodinamico del cual solo las patas traseras y la cola protruden durante la
natacion, la cola ligeramente compresa longitudinalmente con arreglo
complejo de tendones para su control; pies traseros grandes con tela inter-
digital y crestas de pelos que dan una superficie total de cada pie de aproxi-
madamente 3.9 cm-. Ademas, las patas traseras pueden voltear y doblarse de
tal manera que presentan una estructura de “hydrofoil,” con una resistencia
minima contra el agua durante el movimiento hacia adelante (de recobro) al
nadar.
Finalmente, en una forma general y especulativa, una discusion de la
evolucion de habitos semi-acuaticos en pequenos mamiferos es presentada.
Dos cursos principales se pueden proponer: 1) en areas de tierras bajas, donde
el habitat acuatico consiste principalmente de aguas quietas o lentamente
corredizas y la vegetacion acuatica es abundante, los pequenos mamiferos
semi-acuaticos son en su mayoria roedores que comen, principalmente, plantas,
y que tienen tendencia a producir especializaciones morfologicas limitadas
para locomocion acuatica; 2) en areas de montana bien forestadas, donde los
arroyos y riachuelos tienen aguas rapidas que contienen mas animates ac-
uaticos que plantas acuaticas, los habitos semi-acuaticos se han desarrollado
en varios insectivoros y algunos grupos de roedores que han podido adaptarse
a una dieta de pequenos animates. Rheomys underwoodi, asi como otros
roedores ichthyomyines Neotropicales y los roedores hydromyines, Crossomys,
de Nueva Guinea, pertenecen al ultimo grupo y han desarrollado un alto grado
de especializacion morfologica a la locomocion en aguas rapidas, a veces tor-
rentosas, en que ellos viven.
1970
Adaptations of a Water Mouse
13
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Accepted for publication December 1, 1969
contributions
= IN SCIENCE
Number 183
April 17, 1970
OBSERVATIONS AND RECORDS OF
MYOTIS (PIZONYX) VIVESI MENEGAUX
(Chiroptera: Vespertilionidae)
By Donald R. Patten and Lloyd T. Findley
Los Angeles County Museum of Natural History
Los Angeles, California 90007
Exposition Park
CONTRIBUTIONS IN SCIENCE is a series of miscellaneous technical papers
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should be directed to Virginia D. Miller, Los Angeles County Museum of Natural
History, 900 Exposition Boulevard, Los Angeles, California 90007.
Robert J. Lavenberg
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Editor
OBSERVATIONS AND RECORDS OF
MYOTIS (PIZONYX) VIVESI MENEGAUX
(Chiroptera: Vespertilionidae)
By Donald R. Patten' and Lloyd T. Findley-
Abstract: A large number of Myotis (Pizonyx) vivesi
Menegaux were observed at a minimum distance of four nautical
miles from land from the research vessel, Velero IV, trawling in
800 fathoms of water in the Canal de Salsipuedes, Gulf of Califor-
nia, at a speed of four knots. Bats accompanied the vessel for a
minimum of three hours flying into a headwind of 16 knots. Flight
patterns in relation to the vessel and apparent feeding behavior
are described. Contents of the intestinal tracts of three specimens
included finely masticated muscle tissue, fish scales, at least one
piece of a caridean shrimp and a structure thought to be the ab-
domen of an unidentified polychaete larva. New records for
Myotis vivesi are Isla de Alcatraz, Sonora, and Bahia Rosario,
Baja California Sur, the latter being the southernmost distribu-
tional record.
Detection of localized surface disturbances of the water
are discussed as a possible means of food detection. The corres-
pondence between the distribution of M. vivesi and areas of large
scale upwellings and subsequent phytoplankton blooms in the
Gulf of California and adjacent Pacific Ocean is noted.
Three roundworms, genus Trichuris, the mites Spintiirnix
mexicanus Rudnick and Steatonyssus leptus Radovsky and ticks
of the genus Ornithodoros were found as parasites on M. vivesi.
All known ectoparasite records from M. vivesi are presented.
Similarity in morphological and behavioral characteristics
(including piscivorous habits) between M. (Pizonyx) vivesi and
other species of Myotis is noted. The taxonomy of Pizonyx Miller
is discussed, and the taxon is considered a subgenus of Myotis
Kaup.
In the course of a midwater trawl in the Gulf of California on the evening
of November 30, 1967 the research vessel, Velero IV, was nearly engulfed by
a large aggregation of bats. The circumstances and observations of this incident
merit reporting.
Observations and distributional records: At 11:30 pm three non-gravid,
non-lactating females were caught in large dip nets as they flew past the fantail
and were subsequently prepared and identified as Myotis (Pizonyx) vivesi
Menegaux (LACM 28273-28275).
When the bats were caught, the vessel was trawling in a northwesterly
^Curator of Mammalogy, Los Angeles County Museum of Natural History.
^Assistant Curator of Fishes, Department of Biological Sciences, University of
Arizona, Tucson, Arizona 85721. Formerly, Section of Ichthyology, Los Angeles
County Museum of Natural History.
1
9
Contributions in Science
No. 183
direction in 800 fathoms of water in the Canal de Salsipuedes (sometimes
written Sal si Puedes) at a speed of four knots. The exact location was 4.2
nautical mi. W (263° Radar Target) of Isla Salsipuedes, 28° 43' 58" N,
113° 04' 00" W. Barometric pressure was 29.97 in. Hg, sea temperature was
21.1° C (70° F), and the wet and dry bulb air temperatures were 16.1° C
(61° F) and 19.7° C (67.5° F), respectively.
At approximately 1 1 pm a single bat was noticed when it flew up the port
passageway outside the gallery. This was the first time that any of the scientists
were cognizant of the situation. Personnel on the bridge, however, reported
that the bats first appeared at 10 pm and were continuously present until about
2 AM of December 1 when they all left abruptly. A minimum of 17 nautical
miles was traversed during this four hour time interval. At 10 pm the vessel
was a minimum of four nautical miles from the closest land mass, Isla las
Animas, while at 2 am Isla Partida was 4.7 nautical miles to the east, and
Punta de las Animas (mainland Baja California Norte) was 2.5 nautical miles
to the west. The vessel was never closer than four nautical miles to any land
during the first three hours.
Due to its specialized habitat, it is unlikely that M. vivesi is ubiquitously
distributed throughout the islands of the Gulf of California and adjacent land
areas. Maya (1968) found Pizonyx present in January, spring and summer on
the following four islands in the vicinity of the Canal de Salsipuedes: Partida
(= Isla Cordonosa, the type locality) , Raza, Salsipuedes and San Lorenzo.
For a summary of distributional records see Maya (1968), Reeder and Norris
( 1954) , and Banks (1964) . In addition to the Canal de Salsipuedes record,
there are specimens in the Los Angeles County Museum of Natural History
(LACM) mammal collection from two unpublished localities as follows: Isla
de Alcatraz, Bahia Kino, Sonora, Mexico (33 specimens, LACM 13109-
13140, collected by A. L. Gardner, June 25 through July 9, 1960) and Bahia
Rosario, Baja California Sur, Mexico (9 specimens, LACM 28276-28286,
collected by K. E. Stager, March 23, 1963) . The latter specimens were taken
from under rocks on a small island at the south end of Bahia Rosario (24° 15'
N, 1 10° 09' W), ca. 1 mi. SE Punta Coyote, and constitute the southernmost
record for this species.
Reeder and Norris (1954) reported a probable sight record of this
species at sea two miles off Punta Malarrimo, Bahia de Sebastian Vizcaino,
Pacific side of Baja California. Our specimens from Salsipuedes appear to be
the first conclusive evidence of M. vivesi occurring at considerable distances
from land.
According to the ship’s log the wind was from the southeast at 6 knots at
9 PM, “calm” at 10 pm, and thereafter from the northwest at an average
velocity of 16 knots. Several observers noted that the bats first appeared during
the period of calm and for the remaining four hours flew alongside the vessel
into a 16 knot headwind.
1970
Observations of MYOTIS (PIZONYX) VIVESI
3
Despite the absence of moonlight, observation was made possible by the
vessel’s running lights, a portable spotlight, and special lights on the fantail
and A-frame which supported the trawl. Extent of visibility is estimated to
have been 13.7 m (45 ft.) from the vessel. No phosphorescence or biolumi-
nescence was observed in the water.
Approximately two hundred individuals were estimated to be visible
around the fantail at any one moment. The total number is conservatively
estimated to have been at least twice that number. It is not known if the same
individual bats were present during the entire four hours or even if those
visible at any one moment were just part of a much larger, and unobservable
number.
Bats were observed flying all around the vessel except directly in front
of and under the bow, but they appeared to be concentrated around the stern,
presumably because of the lights and wake of the vessel. Most individuals re-
mained more than 2.4 m (8 ft.) from the vessel, but some bats flew diagonally
across the stern of the vessel and maneuvered to avoid the superstructure of
the vessel and the attempts of observers to catch them in dip nets. Those
alongside the vessel appeared to fly repeatedly in an oval pattern, the long axis
parallel to the ship. From the vicinity of the bow individuals would fly adjacent
to the sides of the ship until reaching the stern where they would disperse over
the wake of the ship, reorient into the wind, drop down to the water surface,
and fly forward toward the bow. In the forward flight bats were observed just
above the sea surface until reaching the vicinity of the bow where they abruptly
rose in altitude. Flight speed in the sweep tow'ard the stern was discernibly
faster than that toward the bow. The bats appeared to be definitely interested
in the area of crests produced by the wake of the vessel as well as the choppy
sea surfaces lateral to the vessel.
Bats at the stern were estimated to be flying 2 in. to 1 ft. above the surface
and were definitely observed to dip downward touching the water. Several
observers reported that they actually saw the bats touching the water surface
but could not distinguish which body part made contact. Robert J. Lavenberg
states, that while operating the spotlight, he observed individual bats swoop
down until just above the water surface and then alternately lower and raise
their feet, making contact with the water.
Lavenberg’s observation is pertinent in light of the controversy over the
manner in which food is procured. Miller and Allen (1928) and Burt (1932)
reasoned that the elongated, laterally compressed and recurved feet func-
tioned to capture small fish or cmstacea. On the basis of several observations,
Reeder and Norris (1954) stressed that the tail, uropatagium and calcar
cartilages appeared to be the primary mechanism. Walker (1950) observed a
V-shaped ripple mark remaining after a low swoop by a bat and believed that
this was made by the animal’s chin. To our knowledge no one has recorded
observing M. vivesi actually obtaining prey from the sea, although Maya
4
Contributions in Science
No. 183
(1968, p. 65) observed several individuals . . with fish in their claws.” In
addition to the feet, the tail and uropatagium were considered by Maya (1968)
to be used in capturing prey because these were often wet and frequently bore
fish scales.
Food: After examination of both guano and stomach contents, Burt
(1932, p. 364) found definite evidence of “. . . fish remains, chiefly scales with
an occasional small fish bone.” Burt further reported (p. 363) that one of the
bat stomachs was found to contain fish “. . . belonging to the sardine tribe.”
On the basis of circumstantial evidence. Walker (1950) believed the diet to
consist of small shrimp. He further theorized that upon approaching the water
surface the white venter (atypical for the genus Myotis) served to agitate and
frighten schools of crustaceans, thus facilitating their capture. Reeder and
Norris (1954) recorded in stomach contents small crustaceans and, in one
specimen, many insect fragments. Norris (Reeder and Norris 1954, p. 85)
observed one bat in flight “. . . holding a shining little fish in its hind feet.” Of
an undisclosed number of fecal pellets obtained from roosting sites. Carpenter
(1968, p. 956) found “. . . 62.5 per cent contained remains of only Crustacea,
26.3 per cent contained only fish remains and 11 per cent contained remains
of both.” Maya (1968) found the presence of crustaceans, fish, insects, and
algae in the stomachs of 22 specimens. In droppings he found some indication
of seasonal variations in the relative frequences of the above food items, but
concluded that crustaceans were the major food source. Examination of the
digestive tracts of our three specimens disclosed finely masticated pieces of
muscle tissue, fish scales, at least one piece of a shrimp (Decapoda: Caridea)
and a structure thought to be the abdomen of an unidentified polychaete larva.
Analysis of the fish scales revealed that the majority were from the family
Myctophidae (lantern fishes), comparing well in size, shape and other scale
characters to those of the genera Hygophum, Diaphus and Triphoturus. Thir-
teen of the 65 scales showed a definite, raised center portion which appeared
to be the lens remnant of a myctophid photophore scale whereas 50 additional
scales were non-photophore myctophid scales. Two scales were small and
ctenoid and were not myctophid in origin, those of the latter being larger and
cycloid. Many myctophids are vertical migrators from great depths at night
when they would occur at the surface.
From roosts of M. vivesi Maya (1968) found the dry and partially
chewed remains of the following fishes: File stolifera, Ophisthonema liboleta
[sic, Opisthonema libertate], bothids, and perhaps flying fish, with bothid
remains being the most common. Adult flatfishes of the family Bothidae are
benthic whereas the larvae and prejuveniles live in plankton. Scale develop-
ment is probably slight, if at all, during the planktonic stages. The degree of
scale development of the bothid remains mentioned by Maya (1968, fig. 15)
would indicate that they were probably recently developed juveniles which
1970
Observations of MY OTIS {PIZONYX) V IV ESI
5
inhabit shallow waters near shore. LUe stolifera and Opisthouema lihertate
are herrings (Clupeidae) and are surface schooling fishes.
When removed from the dip net, one of the three bats was discovered to
have a small unidentified shrimp adhering to the surface of the uropatagium.
During the frantic efforts to capture the bats as they flew close to the sides of
the vessel and across the fantail, one of the specimens was caught in a dip net
which subsequently touched the water before being raised to the deck. It is not
known whether or not the specimen bearing the crustacean was the one in-
volved nor is it known if the shrimp was present when the bat was initially
caught or if it was originally adhering to the dip net and subsequently trans-
ferred to the bat.
Food detection: The bats emitted audible sounds described as short, high
pitched squeals, which were readily discernible. The overall intensity of sound
varied in direct proportion to the height of flight, being loudest when the bats
were closest to the water. No noticeable change in pitch was reported by the
observers whereas Maya (1968, p. 66) states that “As they approach the
surface, their vocal chatter, a series of short bursts, becomes high pitched,
more rapid, and finally inaudible.”
Slithers (1965) found that in captivity Noctilio leporinus Linnaeus, a
neotropical bat known to be piscivorous, could readily detect small wires
projecting vertically above the water as well as localized minor disturbances
in the water surface. M. vivesi probably detects prey in the same manner as
N. leporinus although in a subsequent study Slithers (1967) reported a differ-
ence in the orientation sounds produced by these two species. He was unable
to induce captive M. vivesi to take pieces of shrimp from the surface of an
experimental pool.
The behavior and apparent feeding forays by M. vivesi in the areas ad-
jacent to the stern and sides of the vessel may have resulted from disturbances
of the water by the wind and the passage of the vessel. The choppy sea pro-
duced by the wind was similar to the wake of the vessel. Once agitated, any
small fish or crustacean could well have produced small surface disturbances,
thus facilitating their capture by the bats.
Maya (1968, p. 67) mentions that M. vivesi cannot select only small fish,
pointing out that “. . . many [of the fish] caught are frequently too large for
effective feeding.” On another occasion he witnessed a bat “. . . hovering above
and actively pursuing an object” (1968, p. 66). Maya recovered the object
which was an unidentified shrimp (1968, pp. 61 and 66, fig. 16). Judging from
the scale in the figure the shrimp appears about 6 cm. long and too large for
effective feeding by M. vivesi. This would seem to be a logical consequence if
M. vivesi was reacting to a localized surface disturbance rather than dis-
criminating as to the kind and size of organism producing the disturbance.
The known distribution of M. vivesi conforms in general with areas of
the Gulf of California and adjacent areas of the Pacific Ocean subject to large
6
Contributions in Science
No. 183
scale upwellings and subsequent phytoplankton blooms, e.g. the Canal de
Salsipuedes (Parker, 1964, p. 35). Perhaps M. vivesi actively seeks out those
areas where their prey species, higher trophic level consumers, would be rela-
tively more abundant.
Parasites: Three female roundworms of the genus Trichuris (Dorylai-
mida: Trichinellidae) were found as intestinal parasites in one of our three
specimens. In addition, three types of ectoparasites were found: one mite,
Spinturnix mexicanus Rudnick (Acarina: Spinturnicidae) ; two protonymphs
of the mite, Steatonyssus leptus Radovsky (Acarina: Macronyssidae) ; and
four specimens of a tick of the genus Ornithodoros (Acarina: Argasidae),
which apparently are not O. dyeri Cooley and Kohls (Glen M. Kohls, per-
sonal communication).
The following ectoparasites have been recorded from M. (Pizonyx)
vivesi: Ornithodoros dyeri Cooley and Kohls (1940) (Acarina: Argasidae);
Whartonia sonorensis Hotfmann (1960) (Acarina: Trombiculidae) ; Trom-
bicula thompsoni Brennan (1966) (Acarina: Trombiculidae); Speleocola
cortezi Loomis and Webb (1969) (Acarina: Trombiculidae); and the batfly,
Basilia pizonychus Scott (1939) (Diptera: Nycteribiidae) . The latter has also
been reported by Reeder and Norris (1954) and Maya (1968). Rupert L.
Wenzel (personal communication) reports that there are no Streblidae known
from M. vivesi. Whartonia sonorensis, Speleocola cortezi and Basilia pizony-
chus are known only from M. vivesi. Apart from the one record on M. vivesi,
Trombicula thompsoni is known only from marine birds in the Caribbean. At
one time, Ornithodoros dyeri was known only from guano and rock crevices of
bat caves, mines, and tunnels from several localities in the western United
States and Mexico, but it is now known from both M. vivesi and the sac-
winged bat, Balantiopteryx plicata (Kohls, Sonenshine, and Clifford, 1965).
Because it is incomplete, current parasitological information can furnish little
evidence concerning the taxonomic relationships between M. vivesi and the
other species of Myotis.
Taxonomy: We have followed the suggestions of Ellerman and Morrison-
Scott (1951, p. 138) and subsequent investigators that Pizonyx Miller and
Myotis Kaup be regarded as congeneric. These two have been shown to be
essentially similar in hair structure (Benedict, 1957), baculum (Hamilton,
1949), and karyotype (Baker and Patton, 1967). Characteristics used to
distinguish Pizonyx from Myotis have been summarized by Baker and Patton
(1967) and are primarily associated with a piscivorous habit (Miller, 1906
and 1907).
To support the generic retention of Pizonyx, Maya (1968) stressed pis-
civorous behavioral adaptations and specializations in water metabolism.
However, other species of Myotis are now known, or strongly suspected, to be
piscivorous and exhibit many of the same specialized morphological and be-
havioral characteristics formerly regarded as unique to Pizonyx. These species
1970
Observations of MYOTIS (PIZONYX) VIVESI
1
are as follows: M. {Leuconoe) megalopus (Dobson), Africa, (Aellen and
Brosset, 1968); M. (Rickettia) ricketti Thomas in Asia [discussed as Ves~
pertilio pilosus Peters by Allen, 1936]; M. (Leuconoe) macrotarsus Water-
house, Asia, and M. (Leuconoe) daubentoni Kuhl, Europe (Brosset and
Deboiitteville, 1966).
The subgeneric designations follow Tate (1941) and are distinguished
by enlarged feet, Leuconoe, versus enormous feet, Rickettia, in addition to
other characteristics similar to those of Pizonyx. The big-footed condition
according to Tate (1941), at least in Leuconoe, is due to phalangeal elonga-
tion (as in Pizonyx) and not to metatarsal elongation as stated by Maya
(1968). In addition to the aforesaid species, Myotis dasycneme Boie (Europe
and Asia), Myotis macrodactylus Temminck (Japan), and Myotis stalkeri
Thomas (Kei Islands) also have large feet.
Although a white venter is not common in the genus Myotis, it cannot be
considered unique to Pizonyx. Specimens in the American Museum of Natural
History of M. ricketti, M. daubentoni and M. dasycneme all possess a con-
spicuous and extensive white frosting to the venter.
Braun (1965) and Carpenter (1968) have shown that kidney structure
and water metabolism, respectively, of M. vivesi are highly specialized, but
without equivalent information for the other species of Myotis, it is difficult
to interpret the taxonomic significance, if any, of this information. Surely the
desert species of Myotis, if not the others, are also specialized in renal physi-
ology, morphology and water metabolism. Dalquest (1948) reports that both
Myotis californicus (Audubon and Backman) and Myotis yurnanensis (H.
Allen) apparently drink saltwater.
Whether or not the characteristics common to both M. vivesi and the
several old world species indicate convergence or a common phylogenetic
origin is a question yet to be resolved. Certainly the present taxonomic arrange-
ment appears illogical and inconsistent, and pending a revision of all the taxa
concerned, we have considered Pizonyx a subgenus of Myotis.
Acknowledgments
For identification and information concerning the ectoparasites we are
indebted to Deane P. Furman of the Division of Parasitology, Agricultural
Experiment Station, University of California, Berkeley; Glen M. Kohls, Rocky
Mountain Laboratory, Hamilton, Montana; Richard B. Loomis, Department
of Biology, California State College (Long Beach); Frank J. Rodovsky,
George Williams Hooper Foundation, University of California, San Francisco
Medical Center; and Rupert L. Wenzel, Field Museum of Natural History,
Chicago. The nematodes were identified by Murray D. Dailey, Department
of Biology, California State College (Long Beach). Donald B. Bright, De-
partment of Biology, California State College (Fullerton) kindly identified the
invertebrate material in the stomach contents. Robert J. Lavenberg, Los
8
Contributions in Science
No. 183
Angeles County Museum of Natural History, Kristian Fauchald, Allan Han-
cock Foundation, University of Southern California, and crew members of
the Velero IV, Fred Ziesenhenne, Captain, John Falcone, Second Mate, and
Fred Schmidt, Chief Engineer, contributed observations and technical data.
To Andrew Starrett, Charles L. Hogue, Ronald H. Pine, James S. Findley
and Karl F. Koopman we owe a special debt of thanks for suggestions con-
cerning the manuscript. To all of the above individuals we express our
appreciation.
Literature Cited
Aellen, V., AND A. Brosset. 1968. Chiropteres du sud du Congo (Brazzaville).
Rev. Suisse Zook, 75: 435-458, 1 pi.
Allen, G. M. 1936. The status of Vespertilio pilosus Peters. Jour. Mamm., 17;
168-169.
Baker, R. J., and J. L. Patton. 1967. Karyotypes and karyotypic variation of North
American vespertilionid bats. Jour. Mamm., 48: 270-286.
Banks, R. C. 1964. Range extensions for three bats in Baja California, Mexico. Jour.
Mamm., 45: 489.
Benedict, F. A. 1957. Hair structure as a generic character in bats. Univ. Calif. Pub.
Zook, 59: 285-549.
Braun, E. 1965. The gross and microscopic renal anatomy of the fish-eating bat,
Pizonyx vivesi. M.S. Thesis. Univ. Ariz,, 31 pp.
Brennan, J. M. 1966. New records of chiggers (Acarina: Trombiculidae) from
Baja California and islands of the Gulf of California. Jour. Parasitok, 52:
772-775.
Brosset, A., and C. D. Deboutteville. 1966. Le regime alimentaire du vespertilion
de Daubenton Myotis daubentoni. Mammalia, 30: 247-251.
Burt, W. H. 1932. The fish-eating habits of Pizonyx vivesi (Menegaux). Jour.
Mamm., 13: 363-365.
Carpenter, R. E. 1968. Salt and water metabolism in the marine fish-eating bat,
Pizonyx vivesi. Comp. Biochem. Physiol., 24; 951-964.
Cooley, R. A., and G. M. Kohls. 1940. Two new species of Argasidae (Acarina:
Ixodoidea). U.S. Pub. Health Rpk, 55: 925-933.
Dalquest, W. W. 1948. Mammals of Washington. Univ. Kans. Pubis., Mus. Nat.
Hist., 2: 1-444.
Ellerman, j. R., and T. C. S. Morrison-Scott. 1951. Checklist of Palaearctic and
Indian mammals, 1758 to 1946. 2 ed. Brit. Mus. (Nat. Hist.), London, 810 pp.
Hamilton, W. J., Jr. 1949. The bacula of some North American vespertilionid bats.
Jour. Mamm., 30: 97-102.
Hoffmann, A. 1960. Contribuciones al conocimiento de los Trombiculidos Mexi-
canos (Acarina, Trombiculidae) (9a. parte). Acta Zook Mexicana, 4(4): 1-11.
Kohls, G. M., D. E. Sonenshine, and C. M. Clifford. 1965. The systematics of
the subfamily Ornithodorinae (Acarina: Argasidae). II. Identification of the
larvae of the western hemisphere and descriptions of three new species. Entomok
Soc. Amer., Annals, 58: 331-364.
Loomis, R. B., and J. P. Webb, Jr. 1969. A new species of Speleocola (Acarina:
Trombiculidae), off a bat, Pizonyx vivesi, from Baja California, Mexico. So.
Calif. Acad. Sci., Bulk 68: 36-42.
Maya, J. A. 1968. The natural history of the fish-eating bat, Pizonyx vivesi. Ph.D.
Thesis. Univ. Arizona. 106 pp. Univ. Microfilms. Ann Arbor, Mich. (Diss.
Abstr. 69-2016).
1970 Observations of MYOTIS (PIZONYX) VIVESI 9
Miller, G. S., Jr. 1906. Twelve new genera of bats. Biol. Soc. Wash., Proc., 19:
83-86.
. 1907. The families and genera of bats, U.S. Natl. Miis., Bull., 57 : 1-282.
Miller, G. S., Jr., and G. M. Allen. 1928. The American bats of the genera
Myotis and Pizonyx. U.S. Natl. Mus., Bull., 144: 1-214.
Parker, R. H. 1964. Zoogeography and ecology of some macroinvertebrates, par-
ticularly mollusks, in the Gulf of California and the continental slope off
Mexico. Dansk Naturh. Foren., Vidensk. Medd., 126: 1-177.
Reeder, W. G., and K. S. Norris. 1954. Distribution, type locality, and habits of the
fish-eating bat, Pizonyx vivesi. Jour. Mamm., 35: 81-87.
Scott, H. 1939, A new species of Nycteribiidae (Diptera Pupipara) from islands in
the Gulf of California. Allan Hancock Foundation, Allan Hancock Pacific
Expeditions, 2: 167-172.
SuTHERS, R. A. 1965. Acoustic orientation by fish-catching bats. Jour. Exp. Zool.,
158: 319-347.
. 1967. Comparative echolocation by fishing bats. Jour. Mamm., 48: 79-87.
Tate, G. H. H. 1941. Results of the Archbold Expeditions. No. 39. A review of the
genus Myotis (Chiroptera) of Eurasia, with special reference to species occur-
ring in the East Indies. Amer. Mus. Nat. Hist., Bull., 78: 537-565.
Walker, L. W. 1950. The fish bats of Pescadora. Audubon Mag., 52: 294-299.
Accepted for publication February 16, 1970
LOS
ANGELES
COUNTY
MUSEUM
CONTRIBUTIONS
IN SCIENCE
^^UMBER 184
April 17, 1970
i CONOCARDIUM LANGENHEIMI SP. N.
i;
(MOLLUSCA: BIVALVIA) IN THE LOWER PERMIAN SERIES
I OF THE McCLOUD LIMESTONE, NORTHERN CALIFORNIA
i
1
!
I By Edward C. Wilson
I
;t
li
I'
I
Los Angeles County Museum of Natural History • Exposition Park
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CONTRIBUTIONS IN SCIENCE is a series of miscellaneous technical papers
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CONOCARDIUM LANGENHEIMl SP. N.
(MOLLUSC A: BIVALVIA) IN THE LOWER PERMIAN SERIES
OF THE McCLOUD LIMESTONE, NORTHERN CALIFORNIA
By Edward C. Wilson^
Abstract: Conocardiiim langenheimi sp. n, (Mollusca: Bi-
valvia) occurs in the Wolfcampian Stage, Lower Permian Series
of the McCloud Limestone in Shasta County, California. It is a
large shell with a non-carinate posterior end, three pairs of perfo-
rate “shelves” within the gaping anterior end, a tube enclosed dor-
sally in the shell layers of each valve, unique internal hinge
structures, and microstructure unlike most other bivalves. The
functions of these and other characters is unknown. The entire
Order Conocardidoida needs investigation to determine the signi-
ficance of such characters, especially for placing the group in
higher categories of classification.
Introduction
At the suggestion of R. L. Langenheim, Jr., Professor of Geology at the
University of Illinois, I began a study of the paleontology of the McCloud
Limestone in Shasta County, California, in the spring of 1963. Two previous
papers (Wilson, 1967a, 1967b) resulted from this study and this paper is
another contribution.
Acknowledgments
I am grateful for opinions on the morphology and systematic position of
the Conocardidoida from J. Wyatt Durham of the University of California
at Berkeley, John Pojeta, Jr. of the U.S. National Museum, and Norman D.
Newell of the American Museum of Natural History. Professor Durham
kindly visited the type localities with me. Most of the photographs are by
Lawrence S. Reynolds of the Los Angeles County Museum of Natural History.
Field work in 1963 and 1964 was financed in part by the University of
California Museum of Paleontology (UCMP) and the University of California
Committee on Research, both at Berkeley. The Los Angeles County Museum
of Natural History (LACM) provided funds for further field work in 1968.
^Curator of Invertebrate Paleontology, Los Angeles County Museum of Natural
History.
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Contributions in Science
No. 184
SYSTEMATIC DESCRIPTION
Phylum MOLLUSCA
Class BIVALVIA Linnaeus, 1758
Order CONOCARDIDOIDA Neumayr, 1891
Family CONOCARDIIDAE Miller, 1889
Genus Conocardium Bronn, 1834
Remarks: Terminology has not been standardized for the unique char-
acters of the Conocardidoida. I have used where possible the terminology for
other bivalve mollusks, although this is partially unsatisfactory. The structures
called “auricles” are probably not homologous with the auricles of other
bivalves. The “shelves” in the gaping anterior “auricle” are without parallel
in the rest of the Bivalvia. The openings in the “shelves” and the tubes en-
closed in the shells have no formal names. The kind of “teeth” on the ventral
and posterior edges should probably be named. It is not even possible to use
uniformitarianism to orient the shell with certainty. It may seem reasonable
that dorsal and ventral are accurately delineated, but there is no such certainty
about anterior and posterior. The figures and legends indicate usage of mor-
phological terms in this paper.
Conocardium langenheimi, sp. n.
Fig. 1
Description {external) : Valves equivalve, inequilateral, opisthogyrate,
alate (?), not ankylosed; main body of valves inflated, rounded (not carinate
posteriorly); posterior “auricle” small, sloping smoothly into main body of
valves, not gaping; anterior “auricle” separated by constriction from main
body of valves, large, rostrate, gaping anteriorly and ventrally, with large
trigonal aperture having scalloped margins; hinge line straight, except anterior
part of anterior “auricle” may dip to 25° (preservation?), occupied by groove,
shallow in anterior part of anterior “auricle,” deep in posterior part of an-
terior “auricle” and main body of shell; commissure straight on exterior of
ventral parts of shells; holotype shell length 6.5 cm, height 4.3 cm, width 2.9
cm (largest specimen, LACM paratype 2443, is 4 mm. higher, but incomplete
Figures 1-4. Conocardium langenheimi sp. n. UCMP holotype 10589, UCMP
locality D-831, all figures X 1.3; (1) dorsal view, anterior “auricle” at left, posterior
“auricle” at right, main body of shell inflated, openings in umbos caused by post-
burial weathering, double wall caused by dissolution in HCl of calcareous layer
between silicified layers, note absence of posterior carina; (2) ventral view, showing
straight exterior commissure of main body, gape of anterior auricle, internal ridge at
hinge line; (3) lateral view of left valve, sculpture on main body possibly dissolved in
part during etching; (4) ventral view, with piece of shell removed to show dorsal
interior, dorsal ridge along hinge line of main body broken with fragments lying in
right valve, note trigonal cross sections of rdiges remaining in shell wall after etching.
1970 Conocardium langenheimi Sp. N. (Mollusca: Bivalvia)
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Contributions in Science
No. 184
in other dimensions) ; sculpture concentric or cancellate (?) on main body of
valves, with about 4 evenly spaced concentric riblets in one mm., with possible
traces of radial riblets; sculpture radial on “auricles” (UCMP paratype 10590,
juvenile, shows some concentric), about 17 ribs on anterior “auricle” of each
holotype valve, fewer on posterior “auricle” (UCMP paratype 10590, juvenile,
has 9 ribs on each posterior “auricle”) ; ribs straight to very gently sigmoid,
sloping down from hinge line at angles varying from 65° on posterior part of
anterior “auricle” to 25° on anterior part, flat topped with straight to rounded
sides sloping down and out at less than 45°; rib interspaces about equal to rib
widths, lirate, with about 20 dorsally convex lirae in 1 mm, rarely occupied
by single riblet.
Description (internal) ; Main body of valves smooth; pallial line, muscle
scars not observed; ventral and posterior commissure zig-zag, formed by series
of trigonal “teeth” and intervening sockets in each valve, 3 occurring every
5 mm each side of both valves (holotype) , sockets showing plainly only in
section (“teeth” are surficial extensions of radial rods, trigonal in cross sec-
tion, apex pointing inward, that dorsally form part of the internal shell layer) ;
hinge line a complex ridge, low in anterior end, becoming higher posteriorly,
developing horizontal distal bar (inverted T-shaped in cross section) within
main body cavity of valves, where narrow trough borders each side (marking
lateral limits of apparent separate hinge plate in this area), becoming lower
and broader posteriorly, troughs continuing alongside into posterior “auricle,”
where termination not observed because of preservation; 3 pairs of complex
“shelves” lining lateral interior of anterior “auricle,” sloping in and up same
angles as exterior ribs, each pair meeting in midline, dividing “auricle” roughly
horizontally into 4 unequal areas; dorsal pair of shelves beginning about 15
mm back of “auricle” anterior edge, about 8 mm below top of “auricle,” form-
ing large heart shaped opening in cross section above “shelves,” turned up
along midline by lip-like ridge, sloping upward, separating along midline below
Figures 5-9. Conocardium langenheimi sp. n. (5) holotype, anterior view into gaping
“auricle” showing 3 pairs of “shelves,” X 1.3; (6) same specimen, posterior view,
umbos partially weathered away, note zig-zag commissure of “auricle,” straight com-
missure of main body, X 1.3; (7) same specimen, same view as figure 5 with part of
auricle removed, missing parts of wall dissolved during etching, note exterior groove
and interior ridge of hinge line with grooves bordering latter, dorsal “shelves” meet-
ing along midline with opening into main body visible just above, middle shelves
similar but extending anteriorly to form keyhole shaped opening, partly obscured
small ventral “shelves” shaped like middle “shelves,” X 3.6; (8) LACMNH paratype
2443, LACMNH locality 1 133, fragment showing zig-zag interior ventral commis-
sure formed of series of teeth and sockets, note left valve showing original smooth
interior, right valve showing etched layer of shell wall with internal trigonal ridges
which terminate ventrally as teeth, X 0.6; (9) UCMP paratype 10590, UCMP local-
ity D-831, lateral view of right valve of juvenile specimen, note concentric sculpture
on main body and anterior auricle, offset posterior “auricle,” outline of anterior
“auricle” varying from figure 3, X 1.4.
1970 Conocardium langenheimi Sp. N. (Mollusca: Bivalvia)
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hinge line, leaving opening here into main part of shell, shelves continuing
dorsally and posteriorly with lips incurving to form tubes passing into shell
layers before leaving “auricle,” tubes continuing posteriorly in shell above main
body of shell, lateral to hinge line, becoming closer posteriorly and apparently
dropping into external groove above hinge line, with posterior course un-
known due to preservation (no apparent natural external opening observed
in posterior “auricle” or hinge line); middle “shelves” also large, paralleling
dorsal ones, similar in shape and slope, but extending to anterior edge of
“auricle,” forming key-hole shaped opening in anterior ends as well as heart-
shaped opening posteriorly into main body of shell, with anterior lips as in
dorsal “shelves,” extending into grooves in “auricle” walls under dorsal
“shelves” posteriorly, ending at posterior termination of dorsal “shelves,”
without continuing internal tube; ventral “shelves” small, shaped like middle
“shelves.”
Description {microstructure) : Preservation very imperfect, shell par-
tially silicified, some layers separated by silica, some missing (erosion?) in
places; shell calcium carbonate, 1.5 to 3.5 mm thick, with 3 layers throughout
plus 3 highly restricted layers; exterior layer 0.33 to 1.0 mm thick, prismatic,
distal crystal tips reflected ventrally, elsewhere normal to shell surface; middle
layer 0.2 to 0.3 mm thick (generally replaced by silica and difficult to measure) ,
lamellar, showing ventrally convex concentric growth lines crossed by finer
radial lines, separated in places from prismatic layer by dark line (conchio-
lin?); inner layer 2 to 3 mm thick, much wider than other layers, formed of
large calcium carbonate crystals (apparently recrystallized), showing residual
(?) growth lines in some areas, etching in dilute hydrochloric acid in main
body of shell into trigonal ridges (see above), evidence of which not observed
in polished or thin sections; myostracum (?) in 3 bands in each valve, extend-
ing from opposed axial edges of “shelves” to umbos, straight to V-shaped in
Figures 10-18. Conocardium langenheimi sp. n. UCMP paratype 10591, UCMP
locality D-831, specimen ground down from anterior to posterior to show successive
cross sections, all figures X 2.2; (10) section near anterior of anterior “auricle,” note
great shell thickening (left) from thin anterior edge (right); (11) dorsal “shelves”
begin to form, note rod like myostracum (?) protruding from “shelves,” internal
scalloped outline of outer shell layers; (12) three pairs of “shelves” forming, note
deepening grooves between dorsal and middle “shelves”; (13) three pairs of “shelves”
formed here, each with associated myostracum (?); (14) upper “shelves” move
dorsally, curl inwards, middle “shelves” pass into two grooves, lower “shelves” have
disappeared, internal dorsal ridge along hinge line begins to lengthen; (15) dorsal
“shelves” move dorsally, other “shelves” disappear (groove extending from middle
“shelf” on right still present), rods of myostracum (?) move dorsally in shell wall,
teeth and sockets appear ventrally marking ventral beginning of main body of valves;
(16) dorsal ridge lengthens, pre-burial erosion noticeable on dorsal edge of valve at
left; (17) dorsal ridge lengthens; (18) dorsal ridge lengthens, dorsal edge of eroded
valve thins, dorsal “shelves” move higher, accompanied by their myostracum (?)
rods and those from two lower “shelves.”
1970 Conocardium langenheimi Sp. N. (Mollusca: Bivalvia)
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No. 184
cross section (varies), about 0.2 mm thick, apparently exposed at “shelf”
edges (indicates muscles joining opposing “shelves”?), showing concentric
and radial lines as in middle layer.
Name: This species is named for R. L. Langenheim, Jr. in recognition
of his contributions to our knowledge of the paleontology and stratigraphy of
North America and for his inspiration and training of many students in this
work.
Type specimens: Holotype UCMP 10589, paratypes UCMP 10590-
10591, paratype LACM 2443, all UCMP locality D-831 (=LACM locality
1133);paratype UCMP 10592, UCMP locality D-819.
Localities: UCMP locality D-819. 1140 feet stratigraphically above the
lowest exposure of the McCloud Limestone on the first spur south of the
eastern end of the McCloud River bridge as shown in the northwest 14 of the
southwest Vat of section 32, township 35 north, range 3 west of the United
States Geological Survey topographic quadrangle (15 minutes, 1957) of Bol-
libokka Mountain, Shasta County, California. Series of one foot high ledges
of limestone, fine to medium grained, black to medium gray, weathering
medium to light gray; little chert; many silicified fossils, including large cani-
noid corals, Conocardium, Omphalotrochus, fusulinids.
UCMP locality D-831 (= LACM locality 1133). Estimated 200 yards
south of locality D-819 along strike and about 15 feet higher in the section.
Narrow bench of thin-bedded, black, silty limestone below a sheer massive
cliff. Abundant syringoporid corals, very large solitary rugose corals, fusu-
linids, bryozoans, Conocardium.
Age: The fusulinids have the most refined stratigraphic ranges of the
fossils found with Conocardium langenheimi. Two species were identified:
Pseudofusulina soluta Skinner and Wilde, 1965 and Triticites mulleri Skinner
and Wilde, 1965, both having type localities in the McCloud Limestone nearby
the Conocardium localities. Skinner and Wilde (1965, pi. 4) considered the
former fusulinid to indicate definite Permian age (the genus is a Wolfcampian
index fossil) and the latter to be of either Pennsylvanian (Virgilian) or Per-
Figures 19-24. Conocardium langenheimi sp. n. Continuation of sections of same
specimen as figures 10-18, same magnification; (19) dorsal “shelves” have incurved
and passed into tubes in shell wall, marking dorsal beginning of main body of valves,
myostracum (?) rods now wholly within shell wall, dorsal ridge greatly lengthened
and now having horizontal distal end (not clear in this photograph but see fig. 34),
possibly developing into separate plate along hinge line, with lateral grooves inter-
nally, midline groove externally, tube at left opened by erosion; (20) dorsal tubes
narrow, note ventral teeth showing how ventral commissure can be exteriorly straight
yet interior zig-zag, also shows shell not ankylosed; (21 ) dorsal tubes narrow, dorsal
ridge shortens; (22) dorsal tubes narrow and move toward midline, dorsal ridge
shortens; (23) dorsal tubes narrow, valve at left badly eroded; (24) dorsal tubes
smaller, dorsal ridge shorter, dorsal part of valves move ventrally, deepening external
groove.
1970 Conocardium langenheimi Sp. N. (Mollusca: Bivalvia)
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Contributions in Science
No. 184
mian (Wolfcampian) age. It seems clear, therefore, that the interval of the
McCloud Limestone from which C. langenheimi was collected is definitely
occupied by Lower Permian Series, Wolfcampian Stage rocks.
Diagnosis’. C. langenheimi can be separated readily from most other
species because it lacks a carina on the posterior part of the main body of the
shell. It is also larger than most species and the anterior “auricle” is propor-
tionately longer in relation to the rest of the shell than in many other species.
Discussion: C. langenheimi resembles closely only C. ouralicum (Ver-
neuil, in Murchison, Verneuil, and Keyserling, 1845, p. 301, pi. 20, figs. 11a,
1 lb) from Upper Carboniferous rocks of the Ural Mountains. Both species
lack posterior carinae and have elongate anterior “auricles.” However, C.
ouralicum is smaller (length 36 mm) and has an anterior “auricle” that is much
more elongate in proportion to its height than that of C. langenheimi. Internal
structures and shell microstructure of C. ouralicum are unknown as yet.
The microstructure of C. /ang^n/z^/m/ apparently lacks “roofed-over pits”
like those figured by Pannella and MacClintock (1968, pi. 8, fig. 5) in C. sp.
from Upper Pennsylvanian rocks of Oklahoma. The “outer complex-prismatic
shell layer” shown by Pannella, MacClintock, and Thompson (1968, fig. 1 A)
of C. herculeum Konick from Lower Carboniferous rocks of Belgium re-
sembles a similarly positioned layer in C. langenheimi, but lacks the reflected
tips that are present in the latter species.
The presence of complex internal structures in Conocardium was early
reported by Hind (1900, p. 451, pi. 51, fig. 11a) in some detail, although he
apparently worked from specimens somewhat worn internally. Curiously, no
further refined work on them seems to have been published, although Decha-
seaux (1952, p. 319, fig. 162a) and Branson, LaRoque, and Newell (1969,
p. 859, fig. Gl) cursorily discussed and figured some internal characters.
Future workers should examine the internal structure of specimens of Con-
ocardium with great care.
I have tried to find clues that would help determine the habitat and
functional morphology of C. langenheimi. The associated fossils show that the
salinity of the water must have been whatever was normal for the open oceans
in Permian time. Permian near-shore faunas with restricted access to the sea in
western North America generally lacked corals and fusulinids, both of which
Figures 25-29. Conocardium langenheimi sp. n. Continuation of sections of same
specimen as figures 10-24, same magnification; (25) dorsal ridge shorter but hori-
zontal expansions of distal end still present, dorsal tube openings smaller (not seen in
valve at left), some of dorsal part of valve at right shows erosion; (26) dorsal tube
moves inward and downward, dorsal ridge shortens, distal expansions disappear,
erosion of both valves and distortion of valve at left apparent; (27) continuation of
movements seen in figure 26; (28) continuation, myostracum (?) rod (dark chevron)
shows beside eroded tube in valve at right; (29) dorsal tubes move ventrally, hinge
line plate broadens.
1970 Conocardium langenheimi Sp. N. (Mollusca: Bivalvia)
11
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Contributions in Science
No. 184
are abundant at the C. langenheimi localities. The shell itself is sturdy, es-
pecially the thick walled anterior “auricle,” which suggests exposure to some
strong current or wave action. Seemingly eroded parts of some of the speci-
mens indicates that they may have been moved by currents from their original
habitats. However, some fine silty layers in the limestone indicate that sedimen-
tation was generally not rapid. None of the specimens shows evidence of
having been buried in a burrow, which seems to me the most likely habitat by
analogy with Recent bivalves of the most similar external form, size, and
sturdiness (Pholadidae). Branson (1969, p. 860, figs. 3a, 3b, in Branson,
LaRoque, and Newell) figured, but did not discuss, a restoration of a Cono-
car dium apparently occupying a burrow, but having the anterior “auricle”
innermost, which is unlike the Pholadidae. Nicol (1970, p. 70) also compared
Conocardium with Recent bivalves, but reached the different conclusion that
it “was attached by a short, stout byssus . . .” All the specimens of C. langen-
heimi appeared to be lying randomly on surfaces parallel to the bedding planes.
They are all tightly closed pairs, which is unlike other Bivalvia, if one assumes
because of the eroded areas that they were buried after death.
Reconstruction of the soft part morphology of C. langenheimi will not be
simple. I have been unable to create a satisfactory model for incurrent and
excurrent siphons. The complex form of the “shelves” indicates that some
extremely specialized soft part morphology was present. The apparent absence
of muscle scars in the anterior end may be explained by the existence of
muscles between the edges of opposing “shelves,” as suggested by the myo-
stracum (?) exposed there. One would expect posterior muscles also, which
seemingly are not represented by muscle scars or a posterior myostracum.
Perhaps the function of the unique hinge line structures and the ventral and
posterior “teeth” rendered posterior muscles unnecessary. The internal ridge
along the hinge line looks greatly like a support, but one wonders for what,
since the other bivalves seem to function well without such a rigid support for
the ctenidia, alimentary canal, or other systems. It is tempting to postulate a
Figures 30-34. Conocardium langenheimi sp. n. (30) continuation of sections of
same specimen as figures 10-29, same magnification, dorsal tubes drop into exterior
grooves bordering hinge line, dorsal ridge disappears; (31) same specimen, same
magnification, dorsal tubes not present (preservation?), external grooves coalesce,
with narrowed opening above, hole in shell at left caused by erosion, shell not pre-
served posterior to this section and nature of structures there unknown; (32) thin
section made from polished section shown in fig. 11, note especially exposed end of
myostracum (?) at edge of “shelf” at left and indeterminate nature of hinge articula-
tion, X 3.2; (33) thin section made from polished section shown in fig. 31, showing
nature of hinge and apparent absence of dorsal tubes seen in figs. 19-30, X 3.2;
(34) thin section made from polished section shown in fig. 21, showing dorsal tubes,
myostracum (?) layers, elongate internal hinge line ridge with horizontal distal bar,
exterior ligament (?) groove, and apparent separate nature of plate along hinge
line, X 3.2.
1970 Conocardium langenheimi Sp. N. (Mollusca: Bivalvia)
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Contributions in Science
No. 184
function such as brood chambers to the tubes in the dorsal parts of the valves,
but, again, close analogies in living bivalves seem absent. Some of the grooves
in the shell suggest ciliated tracts, but there is no certainty about this.
Conclusions
C. langenheimi is a member of the Conocardidoida, which is externally
like the Bivalvia, has a world wide distribution, and ranges from Ordovician
through Permian (Triassic?) rocks. The degree of complexity shown by the
shell microstructure definitely appears to be molluscan. The paired valves sug-
gest that the Order belongs in the Class Bivalvia, however the internal mor-
phology and shell microstructure is unlike others in that class. If typical of
the order, these suggest need for a reconsideration of the systematic position
at the class or higher level.
Literature Cited
Branson, C. C., A. LaRocque, and N. D. Newell. 1969. Order Conocardioida, in
Treatise on invertebrate paleontology, R. C. MOORE, ed., Bivalvia: Part N.
Geol. Soc. America and Univ. Kans. Press, Lawrence, Kansas. 2:491-951,
figures.
Dechaseaux, Colette. 1952. Etude systematique, in Traite de Paleontologie, J.
PIVETEAU, ed. Masson et Cie., Paris. 2:261-364, figs. 35-215.
Hind, W. 1900. A monograph of the British Carboniferous Lambellibranchiata, part
V. Paleont. Soc., London. 361-476, pis. 40-54.
Linnaeus, Carolus. 1758. Systema naturae. Editio decima. Laurentii Salvii, Stock-
holm. 1-824.
Miller, S. A. 1889. North American geology and palaeontology for the use of
amateurs, students, and scientists. Western Methodist Book Concern, Cincinnati.
664 pp., 1 194 figs.
Murchison, R. I., E. De Verneuil, and A. De Keyserling. 1845. Geologic de la
Russie d’Europe et des Montagnes de I’Oural. Paleontologie. John Murray,
London. Bertrand, Paris. 2(3) : 1-512, 50 pis.
Neumayr, M. 1891. Beitrage zur morphologischen Einteilung der Bivalven. Denk-
schr. Acad. Wiss. Wien Math.-Nat. Cl. 58:701-801.
Nicol, David. 1970. Mode of life of Conocardiiim, a Paleozoic pelecypod. Nautilus
83:77-80, figs. 1-5.
Pannella, G. and C. MacClintock. 1968. Biological and environmental rhythms
reflected in molluscan shell growth. Jour. Paleontol. 42:64-80, pis. 1-9.
Pannella, G., C. MacClintock, and M. N. Thompson. 1968. Paleontological evi-
dence of variations in length of synodic month since Late Cambrian. Science
162:792-796, figs. 1, 2.
Skinner, J. W. and G. L. Wilde. 1965. Permian biostratigraphy and fusulinid faunas
of the Shasta Lake area, northern California. Univ. Kans. Paleontol. Contr.,
Protozoa 6:1-98, pis. 1-65.
Wilson, E. C. 1967a. Fusulina gracilis Meek, 1864 (Foraminiferida) : proposed
invalidation of neotype-material, designation as type-species of Eoparafusulina
Coogan, 1960, and related actions. Bull. Zool. Nomencl. 24:234-236, pi. 1.
. 1967b. Primary type-material of the Early Permian archaeogastropod
Omphalotrochus whitneyi (Meek, 1864). Jour. Paleontol. 41 : 1 113-1 1 15, pi. 142.
Accepted for publication February 17, 1970
! roH.es CONTRIBUTIONS
z::z IN SCIENCE
Dumber 185 April 17, 1970
: FIVE NEW EPITONIID GASTROPODS FROM THE
|i WEST COAST OF THE AMERICAS
i|
ji
By Helen DuShane
ii
ii
!i Los Angeles County Museum of Natural History • Exposition Park
:j 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 sep-
arately, and numbers run consecutively regardless of subject matter. Number 1 was
issued January 23, 1957. The series is available to scientific institutions and scien-
tists on an exchange basis. Copies may also be purchased at a nominal price. Inquiries
should be directed to Virginia D. Miller, Los Angeles County Museum of Natural
History, 900 Exposition Boulevard, Los Angeles, California 90007.
Robert J. Lavenberg
Managing Editor
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. Al-
though priority will be given to manuscripts by staff members, or to papers dealing
largely with specimens in the Museum’s collections, other technical papers will be
considered. All manuscripts must be recommended for consideration by the curator
in charge of the proper section or by the editorial board. Manuscripts must conform
to those specifications listed below and will be examined for suitability by an Edi-
torial Committee. They may also be subject to review by competent specialists out-
side the Museum.
Authors proposing new taxa in a CONTRIBUTIONS IN SCIENCE must
indicate that the primary type has become the property of a scientific institution of
their choice and cited by name.
MANUSCRIPT FORM.— ( 1 ) The 1964 AIBS Style Manual for Biological
Journals is to be followed in preparation of copy. (2) Double space entire manu-
script. (3) Footnotes should be avoided if possible. Acknowledgments as footnotes
will not be accepted. (4) Place all tables on separate pages. (5) Figure legends and
unavoidable footnotes must be typed on separate sheets. Several of one kind may
be placed on a sheet. (6) An abstract must be included for all papers. This will be
published at the head of each paper. (7) A Spanish summary is required for all
manuscripts dealing with Latin American subjects. Summaries in other languages
are not required but are strongly recommended. Summaries will be published at
the end of the paper. (8) A diagnosis must accompany any newly proposed taxon.
(9) Submit two copies of manuscript.
ILLUSTRATIONS.- — All illustrations, including maps and photographs, will
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will be included with the galley proof.
Virginia D. Miller
Editor
FIVE NEW EPITONIID GASTROPODS FROM THE
WEST COAST OF THE AMERICAS
By Helen DuShane^
Abstract: Five epitoniids (Mollusca: Gastropoda) are de-
scribed as new, including {Asperiscala) longinosanum from the
west coast of Baja California, Mexico; Epitonium (Asperiscala)
macleani from Ventana Bay, Baja California, Mexico; Epitonium
(Acirsa) cerralvoensis from Cerralvo Island, Baja California,
Mexico; Epitonium (Acirsa) murrha from Panama; and Amaea
contexta from Guerrero, Mexico.
The epitoniid collections of the Los Angeles County Museum of Natural
History (LACM), including those now on loan from the Allan Hancock
Foundation, contain five new species described here. Two of the new species
were dredged by the R/V Valero III in the 1930’s and are from the Hancock
Foundation Collection. Two are from a 1966 Museum expedition to the Cape
San Lucas area of Baja California, sponsored by Mr. Richard Dwyer of
Corona del Mar, on the R/V Sea Quest. One other new species was collected
by Fred E. Lewis and remained unidentified until now.
Acknowledgments
I am grateful to the Los Angeles County Museum of Natural History for
the opportunity to study their collections and for the help rendered by Dr.
James H. McLean and Mr. Gale Sphon of the Invertebrate Zoology section. I
also thank the photography department of the Museum for the illustrations.
Epitonium (Asperiscala) longinosanum, sp. nov.
Fig. 1
Diagnosis: An elongate species with 1 1 low varices differing from all
other epitoniids from the Panamic molluscan province in having a brown shell
with white ribs and unevenly spaced minute spiral striae.
Description: Shell large, tall and slender, thin but strong, medium brown
in color, shining, with 1 1 gradually enlarging, rounded whorls, nuclear whorls
missing; suture distinct, varices white, 11, low and slightly reflected with a
small, sharp spine on shoulder of whorl, varices continuous from whorl to
whorl, an occasional costa thicker than others; spiral striations uneven, visible
only under 10 X magnification; columellar lip curved, peritreme continuous,
reflected over the ends of the varices, outer lip thickened by last varix; um-
^Research Assistant in Invertebrate Zoology, Los Angeles County Museum of
Natural History, Los Angeles, California 90007.
1
2
Contributions in Science
No. 185
Figures 1-2. 1, Epitonium (Asperiscala) longinosanum sp. nov., holotype, LACM
1 196. X 3; 2, Epitonium (Asperiscala) macleani sp. nov., holotype, LACM 1 197. X 9.
bilious lacking, operculum missing in holotype. Dimensions (in mm) : length
16.5, width 5.5.
Type Locality: Rancho Inocentes, Baja California del Sur, Mexico, 23°
45' N, 1 10° 40' W. Collected by Captain Fred E. Lewis prior to 1945, one
specimen.
Type Material : Holotype, LACM 1196.
Discussion: The coloring and 1 1 varices with sharp, shoulder spines
distinguish E. longinosanum from all other west American epitoniids. It differs
from E. appressicostatum Dali, 1917 (p. 482) , which it most resembles, in
having fewer costae, a less glassy surface, and fine spiral striations between
the costae.
Epitonium longinosanum is named for Jose Longinos Martinez who es-
tablished a museum of natural history in Mexico City in 1791 and the follow-
ing year collected shells from La Paz to San Francisco (Longinos Martinez,
1938).
1970
Five New Gastropods
3
Epitonium (Asperiscala) macleani, sp. nov.
Fig. 2
Diagnosis: A small species differing from all others in having rounded
whorls, numerous low axial costae of differing strengths and numerous fine
spiral ribs.
Description: Shell small, white, fragile, with approximately 32 fine costae
of varying size, nuclear whorls two, smooth, rounded, light brown, first
postnuclear whorl with axial striations; remaining whorls six; suture distinct,
deeply impressed; umbilicus small and nearly covered by peritreme; surface
between whorls with raised spiral threads, approximately 28 on the body
whorl; whorls rapidly expanding, rounded, basal disk or cord lacking; aperture
oval (lip broken in holotype) ; operculum unknown. Dimensions (in mm) :
length 7.0; width 3.5 (holotype) .
Type Locality: Off Punta Ventana, Baja California del Sur, Mexico
(opposite Cerralvo Island), 24° 4' N, 109° 49' W, 15-25 fms, sand bottom,
LACM Invertebrate Zoology locality 66-23, James H. McLean, Peter M.
Oringer, and Louis Marincovich, R/V Sea Quest, 8 April 1966, ten specimens.
Type Material: Holotype, LACM 1197; 8 paratypes, LACM 1198; 1
paratype, DuShane Collection.
Discussion: This species differs from all others known from the Panamic
province. Less globose than the brownish Epitonium (Asperiscala) huffmani
DuShane and McLean, 1968 (fig. 1), it is white and has somewhat similar
sculpture.
Epitonium (Asperiscala) macleani is named for Dr. James H. McLean,
Curator of Invertebrate Zoology, Los Angeles County Museum of Natural
History.
Epitonium (Acirsa) cerralvoensis, sp. nov.
Fig. 3
Diagnosis: A slender, pale brown species differing from other epitoniids
from the Panamic molluscan province in having reduced axial costae and
unevenly spaced spiral striae.
Description: Shell of medium size, tall and slender, thin but strong, light
brown in color, with 10-12 gradually enlarging, rounded whorls, two opaque
nuclear whorls; suture moderately impressed; axial ridges pale brown, 16 on
the fourth whorl, gradually becoming obsolete on succeeding whorls, spiral
ridges wider than the interspaces, 19 on last whorl; peritreme incomplete, lip
thin, columella arched; no basal ridge; operculum horn colored, paucispiral.
Dimensions (in mm) : length 1 1.8, width 3.7 (holotype) .
Type Locality : West side, Cerralvo Island, Baja California del Sur,
Mexico, 24° 11' N, 109° 55' W, 7 fms, sand bottom, LACM Invertebrate
Zoology locality 66-24, James H. McLean, Peter M. Oringer, and Louis
Marincovich, R/V Sea Quest, 9 April 1966, two specimens.
4
Contributions in Science
No. 185
Figures 3-5. 3, Epitonium (Acirsa) cerralvoensis sp. nov., holotype, LACM 1200.
X 5; 4, Epitonium (Acirsa) murrha sp. nov., holotype, LACM 1232. X 3; 5, Amaea
(s.l.) contexta sp. nov., holotype, LACM 1 199. X 4.
Type Material : Holotype, LACM 1200; 1 paratype, LACM 1201. Al-
though larger (15.3 mm in length), the paratype is worn and has several holes
in the shell.
Referred Material : LACM 66-17, 10-20 fms off Cape San Lucas, Baja
California, 1 specimen; LACM 66-17, 10-20 fms off Rancho Palmilla, Baja
California, 1 specimen; LACM 66-20, 4 fms, Pulmo Bay, Baja California, 1
specimen; AHF 1051-40, 21 fms off Puerto Refugio, Angel de la Guardia
Island, Baja California, 1 specimen.
Discussion: No other epitoniid from the west coasts of the Americas has
sculpture like this species.
Epitonium (Acirsa) cerralvoensis is named for Cerralvo Island, its type
locality.
Epitonium (Acirsa) murrha, sp. nov.
Fig. 4
Diagnosis: A large species with 19 low axial ribs over which the spiral
sculpture crosses; differing from other Acirsa from the west coasts of the
Americas in having more pronounced axial sculpture on the body whorl.
Description: Shell of medium size, white, texture of opaque china;
1970
Five New Gastropods
5
nuclear whorls two, opaque (first nuclear whorl missing in holotype), first
postnuclear whorl with faint indication of axial sculpture; second postnuclear
whorl with incised spiral lines; remaining convex whorls 10, gradually en-
larging; 19 retractive axial ridges on body whorl giving a sinuous appearance;
penultimate whorl with 15 spiral cords, uneven in width, separated by narrow,
incised lines riding over the axial ridges; suture moderately impressed; basal
ridge with 10-11 flat cords separated by incised lines; aperture oval, peritreme
incomplete, umbilicus lacking, operculum unknown. Dimensions (in mm) :
length 19.1, width 6.9 (holotype).
Type Locality: Ten miles southwest of Secas Islands, Panama, 7° 5T 10"
N, 82° 12' 05" W, 30 fms, gray sand, R/V Velero III station 944-39, 27 March
1939, one specimen.
Type Material: Holotype, LACM-AHF 1232.
Discussion: No other epitoniid from the west American coast has the
color or sculpture of this species. The north Atlantic species, Acirsa costulata
Mighels and Adams, 1842, figured by Clench and Turner (1950, p. 230) is
similar in proportions but has weaker axial and spiral sculpture.
The name murrha, a Latin noun, meaning porcelain clay, refers to the
chinalike surface of the shell. It is intended as a noun in apposition.
Amaea (s.L) contexta, sp. nov.
Fig. 5
Diagnosis: An epitoniid differing from all other species in having many
whorls, a dark brown color, numerous low axial ribs, and coarse spiral ribs.
Description: Shell of medium size, slender, dark brown, with four
smooth, white, glassy nuclear whorls (missing in holotype), first postnuclear
whorl axially ribbed; the following 10 whorls dark brown, convex; suture
deeply impressed; axial sculpture of thin, strong ribs, about 24 on the body
whorl, with occasionally a very heavy rib; these ribs continue to the columellar
wall; spiral sculpture of about 18 ridges per whorl, beaded on the anterior face
of the ribs; basal ridge present; outer lip formed by the extension of the last
varix, aperture oval, umbilicus lacking; operculum lacking in holotype. Dimen-
sions (in mm) : length 15.4, width 4.5 (holotype).
Type Locality: Petatlan Bay, Guerrero, Mexico, 17° 31' 45" N, 101° 27'
34" W, 5-10 fms, hard sand and shell bottom, R/V Velero III station 265-34,
3 February 1934, one specimen.
Type Material: Holotype, LACM 1199.
Referred Material : Two smaller, damaged specimens with intact nuclear
whorls were dredged by Laura and Carl Shy off Manzanillo, Colima, Mexico,
in 5-10 fms, November 1967. These two specimens are uniformly dark brown,
suggesting that the color of the holotype has faded.
Discussion : No other epitoniid from the west American coast is related
6
Contributions in Science
No. 185
to this species. Sculpture is unique, making the generic allocation uncertain.
Since the general features suggest Amaea H. and A. Adams, 1853, it is placed
here provisionally.
The Latin name is derived from contextus, meaning interwoven, and
refers to the woven texture of the axial and spiral sculpture.
Resumen
Cinco epitonidos (Mollusca: Gastropoda) se describen como nuevas
especies, incluyendo (Asperiscala) longinosanum de la costa oeste de la Baja
California, Mexico; Epitonium {Asperiscala) macleani de la Bahia de Ventana,
Baja California; Mexico; Epitonium (Acirsa) cerralvoensis de la Isla Cer-
ralvo, Baja California, Mexico; Epitonium (Acirsa) murrha procedente de
Panama; y Amaea contexta de Guerrero, Mexico.
Literature Cited
Clench, W. J., and R. D. Turner. 1950. The genera Sthenorytis, Cirsotrema,
Acirsa, Opalia and Amaea in the western Atlantic. Johnsonia, 2: 221-248.
Dale, W. H. 1917. Notes on the shells of the genus Epitonium and its allies of the
Pacific coast of America, U.S. Nat. Mus., Proc., 53: 471-488.
DuShane, Helen, and J. H. McLean. 1968. Three new epitoniid gastropods from
the Panamic Province. L.A. County Mus. Nat. Hist., Contrib. Sci., 145: 1-6.
Longinos Martinez, Jose. 1938. California in 1792. The expedition of Jose Longi-
nos Martinez, transl. by L. B. Simpson. Huntington Library, San Marino, Calif.
1 1 1 pp.
Accepted for publication February 11, 1970
LOS
ANGELES
COUNTY
MUSEUM
CONTRIBUTIONS
IN SCIENCE
UMBER 186
April 17, 1970
A GENERIC REVISION AND CATALOG OF THE WESTERN
HEMISPHERE GLENURINI WITH THE DESCRIPTION OF A
NEW GENUS AND SPECIES FROM BRAZIL
(NEUROPTERA : MYRMELEONTIDAE)
By L. A. Stance
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 sep-
arately, and numbers run consecutively regardless of subject matter. Number 1 was
issued January 23,. 1957. The series is available to scientific institutions and scien-
tists on an exchange basis. Copies may also be purchased at a nominal price. Inquiries
should be directed to Virginia D. Miller, Los Angeles County Museum of Natural
History, 900 Exposition Boulevard, Los Angeles, California 90007.
Robert J. Lavenberg
Managing Editor
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. Al-
though priority will be given to manuscripts by staff members, or to papers dealing
largely with specimens in the Museum’s collections, other technical papers will be
considered. All manuscripts must be recommended for consideration by the curator
in charge of the proper section or by the editorial board. Manuscripts must conform
to those specifications listed below and will be examined for suitability by an Edi-
torial Committee. They may also be subject to review by competent specialists out-
side the Museum.
Authors proposing new taxa in a CONTRIBUTIONS IN SCIENCE must
indicate that the primary type has become the property of a scientific institution of
their choice and cited by name.
MANUSCRIPT FORM. — ( 1 ) The 1964 AIBS Style Manual for Biological
Journals is to be followed in preparation of copy. (2) Double space entire manu-
script. (3 ) Footnotes should be avoided if possible. Acknowledgments as footnotes
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A GENERIC REVISION AND CATALOG OF THE WESTERN
HEMISPHERE GLENURINI WITH THE DESCRIPTION OF A
NEW GENUS AND SPECIES FROM BRAZIL
( NEUROPTER A : M YRMELEONTID AE )
By L. A. Stance^
Abstract: Studies of the type species of described glenurine
genera have led to considerable new synonymy as follows: Elach-
yleon Esben-Petersen 1927 {—Sericoleon Esben-Petersen 1932);
Eremoleon Banks 1901 (-^zlncamoleon Banks 1913, —Sosa Navas
1914, =Cortesiiis Navas 1914, =Dobla Navas 1926, =Joergenia
Esben-Petersen 1932, =Antilloleon Banks 1943); Psammoleon
Banks 1899 {=Diaziis Navas 1914). Pachyleon alvarengai, new
genus and new species, is described from Mato Grosso, Brazil.
Eight genera are now recognized and a key is given for their iden-
tification. Each genus is characterized and briefly discussed, in-
cluding new synonyms. A complete bibliography and catalog is
provided for the Western Hemisphere Glenurini. New informa-
tion contained in this catalog includes the following new species
synonyms: Psammoleon caiitiis (Walker) 1853 {—Feinenis neb-
ulosus Navas 1922) ; Psammoleon serrei (Navas) 1920 (=For-
micaleo chaperi Navas 1922); Gleniinis peculiaris (Walker)
1859 {—Glenunis brasiliensis Navas 1920); Eremoleon macer
(Hagen) 1861 (=Hesperoleon atomariiis Navas 1933). Also a
number of new combinations are indicated. The tribe Dimarellini
Markl 1954 is now considered a synonym of the Glenurini Banks
1928. The tribe Glenurini is now referred to the subfamily Myr-
meleontinae.
Introduction
The tribe Glenurini is one of the largest tribes of antiions in the world in
numbers of described genera. Markl (1954:246) lists 30 genera of Glenurini
from all major zoogeographical regions of which sixteen are restricted in dis-
tribution to the Western Hemisphere. However, no modern comprehensive
generic study has ever been made of the Glenurini. Furthermore the definition
of the tribe is probably deficient judging from the paucity of taxonomic char-
acters given by Markl (1954, p. 245) and earlier authors. The present account
deals only with the glenurine genera of the Western Hemisphere. An attempt
is made to signal taxonomic characters of importance and to point out the
variation that exists in others that have been used in the classification of the
tribe. Terminology is adopted from Stange ( 1970) .
^Department of Zoology, Instituto Miguel Lillo, San Miguel de Tucuman, Argentina.
1
2
Contributions in Science
No. 186
Acknowledgments
Study of nearly all of the types of the type species of glenurine genera by
me was made possible only by the cooperation of numerous individuals and
organizations. I am especially indebted to the Society of Sigma Xi for a Grant-
in-Aid of research which enabled me to study type material in Europe during
the summer of 1964. I would also like to thank Dr. S. L. Tuxen, Copenhagen
University, and Dr. Max Beier, Vienna Museum, for the loan of type material.
The following list of collections mentioned in the present study also includes
the names of persons who facilitated my studies: AMNH, American Museum
of Natural History, Dr. Jerome Rosen; Barcelona, Museo de Zoologia, Barce-
lona, Dr. Francisco Espahol Coll.; Berlin, Institut fur spezielle Zoologie und
zoologisches Museum, East Berlin, Dr. E. Konigsmann; BM(NH), British
Museum (Natural History), London, Mr. D. E. Kimmins; CAS, California
Academy of Sciences, Dr. Edward Ross; Copenhagen, Universitets Zoologiske
Museum, Copenhagen, Dr. S. L. Tuxen; Greifswald, Zoologisches Institut und
Museum, Ernst-Moritz-Arndt-Universitat, Greifswald. Dr. Ilselotte Groth.
Halle, Zoologisches Institut, Martin-Luther-Universitat, Halle-Saale, Dr. J. O.
Hiising; Havana, Institute de Biologia, La Habana; LACM, Los Angeles
County Museum of Natural History, Dr. Charles Hogue; La Plata, Museo de
La Plata, Facultad de Ciencias Naturales, La Plata, Prof. Luis De Santis;
MACN, Museo Argentine de Ciencias Naturales, Buenos Aires, Dr. Manuel
Viana; MCZ, Museum of Comparative Zoology, Harvard University, Dr.
Howard Evans; Paris, Museum National d’Histoire Naturelle, Paris, Miss S.
Kelner-Pillault; Hamburg, Zoologisches Staatsinstitut und Zoologisches Mu-
seum, Hamburg; Turin, Institute e Museo di Zoologia, Universita di Torino,
Turin; USNM, United States National Museum, Dr. Oliver Flint; Vienna,
Naturhistorisches Museum, Vienna, Dr. Max Beier; Zaragoza, Colegio del
Salvador, Zaragoza.
Systematic Position of the Glenurini
Banks (1943, p. 166) classified genera of the Glenurini in the subfamily
Macronemurinae. Markl (1954) published a comprehensive world classifica-
toin of the Myrmeleontidae but eliminated subfamilies and only dealt with
tribes. Stange (1968) placed the Glenurini in the subfamily Dendroleontinae.
My attempts to provide a satisfactory definition of the Dendroleontinae from
the Myrmeleontinae have now convinced me that this separation can not be
maintained. Therefore I consider the subfamilies Dendroleontinae and Macro-
nemurinae to be synonyms of the Myrmeleontinae. Essentially only two vena-
tional characters have been employed in the past for separating these subfami-
lies. One is the condition of vein 2A of the forewing. In some groups {Den-
droleon and Brachynemurus) vein 2A is evenly curved from its base to its
termination whereas it is strongly angled in Myrmeleon L, Macronemurus (an
old world genus), and in nearly all of the Glenurini. However the close struc-
1970
Catalog of the Western Hemisphere Glenurini
3
Figures 1-7: Pachyleon alvarengai. 1. Head of holotype. 2. Base of forewing. 3. Male
terminalia (right ectoproct removed). 4. Male genitalia (a) side view; (b) posterior
view. 5. Female terminalia, ventral view. 6. Foreleg of holotype, apex of femur, tibia
and tarsus. 7. Hindleg of holotype, apex of tibia and tarsus.
4
Contributions in Science
No. 186
tural resemblance between Dendroleon and some genera of the Glenurini (i.e.
Glenurus) argues against such an extreme separation. Furthermore vein 2A
shows some variation in the Glenurini. In a new genus to be described later in
this paper, vein 2A is closely associated with 3A and not with lA as is typical
in the tribe. Also in an undescribed species (Mystroleon praedator Sensu
Banks) vein 2 A is intermediate in condition between typical Glenurini and
Dendroleontini. For these reasons I consider this character insufficient for
separating subfamilies. The second venational character that has figured prom-
inently in subfamily classifications is the number of hindwing presectoral cross
veins. In the Myrmeleontinae there are four or more such crossveins whereas
the typical number in the Dendroleontinae is one. However, within the tribe
Brachynemurini, which I consider closely related to the Glenurini, there are
sometimes several crossveins depending upon the point of origin of the radial
sector. This indicates that the number of presectoral crossveins is of limited
value as a subfamily character.
Markl (1954) recognizes numerous tribes that are referrable to the Myr-
meleontinae. Probably many of these are invalid. Markl defines the Glenurini
almost entirely on wing venational characters some of which I have found to
be variable. Markl emphasizes the forking of forewing veins 2A and 3 A. In the
Glenurini, Nyutini (restricted to Africa) and Dimarellini (restricted to the
Neotropical Area), vein 2A is forked and 3 A is simple. Other related tribes,
all restricted to the Eastern Hemisphere, have vein 2A simple and 3A forked.
This condition is given for the Formicaleontini, Creoleontini, Obini, Nemo-
leontini and Protoplectrini. However this character is variable, even at the
specific level in some instances. In the glenurine genus Eremoleon, most spe-
cies fit the concept of Markl (vein 2 A forked, 3 A simple in insipidus, longior,
etc.) but other species show the opposite condition (as in some nigribasis) .
Therefore this character is of limited taxonomic importance. Another vena-
tional character that Markl uses in differentiating the Glenurini (from the
Dimarellini and Nyutini) is the presence of the anterior banksian line. This
prominent longitudinal line is found in either wing and is formed by the bend-
ing of the branches of the radial sector. It is highly variable in its development
in the genus Brachynemurus, a member of the related tribe Brachynemurini.
In the tribe Glenurini, nearly all of the Western Hemisphere species that I
have examined show no development of the banksian lines. It is apparent that
Markl was in error employing this character in the diagnosis of the Glenurini
(at least the Western Hemisphere component).
Markl (1954, p. 247) defined the tribe Dimarellini on venational char-
acters which Stange (1963, p. 810) criticized. Stange offered the character of
the pretarsal claws as a possible means of separating the Dimarellini from the
Glenurini. With this new criterion the genera Elachyleon and Navasoleon were
removed from the Glenurini and placed with Dimarella in the Dimarellini. In
these three genera the pretarsal claws are capable of closing against the distal
1970
Catalog of the Western Hemisphere Glenurini
5
tarsomere whereas the claws are ordinary in other genera. However it is now
apparent that this specialized condition of the pretarsal claws have evolved
several times in unrelated groups of Myrmeleontidae as evidenced by the
occurrence of this character in the Dendroleontini {Tricholeon Esben-Petersen
from Africa; Froggattisca Esben-Petersen from Australia) and in the Glenu-
rini {Megistopus Rambur from Europe). The genus Elachyleon, possessor of
the specialized pretarsal claws, agrees with the glenurine genus Eremoleon in
all generic level taxonomic characters except the condition of the pretarsal
claws. Thus, I now consider the pretarsal claw character to be a generic charac-
ter. In search of a possible new means of maintaining the tribe Dimarellini I
have reconsidered all the specialized features of the genus Dimarella. The mem-
bers of this genus differ from other Glenurini in the following characters: (1)
wing venation (CuP + lA run parallel with the posterior fork of forewing
CuA) ; (2) antennal fossa widely separated from eye; (3) male ectoproct with
elongate postventral lobe; (4) form of the male gonarcus (large lateral expan-
sions) and shape of the parameres (not in the form of flat plates). A new
genus to be described later in this paper also shares these characters (but has
ordinary pretarsal claws). However recent study of the poorly known species
Psammoleon-cautus has revealed that this species is intermediate between the
genus Dimarella and the glenurine genus Psammoleon. This species appears
to be a rather ordinary Psammoleon in regards to the wing venation, head
structure and legs but possesses the elongate postventral lobe of the male ecto-
proct as in Dimarella as well as comparable development of the male gonarcus
and parameres.
From the foregoing discussion it is apparent that we are confronted with
a weak tribal classification of the Myrmeleontinae. More study is needed of
the Old World genera of the Glenurini as well as of the related Old World
tribes Formicaleontini, Creoleontini, Obini, Nemoleontini, Protoplectrini and
Nyutini before a satisfactory world-based classification can be given. The new
world tribe Dimarellini can not be maintained and I now consider Dimarella
as a rather specialized genus of the Glenurini.
In the Western Hemisphere there is less diversity in the Myrmeleontinae,
at least from the point of view of described tribes. The following key will dis-
tinguish the Glenurini from other Western Hemisphere groups of antiions.
1. Sensory pit of the distal labial palpomere slitlike; hindleg with femoral sense
hair or femoral sense hair absent Palparinae and Acanthaclisinae
Sensory pit of the distal labial palpomere oval-shaped; femoral sense hair
present on foreleg and midleg (except few species of Brachynemurus) but
not on hindleg ’ (Myrmeleontinae) 2
2. Forewing vein 2A evenly curved from base to hind margin, widely separated
from 3A at least at basal one-half Dendroleontini and Brachynemurini
Forewing vein 2A runs close to vein CuP + 1 A for a short distance, then
6
Contributions in Science
No. 186
abruptly angles toward hind margin or veins 2A and 3A closely associated....
3
3. Radial sector of hindwing arises beyond cubital fork, four or more presec-
toral crossveins present; pilula axillaris present Myrmeleontini and
Porrerini
Radial sector of hindwing arises well before cubital fork, one (rarely two or
three) presectoral crossveins present; pilula axillaris absent Glenurini
Western Hemisphere Genera of the Glenurini
All of the glenurine genera known from the Western Hemisphere are
restricted to that area. It is possible that this reflects the regionalism of former
workers on this group since no modern world study has been made. Even the
Nearctic and Neotropical faunas of the Glenurini have not been treated to-
gether. Banks provided keys to the Nearctic genera (1928) and the Neotropical
genera (1943). The Nearctic fauna is relatively small. The three genera repre-
sented in this region are also found in both the North American and South
American Neotropical areas. All of the new world glenurine genera are found
in South America and three of these are restricted to that continent. Twenty-
four generic names have been proposed for the Western Hemisphere Glenu-
rini. Some of these have been placed in synonymy. My study of nearly all of
the type species of Western Hemisphere genera of the Glenurini has led to the
discovery of further generic synonymy. In the following account of the glen-
urine genera this synonymy will be discussed.
Araucaleon Banks
Araucaleon Banks, 1938. Proc. Ent. Soc. Wash. 40:127.
Type-species: Araucaleon inca Banks, by original designation.
Description: Antennal fossa separated from ocular rim by less than great-
est diameter of pedicel; antenna long and slender; pronotum longer than wide;
mesoscutellum smaller than metanotum; all legs about equal in length; tibial
spurs longer than forecoxal length; pretarsal claws not capable of closing
against distal tarsomere; forewing with anterior margin abruptly angled and
swollen near coalescing of subcostal and radial veins, costal area at this point
nearly twice as high as at middle of wing; forewing radial sector originates well
basad to forking of CuA; posterior fork of forewing vein CuA at an oblique
angle to hind margin; forewing vein 2 A widely separated from normal 3 A be-
fore strong angle; hindwing vein CuA ends well before midpoint to forking
of MP9; male ectoproct simple, without postventral lobe; male paramere in
form of rigid plate; male gonarcus not greatly expanded laterally.
Discussion: This genus is rare in collections. I have seen only two males
of the type-species. In addition, Glenurus withycombei Esben-Petersen, de-
scribed from Trinidad, belongs to Araucaleon. This species is known only from
the holotype female. This genus appears to be allied with Eremoleon Banks
1970
Catalog of the Western Hemisphere Glenurini
7
but differs from that genus and other known glenurine genera by the swelling
of the forewing costal margin near the point of coalescing of the subcostal and
radial veins and by the termination of vein CuA of the hindwing well before
the midpoint between the wing base and forking of MP2.
Dimarella Banks
Dimarella Banks, 1913. Trans. Amer. Ent. Soc. 39:229.
Type-species: Eremoleon angiistus Banks, by original designation.
Description: Antennal fossa separated from ocular rim by more than
greatest diameter of pedicel; antenna variable in length; pronotum wider than
long; mesoscutellum as long as metanotum; legs different in lengths, often
midleg shortest; tibial spurs much shorter than forecoxal length; pretarsal claws
capable of closing against distal tarsomere; forewing with anterior margin
evenly curved toward apex, costal area at point of coalescing of subcostal and
radial veins usually lower than at middle of wing; forewing radial sector orig-
inates well beyond forking of CuA; forewing vein CuP + 1 A runs parallel with
posterior branch of CuA for a long distance; forewing vein 2A usually widely
separated from normal 3A before strong angle toward posterior margin; hind-
wing vein CuA ends near forking of MP._,; male ectoproct with elongate post-
ventral lobe; male paramere with hook; male gonarcus expanded laterally as
a round plate; female ectoproct with strong digging setae ventrally, shorter
than on lateral gonapophyses.
Discussion: Dimarella is limited to the Neotropical Region. Synonymical
notes on the genus were given by Stange (1963; 1968). The Mexican species
were revised by Stange (1963). This genus stands apart from other described
genera by the prominent postventral lobe of the male ectoproct, the parallel
course of veins CuP + lA and CuA of the forewing and the wide space be-
tween the ocular rim and antennal fossa. All of these features are shared with
a new genus to be described later in this paper. The specialized pretarsal claws
is a morphological trait shared only with Navasoleon and Elachyleon.
Elachyleon Esben-Petersen
Elachyleon Esben-Petersen, 1927. Ann. Mag. nat. Hist. (Ser. 9) 20:348. Type-
species: Elachyleon punctipennis Esben-Petersen, by original designation.
Sericoleon Esben-Petersen, 1932. Vidensk. Medd. Dansk. naturh. Foren
94:114. Type-species: Sericoleon paessleri Esben-Petersen, by original
designation. New Synonymy.
Description: Antennal fossa separated from ocular rim by less than great-
est diameter of pedicel; antenna long and slender; mesoscutellum shorter than
metanotum; all legs about equal in length; tibial spurs shorter than fore-
coxal length; pretarsal claws capable of closing against distal tarsomere;
forewing with anterior margin evenly curved toward apex, costal area at point
8
Contributions in Science
No. 186
of coalescing of subcostal and radial veins either somewhat lower or higher
than at middle of wing; forewing radial sector originates only somewhat before
forking of CuA or well beyond; posterior fork of forewing vein CuA at an
oblique angle to hind margin; forewing vein 2 A widely separated from normal
3 A before strong angle toward posterior margin; hindwing vein CuA extends
nearly to forking of MP2; male ectoproct simple, without postventral lobe; male
paramere in form of rigid plate; male gonarcus not greatly expanded laterally;
female ectoproct usually with weak digging setae; posterior gonapophysis
longer than wide, digitiform.
Discussion: I have studied one of the paratypes of the type-species of
Sericoleon, S. paessleri Esben-Petersen, described from Chile. This species
possesses all of the generic characters of Elachyleon and therefore I am placing
Sericoleon in synonymy with Elachyleon. Elachyleon can be distinguished
from all other Western Hemisphere Myrmeleontidae by the specialized pre-
tarsal claws, except for Navasoleon (which lacks tibial spurs) and Dimarella
(which has forewing veins CuP + 1 A and CuA parallel) . This genus agrees
closely with Eremoleon in both morphological and biological characters. I have
reared an undescribed species from Mexico and from Argentina. Both species
are found in the silt floors of caves. This is the same habitat for various species
of Eremoleon. Elachyleon is now constituted of four described species. The
most widespread species, E. punctipennis (ranges from Mexico to Argentina)
has been illustrated in part by Stange ( 1963) . Additional generic synonymy
is given by Stange ( 1 968 ) .
Eremoleon Banks
Eremoleon Banks, 1901. Trans. Amer. Ent. Soc. 27:366. Type-species: Myr-
meleon macer Hagen, by original designation.
Incamoleon Banks, 1913. Trans. Amer. Ent. Soc. 39:229. Type-species:
Psammoleon punctipennis Banks, by original designation. New Syn-
onymy.
Sosa Navas, 1914. Broteria 12:218. Type-species: Sosa conspicuus Navas, by
original designation. New Synonymy.
Cortesius Navas, 1924. Broteria 21 : 107. Type-species: Cortesius genini
Navas, by original designation. New Synonymy.
Dobla Navas, 1926. Deutsch. ent. Zeitschr. :428. Type-species: Dobla arcuata
Navas, by original designation. New Synonymy.
Joergenia Esben-Petersen, 1932. Vidensk. Medd. Dansk. naturh. Foren
94:118. Type-species: Joergenia pulchra Esben-Petersen, by original des-
ignation. New Synonymy.
Antilloleon Banks, 1 943. Bol. ent. venezolana 2: 168. Type-species: Glenurus
cerverai Navas, by original designation. New Synonymy.
Description: Antennal fossa separated from ocular rim by less than great-
1970
Catalog of the Western Hemisphere Glenurini
9
est diameter of pedicel; antenna long and slender; pronotum variable, usually
longer than wide; mesoscutellum shorter than metanotum; legs about equal in
length except usually hindlegs longer; tibial spurs variable in length, usually
shorter than forecoxal length; pretarsal claws not capable of closing against
distal tarsomere; forewing with anterior margin evenly curved toward apex,
costal area at point of coalesing of subcostal and radial veins usually lower
than at middle of wing; forewing radial sector originates somewhat before
forking of CuA or well beyond; posterior fork of forewing vein CuA at an
oblique angle to hind margin; forewing vein 2A widely separated from normal
3A before strong angle toward posterior margin; hindwing vein CuA extends
nearly to forking of MP2 or somewhat beyond; male ectoproct simple without
postventral lobe; male paramere in form of rigid plate; male gonarcus not
greatly expanded laterally; female ectoproct usually with weak digging setae;
posterior gonapophysis longer than wide, digitiform.
Discussion: There is considerable new generic synonymy in addition to
that reported by Navas (1916) Banks (1928) & Adams (1957, 1958). These
new synonyms will be discussed in chronological order.
Incamoleon Banks 1913 — The type-species of this genus, punctipennis
Banks, was originally described in the genus Psammoleon. Banks separated
this species in its own genus apparently based on the longer, more slender legs
of punctipennis. Also the wing shape is more slender than in Psammoleon. All
of these features are found in the genus Eremoleon. Banks (1943:166) dis-
tinguishes Incamoleon from Eremoleon {—Glenopsis Banks) by the relative
length of the tarsomeres. In Incamoleon the distal tarsomere is about equal to
the other tarsomeres together whereas in Eremoleon the distal tarsomere is
shorter. However my studies on Incamoleon puftctipennis have revealed that
this tarsomere proportion character is not constant and although useful for
identification it is not important enough for generic distinction. Further studies
have revealed two characters found in Incamoleon that are unknown in other
Eremoleon. First, there is marked sexual dimorphism. The female wings usually
have conspicuous dark spots, whereas the wings are not spotted (or only
weakly so) in the males. Also the female genitalia appear somewhat distinct
in that the posterior gonapophysis is more inflated than in other members of
the genus Eremoleon. However I am of the opinion that these two characters
are not sufficient to maintain Incamoleon as a distinct genus so that I am now
referring punctipennis to the genus Eremoleon.
Sosa Navas 1914 — I studied the holotype female of the type-species, Sosa
conspicuus, in 1962 from the collections of the Vienna Museum. Navas gave
few notes in his original description as to the basis of this new genus but did
point out that it was related to Eormicaleon (an old world genus) but with
longer and more slender legs and that the genus might belong to the Dendro-
leontini. However my studies of the holotype revealed no important differences
from the current generic concept of Eremoleon Banks. Sosa conspicuus re-
10
Contributions in Science
No. 186
sembles Eremoleon macer (Hagen) except that the wings are more slender.
This species is now placed in Eremoleon.
Cortesius Navas 1924 — I studied the two syntypes of the type-species,
Cortesius genini Navas, in the Paris Museum in 1964. Navas (1924:107) did
not give much information in his original description as to the relationships
of this genus except to point to a similarity with the genus Glenurus Hagen.
My studies revealed that Cortesius genini agrees with all generic characters
of Eremoleon. The only unusual feature of this species is that the forewing
costal area is biareolate to triareolate. Interconnected crossveins in the costal
area are found otherwise only in the genus Psammoleon among the Glenurini.
Dobla Navas 1926 — The holotype female of the type-species, Dobla ar-
cuata Navas, is located in the Paris Museum. My studies of the holotype indi-
cate that it is a synonym of Cortesius genini Navas. Therefore the genus Dobla
is a synonym of Eremoleon Banks.
Joergenia Esben-Petersen 1932 — I have just recently received for study
the holotype female of the type-species, Joergenia pulchra Esben-Petersen, on
loan from the Copenhagen Museum. Although this species is quite distinctive
by virtue of the unusual narrowing of the wings, especially of the costal area,
I have not found characters that would warrant generic separation from Ere-
moleon. There is considerable range in the relative broadening of the costal
area in Eremoleon and I am of the opinion that J. pulchra represents one ex-
treme. The other extreme, broadening of the costal area, is exemplified by
E. genini (Navas) and E. anomalus (Rabur), whereas intermediate conditions
are found in E. conspicuus (Navas), E. gracile Adams and many others. I have
also seen an undescribed species of Eremoleon from Venezuela which has the
costal area nearly as narrow as in /. pulchra.
Antilloleon Banks 1943 — I have studied the type material of the type-
species, Glenurus cerverai Navas, in the collections of the Museum of Com-
parative Zoology, Harvard University. Banks (1943:167) pointed out the
unusual venational feature present in the Caribbean species. The radial sector
originates only a little beyond the level of forking of CuA in the forewing. In
most other species of the Glenurini the radial sector originates well beyond
the forking of CuA. However in the genus Araucaleon Banks, the radial sector
originates well before the forking of CuA, a peculiarity in the Glenurini. In
other respects A. cerverai Navas agrees well with species of the genus Eremo-
leon. Considerable variation is present in the genus Eremoleon in the relative
position of the origin of the radial sector and forking of CuA. I am, therefore,
of the opinion that Glenurus cerverai belongs to the genus Eremoleon, Antil-
loleon thus falling as a synonym of Eremoleon.
Revisionary studies are needed of the genus Eremoleon which appears to
be a complex taxon. Adams (1957) presented a key to most of the Nearctic
species. There is no one key character of the genus which permits easy recog-
nition, although the genus differs considerably from other glenurine genera
1970
Catalog of the Western Hemisphere Glenurini
11
except Elachyleon Esben-Petersen, which has specialized pretarsal claws. I have
reared four species of Eremoleon (from Arizona, Mexico and Argentina) and
all exhibit similar biological characteristics. The larvae live in the silt of caves
or rock crevices and seem to be good climbers.
Glenurus Hagen
Glenurus UdigQn 1866. Stettin, ent. Ztg. 'll '312.
Type-species: Formicaleo grata Say.
Descripti'on: Antenna fossa separated from ocular rim by less than great-
est diameter of pedicel; antenna long and slender; mesoscutellum shorter than
metanotum; all legs about equal in length except hindlegs usually somewhat
longer; tibial spurs longer than forecoxal length; pretarsal claws not capable
of closing against distal tarsomere; forewing with anterior margin evenly
curved toward apex, costal area at point of coalescing of subcostal and radial
veins somewhat lower or higher than at middle of wing; forewing radial sector
originates well beyond forking of CuA; posterior fork of forewing vein CuA
at an oblique angle to hind margin; forewing vein 2 A widely separated from
normal 3 A before strong angle toward posterior margin; hindwing vein CuA
extends nearly to or somewhat beyond forking of MP2; male ectoproct simple
without postventral lobe; male paramere in form of rigid plate; male gonarcus
not greatly expanded laterally; female ectoproct with prominent digging setae;
posterior gonapophysis weakly produced as a swelling.
Discussion: This is a very homogeneous genus easily recognized by the
considerable brown suffusion of the wing apices. G. heteropteryx is the only
known species that lacks this prominent suffusion in the forewing, although the
hindwing conforms with the rest of the genus. Structurally Glenurus seems
close to Eremoleon but differs from that genus as well as all other glenurine
genera in having the posterior gonapophysis poorly developed. Banks (1922)
has provided a key to the South American species and (1928) to the Nearctic
species.
Navasoleon Banks
Navasoleon Banks 1943. Bol. ent. Venez. 2:168. Type-species: Gymnocnemia
boliviana Banks, by original designation.
Description: Antenna fossa separated from ocular rim by less than great-
est diameter of pedicel; antenna long and slender; mesoscutellum shorter than
metanotum; foreleg (at least females) much longer than midlegs and hindlegs;
tibial spurs absent; pretarsal claws capable of closing against distal tarsomere;
forewing with anterior margin evenly curved toward apex, costal area at point
of coalescing of subcostal and radial veins lower than at middle of wings; fore-
wing radial sector originates well beyond forking of CuA; posterior fork of
forewing vein CuA at an oblique angle to hind margin; forewing vein 2 A
widely separated from normal 3 A before strong angle toward posterior margin;
12
Contributions in Science
No. 186
hindwing vein CuA extends somewhat beyond forking of MP2; female ecto-
proct with small digging setae; posterior gonapophysis swollen thumb-like.
Discussion: Navasoleon is a poorly known genus and male specimens
are unavailable for study. The genus appears very distinct. The lack of tibial
spurs is diagnostic and the specialized pretarsal claws is found only in Elachy-
leon and Dimarella. In the type-species of the genus and also in an undescribed
species the foreleg is unusually long. The species known from Argentina has
shorter front legs but the specimen is damaged to such an extent that the sex
is undeterminable.
Pachyleon Stange, new genus
Diagnosis: This genus is distinguished from other glenurine genera in
the Western Hemisphere except Dimarella by the following characters: (1)
forewing vein 2A is closely associated with vein 3A for its entire course (see
Fig. 2) ; (2) radial and subcostal veins in both wings crowded together; (3)
antennal fossa separated by more than greatest diameter of scape from ocular
rim; (4) forewing veins CuP + lA and posterior branch of CuA run parallel
for a long distance. Pachyleon is closely allied to Dimarella but differs from
that genus in two important characters: (1) pretarsal claws simple in Pachy-
leon, specialized in Dimarella (capable of closing against tarsomere); (2)
hindwing broadly triangular in Pachyleon (greatest breadth about equal to
forewing) whereas it is more slender in Dimarella (greatest breadth of hind-
wing much less than that of forewing). The structural gap between Pachyleon
and Dimarella would be even greater if it were not for a recently discovered
(apparently undescribed) species of Dimarella which has the condition of fore-
wing vein 2A (closely associated with 3A) very similar to that in Pachyleon.
Pachyleon and Dimarella share so many characters that are not found in
almost all other Glenurini that they form a group apart. However Psammoleon
cautus (Walker) represents an intermediate type between Psammoleon and
these two genera. The unusual shape of the hindwing of Pachyleon provides a
key character for the genus. It remains to be seen whether or not the broad
hindwing is of generic or specific importance since considerable variation
exists in the relative broadening of the hindwing in the allied genus Dimarella.
Description: Antennal fossa separated from ocular rim by more than
greatest width of antennal scape; antenna short and thick; pronotum much
broader than long; thorax depressed, mesoscutellum as long as metanotum;
foreleg moderately short and thickened, midleg greatly swollen and short,
hindleg relatively long, about twice as long as foreleg; tarsomeres of foreleg
and midleg about equal, those of hindleg much longer; tibial spurs present,
much shorter than forecoxal length; pretarsal claws not capable of closing
against distal tarsomere; femoral sense hair of foreleg short, much less than
one-half length of femur; banksian lines absent; subcostal and radial veins
crowded together, nearly touching well before their apical fusion; forewing
1970
Catalog of the Western Hemisphere Glenurini
13
Figure 8. Photo of holotype (male terminalia removed).
vein CuP + 1 A runs parallel with posterior branch of CuA for a long distance;
forewing vein 2A widely separated from lA, closely associated and almost
fused with vein 3A, not greatly angled; male ectoproct with elongate postven-
tral lobe; male genitalia not rigid, paramere with hook; gonarcus expanded
laterally as a large round plate; female ectoproct simple with strong digging
setae ventrally; lateral gonapophyses separated, with longer digging setae than
on ectoproct; posterior gonapophysis digitiform, longer than wide.
The generic name is from the Greek words Pachys meaning wide and
leon meaning lion, in reference to the broad wings, pronotum and head. The
following new species is the type-species of the genus.
Pachyleon alvarengai Stange, new species
Diagnosis: As the only species in the genus, the generic characters serve
to distinguish this species. The unusually short and broadened wings, espe-
cially the rather triangular hindwing (see Fig. 8) is distinctive among all
known species of ant-lions in the Western Hemisphere. This unusual hindwing
shape is also unknown to me in Old World species but I do not have a perfect
knowledge of the Old World fauna. The most similar species is a Dimarella
(probably undescribed) in which the wings are similar in shape although not
as extreme as in P. alvarengai and in which vein 2A is closely associated with
vein 3A.
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Contributions in Science
No. 186
Description (Holotype male): Length of body about 19 mm, forewing
15 mm, hindwing 1 1 mm, greatest wing width (about equal in forewing and
hindwing) 4.75 mm.
Color: Face (Fig. 1) pale yellowish with A-shaped dark brown mark at
middle below antennae and crescent-shaped dark brown mark above each
antenna, sublateral spot on each side of clypeus, larger dark brown area
laterad of antenna toward eye, extending ventrally and dorsally nearly to first
row of vertex scars; vertex with grayish bloom, scar pattern about as in Fig. 8;
antenna with scape and pedicel pale yellow with dark brown behind, flagello-
meres dark brown basally and much paler distally; maxilla and labius nearly
all pale yellow except distal palpomere and cardo with dark brown; postgena
mostly dark brown; pronotum mostly dark brown with grayish bloom inter-
rupted by irregular paler areas; notum II and III dark brown with grayish
bloom, darker velvety areas as follows: submedial triangular area on each side
of prescutum II; dark brown oval spot each side of middle and touching, sur-
rounded by golden colored microtrichia on prescutum III; large curved dark
brown mark sublaterally near anterior margin of scutum; postscutellum II and
III nearly all black; cervical sclerites pale yellow; pleuron and sternum mostly
dark brown with grayish bloom, paler brown areas mostly in front of midcoxa
and hindcoxa, mesopleural wing process yellowish; forecoxa mostly pale
brown, other coxae darker brown; forefemur pale brown with numerous dark
brown spots, spots becoming confluent on most of exterior face and on distal
one-third of closing face; foretibia pale brown with dark brown spots and small
dark brown stripes at basal one-third of exterior face, larger dark brown areas
at middle, mostly complete but smaller apical rings, dark brown areas inter-
connected along closing face; midtibia and midfemur with large, mostly
separated dark brown spots on exterior face, somewhat denser and forming
short transverse dark brown bands on closing face of tibia, subapical inter-
rupted dark brown ring on femur, solid dark brown one on tibia; hindfemur
pale brown with dark brown spots at setal bases, infuscated between spots
especially on exterior face, large subapical dark brown band; hindtibia pale
brown with large, mostly separated dark brown spots at setal bases on closing
face, small dark brown stripe near base, spots becoming more stripelike dis-
tally, exterior face nearly all pale brown except dark brown apical ring and
smaller dark brown area toward base; tarsi of all legs similarly patterned, pale
brown with apical dark brown ring on tarsomeres; wings nearly without suffu-
sion, pale white stigmal area preceded by small darkish brown area in fore-
wing; wing veins mostly pale brown, especially hindwing, with dark brown
streaks along longitudinal veins principally at crossvein junctions, many cross-
veins dark brown especially apically in forewing; abdomen with tergites mostly
pale brown with complicated pattern formed by dark brown areas (see Fig. 8),
sternites mostly dark brown with a prominent although narrow median longi-
tudinal pale brown line.
1970
Catalog of the Western Hemisphere Glenurini
15
Chaetotaxy: Head with few prominent setae except on most of clypeus,
white ones on dark brown areas of frons, longer ones on labrum and many
long white setae on mentum and stipes; vertex with few inconspicuous ap-
pressed dark setae; pale with very sparse, short dark brown setae; pronotum
with some short setae on disc, longer dark brown ones at lateral and posterior
margin; row of five long white setae on each side of middle along anterior
margin of scutum II, several smaller dark brown setae on prescutum II, else-
where nearly absent or very short; pleuron and coxa with rather conspicuous
long white hairlike setae; forefemur with dense, mostly appressed white setae
on exterior surface, some black and some white erect setae distally, closing
surface with fewer setae and with more black ones; foretibia with subbasal
row of three black bristles, followed by another similar row, then one of two
bristles and finally a subapical black bristle on exterior face, interspersed with
shorter, mostly dark setae, closing surface with many stout setae, predomi-
nately white; midfemur with row of white bristles on exterior surface, many
other setae elsewhere but more hairlike and white; midtibia with mostly black
setae of various sizes, larger black ones toward apex of lateral face, larger
white ones near base of median face; hindfemur with many short, mostly
appressed white setae; hindtibia with many setae along closing surface, white
ones mostly concentrated in middle, exterior surface nearly without setae, only
a few scattered minute ones; femoral sense hair of foreleg and midleg shorter
than width of femur at that point; tarsi with rather dense, dark brown setae on
closing surface flanked (typically) on the four basal tarsomeres by one white
seta near apex, exterior surface with fewer setae, especially hindtarsus; ab-
domen with only microsetae on tergites I-V; starting with tergite VI, dark
brown setae mostly laterally increasing in abundance toward terminalia; ster-
nites with numerous, mostly white hairlike setae; postventral lobe with rather
prominent black bristles toward apex (Fig. 3).
Structure-. Head with vertex not much raised above eyes; greatest ocular
width about one-third interocular distance (measured just below antennal
fossae); antenna short and rather flattened with 19 flagellomeres, flagellomere
I longer than wide, II about 1.5 times wider than long (in flat view) with
increasing width toward apex; distal palpomere of labius rather slender, not
much swollen, sensory pit oval and close to base; pronotum about twice as wide
as long; femur of all legs broadened beyond base, tibia much more slender
except midtibia, which is subequal to midfemur; midleg much shorter than
either foreleg or hindleg; wing venation and shape as in Fig. 8; abdomen
shorter than forewing, segments larger basally, decreasing in diameter begin-
ning with segment IV; tergites I-V and sternite II with numerous scalelike
spicules; terminalia as in Fig. 3, genitalia as in Figs. 3 and 4.
Types: Male holotype and 1 female paratype from Jacare (N. Xingu),
Mato Grosso, Brazil, XI 1961, M. Alvarenga collector 52° 24' W; 12° 00' S.
16
Contributions in Science
No. 186
Holotype deposted in the collections of the Los Angeles County Museum of
Natural History. Paratype female retained in the author’s collection.
Variation: The female paratype agrees well in color, chaetotaxy and
structure with the male except that the scalelike spicules on the abdomen are
lacking. Female terminalia as in Fig. 5.
Discussion : This species is dedicated to Moacir Alvarenga, Brazilian
coleopterist, for his efforts in securing critical ant-lion material from Brazil.
The holotype male upon arrival at the depository was found damaged in
transit. Most of the broken parts (pieces of wings, legs and antenna) are
associated with the specimen and no difficulty in identification is foreseen
although this was an unfortunate mishap. The photo (Fig. 8) shows the
holotype before shipment.
Psammoleon Banks
Psammoleon Banks 1899. Canad. Ent. 31 :69. Type-species: Myrmeleon in-
geniosus Walker, by original designation.
Diazus Navas 1914. Broteria 12:220. Type-species: Diazus clavatus Navas,
by original designation.
Description: Antennal fossa separated from ocular rim by much less
than greatest diameter of scape; antenna variable; pronotum wider than long;
mesoscutellum smaller than metanotum; legs about equal in length, hindleg
sometimes longer than foreleg; tibial spurs variable, usually shorter than fore-
coxal length; pretarsal claws not capable of closing against distal tarsomere;
forewing with anterior margin evenly curved toward apex, costal area at point
of coalescing of subcostal and radial veins lower than at middle of wing; fore-
wing radial sector originates well beyond forking of CuA; posterior fork of
forewing vein CuA at an oblique angle to hind margin; forewing vein 2 A
widely separated from normal 3A before strong angle toward posterior mar-
gin; hindwing vein CuA extends nearly to forking of M?2 or somewhat beyond;
male ectoproct simple without postventral lobe, except in P. cautus (Walker) ;
male paramere in form of rigid plate (except cautus) ; male gon arcus not
greatly expanded laterally (except cautus) ; female ectoproct without digging
setae (sometimes long bristles present) , often produced ventrally as a lobe;
posterior gonapophysis well developed, often somewhat inflated.
Discussion: I studied the holotype male of the type-species of Diazus
(D. clavatus Navas) in 1962 on loan from the Vienna Museum. The specimen
is in good condition except that the hindlegs and abdominal terminalia are
missing. My studies of this type indicate that D. clavatus is a typical Psammo-
leon, rather similar to Psammoleon minor Banks. Navas (1914:220) gave few
remarks concerning the diagnostic features of Diazus except to point to a
relationship with Incamoleon and Glenopsis. Probably the swollen femora of
D. clavatus prompted him to consider this species generically distinct. However
it is clear that Diazus is a synonym of Psammoleon.
1970
Catalog of the Western Hemisphere Glenurini
17
The genus Psammoleon is the largest genus in the Western Hemisphere
Glenurini, with twenty-seven described species. Also I have seen a number of
undescribed species, mainly from Mexico. Most species are found in North
America. The genus is unknown south of Mato Grosso, Brazil. The long fore-
femoral sense hair (over one-half length of femur) is diagnostic. Also the
rather swollen forefemur is characteristic except for Pachyleon and Dimarella.
P. cautus (Walker) is intermediate between Psammoleon and Dimarella and
perhaps should be placed in a distinct genus. However this situation suggests
a relationship between these genera. The modified female ectoproct in many
species of Psammoleon is also without known counterpart among Western
Hemisphere Glenurini. Considerable variation is present between species in
the development of the tarsus, forewing costal area and in chaetotaxy.
Key to Genera of Western Hemisphere Glenurini
1. Pretarsal claws capable of closing against distal tarsomere 2
Pretarsal claws not capable of closing against distal tarsomere 4
2. Posterior fork of forewing vein CuA and vein CuP + 1 A parallel with each
other and hind margin for a long distance; antennal fossa separated by more
than greatest width of antennal pedicel from ocular rim; male ectoproct
with elongate postventral lobe Dimarella
Posterior fork of forewing vein CuA at an oblique angle to hind margin;
antennal fossa separated by less than greatest width of antennal pedicel
from ocular rim; male ectoproct without elongate postventral lobe 3
3. Tibial spurs present; foreleg about equal or shorter than \\md\Qg..Elachyleon
Tibial spurs absent; foreleg (at least in female) usually longer than hindleg
Navasoleon
4. Antennal fossa separated by more than greatest width of antennal pedicel
from ocular rim; hindwing about twice as long as wide, much shorter than
forewing (Fig. 8); forewing vein 2A closely associated with 3 A, not much
angled (Fig. 2); subcostal and radial veins of both wings nearly touching,
especially before their apical fusion; male ectoproct with elongate post-
ventral lobe Pachyleon
Antennal fossa separated by less than greatest width of antennal pedicel
from ocular rim; hindwing over three times longer than wide, about equal
in length to forewing; forewing vein 2A widely separated from normal 3A
before strong angle toward hind margin; subcostal and radial veins widely
separated until their apical fusion; male ectoproct without postventral lobe
(except P. cautus) 5
5. Forewing with anterior margin abruptly angled and swollen near coalescing
of subcostal and radial veins, costal area at this point nearly twice as wide
as at middle of wing; hindwing vein CuA ends well before midpoint to fork-
ing of MP2; forewing radial sector originates well basad to forking of
CuA Araucaleon
18
Contributions in Science
No. 186
Forewing with anterior margin evenly curved toward apex, costal area at
point of coalescing of subcostal and radial veins usually somewhat narrower
than at middle of wing; hindwing vein CuA extends nearly to forking of
MPo or beyond; forewing radial sector originates near or well beyond
forking of CuA 6
6. Forefemur swollen, at widest point much wider than interantennal distance,
with abundant pubescence, especially on closing surface; femoral sense hair
or foreleg elongate, over one-half length of femur; female ectoproct often
with ventral projection Psammoleon
Forefemur slender and elongate, at widest point about equal at most to in-
terantennal distance, with sparser pubescence; femoral sense hair of foreleg
less than one-third length of femur; female ectoproct not produced ven-
trally 7
7. Apical one-fifth of hindwing and usually forewing with predominant dark
brown suffusion in marked contrast to basal one-fifth; posterior gonapophy-
sis of female weakly produced as a swelling Glenurus
Apical one-fifth of wings predominantly transparent, sometimes many
scattered dark brown spots present; posterior gonapophysis of female well
developed, digitiform Eremoleon
Synoptic Catalog of the Western Hemisphere Glenurini
WITH Type Information
Genus Araucaieon Banks
Araucaleon Banks 1938b: 127. Type-species: Araucaieon inca Banks, orig.
design.
inca Banks 1938b: 128. Hoi. 5, Huachi, Bolivia, IX.1925 (USNM).
withycombei (Esben-Petersen) 1927:347 (as Glenurus). Hoi. $, St. Augus-
tine, Trinidad, VI. 1924 (BM). New Comb.
Genus Dimarella Banks
Dimarella Banks 191 3a: 229. Type-species: Eremoleon angustus Banks, orig.
design.
=Nobra Navas 1915b:6. Type-species: N o bra mar tinsi as, ong. design.
=Furgus Navas 1921:51. Type-species: Nobra riparius Navas, orig. design.
=Mystroleon Banks 1924:436. Type-species: Myrmeleon praedator^ a\kev .
Taxonomy: Banks ( 1943: 166) ; Stange (1963:810-812; 1968:59).
angusta (Banks) 1908:31 (as Eremoleon). Hoi. 9, Santa Elena, Ecuador
(MCZ).
=Nobra nevermanni Navas 1936:167. Syntypes, Costa Rica, Farm Ham-
burg am Reventazon, V.2.1934 (Zaragoza).
Taxonomy: Banks (1913a:230; 1943 : 168) ; Stange (1963:812).
1970
Catalog of the Western Hemisphere Glenurini
19
Distribution: Neotropical (Costa Rica, Colombia, Ecuador, Venezuela,
Brazil, Peru).
campestris (Navas) 1930:107 (as Furgus). Hoi. $ , Guayaquil, Ecuador 1930
(Barcelona). New Comb.
garciai (Navas) 1932:12 (as Nobra). Hoi. 9, Vinces, Ecuador (Turin).
New Comb.
martinsi (Navas) 1915b:6 (as Nobra). Hoi. 9, Rio Purus, Brasil 1904
(Paris). Taxonomy: Stange (1963:812).
menkei Stange 1963:814. Hoi. 5, Rio Cuchujachi, 10 miles S.E. Alamos,
Sonora, Mexico, V.22.1962 (CAS).
praedator (Walker) 1853:391 (as Myrmeleon) . Hoi. 9, Santarem, Brazil
(BM).
Taxonomy: Hagen (1860:364) ; Stange (1963:812).
psammophila Stange 1963:814. Hoi. 9, Veracruz, Mexico, IV.29.1962
(CAS).
riparius (Navas) 1918b: 16 (as Nobra). Lect. 5, Santa Fe, Argentina, 1.9.
1917 (Barcelona).
—Dimarella pallida Navas 1933a:89. Hoi. 9, S. Miguel, Argentina, XI. 25.
1932 (Barcelona).
Taxonomy: Stange (1968:59).
Distribution: Neotropical (Uruguay, Argentina).
silvaticus (Navas) 1918b:6 (as Nobra). Hoi. 9, Mato Grosso, Brasil (La
Plata). N. Comb.
tarsalis (Guilding) 1829:47 (as Formicaleo) . Type (?s), ?Demerara (not
located) .
^Myrmeleon efferus Walker 1853:388. Hoi. 9, Para, Brazil (BM).
Taxonomy: Hagen (1860:364) ; Banks ( 1913a:229; 1913b:86; 1943:168);
Stange (1963:812).
Genus Elachyleon Esben-Petersen
Elachyleon Esben-Petersen 1927:348. Type-species: Elachyleon piinctipennis
Esben-Petersen, orig. design.
=Oroleon Navas 1927:49. Type-species: Oroleon serranus Navas, orig. de-
sign.
=Sericoleon Esben-Petersen 1932:114. Type-species: Sericoleon paessleri
Esben-Petersen, orig. design. New Synonymy.
Taxonomy: Banks ( 1943 : 166) ; Stange (1963:810; 1968:56).
gentilis (Banks) 1943:171 (as Sosa). Hoi. $, Kartabo, British Guiana, XI.4
(MCZ). New Comb.
paessleri (Esben-Petersen) 1932:115. (as Sericoleon) . Hoi. 9, Taltal (Anto-
fagasta), Chile, 1.30.1906 (Copenhagen). New Comb,
punctipennis Esben-Petersen 1927:348. Hoi. (sex undet.), St. Augustine,
Trinidad, V.20. 1925 (BM).
20
Contributions in Science
No. 186
=Elachyleon punctipennis pulchellus Esben-Petersen 1932:113. Syntypes:
9 , Las Mercedes, Costa Rica, IV.27.1922 (Hamburg, destroyed) ; sex un-
det., Brazil (Berlin?).
Taxonomy: Banks ( 1943: 168) ; Stange (1963:815; 1968:56).
Distribution: Neotropical (Mexico to Argentina).
serranus (Navas) 1927:50 (as Oroleon) . Hoi. 9, Alta Gracia, La Granja,
Sierras de Cordoba, Argentina, 1.8.1928 (Buenos Aires).
Taxonomy: Stange (1968:57).
Genus iremoleon Banks
Eremoleon Banks 1901:366. Type-species: Mynneleon macer Hagen, orig.
design.
=Incamoleon Banks 191 3a: 229. Type-species: Psammoleon punctipennis
Banks, orig. design. New Synonymy.
—Glenopsis Banks 191 3a: 229. Type-species: Myrmeleon anomalus Ram-
bur, orig. design.
—Sosa Navas 1914a:218. Type-species: Sosa conspicuus Navas, orig. de-
sign. New Synonymy.
—Segura Navas 1914c: 18. Type-species: Segura vitreus Navas, orig. de-
sign.
—Belen Navas 1921:119. Type-species: Belen cerverinus Navas, orig. de-
sign.
=Cortesius Navas 1924:107. Type-species: Cortesius genini Navas, orig.
design. New Synonymy.
=Novulga Navas 1925:189. Type-species: Novulga mexicanaNdiwks, orig.
design.
—Dobla Navas 1926:428. Type-species: Dobla arcuata Navas, orig. design.
New Synonymy.
=Joergenia Esben-Petersen 1932:118. Type-species: Joergenia pulchra
Esben-Petersen, orig. design. New Synonymy.
=Antilloleon Banks 1943:168. Type-species: Glenurus cerverai Navas,
orig. design. New Synonymy.
Taxonomy: Navas (1916:232; 1917:275); Banks (1928:69-71; 1942:
144); Adams (1957:85; 1958:6); Alayo (1968:62, 70); Stange (1968:
57).
anomalus (Rambur) 1842:388 (as Myrmeleon) . Type (?s), Colombia (not
located) .
— Glenurus mollis Gerstaecker 1888:101. Hoi. (sex undet.) Colombia
(Greifswald) .
Taxonomy: Banks (1913a:229; 1943:171); Navas (1916:232); Stange
(1968:57).
Distribution: Neotropical (Colombia, Venezuela, Brazil, Argentina).
1970
Catalog of the Western Hemisphere Glenurini
21
capitatus (Navas) 1913:52 (as Formicaleo) Hoi. (sex undet.), Rio de Ja-
neiro, Brasil, VI. 1911 (not located). New Comb,
cerverai (Navas) 1921:118 (as Glenurus). Hoi. 9, Santiago (de las) Vegas,
Habana, Cuba, VIII.23.1907 (MCZ). New Comb.
Taxonomy: Banks ( 1943 :68) ; Alayo (1968:68).
cerverinus (Navas) 1921:120 (as Belen) . Hoi. 9, Rio Almendares, Habana,
Cuba, VIII.1.1915 (MCZ).
Taxonomy: Adams ( 1957:6) ; Alayo (1968:70).
conspicuus (Navas) 19 14a: 21 9 (as Sosa). Hoi. Brasil (Vienna). New
Comb.
femoralis (Banks) 1942:146 (as Psammoleon) . Hoi. (sex undet.), 20 miles
N.W. La Paz, Lower California VII. 16. 1938 (CAS). New Comb,
genini (Navas) 1924:108 (as Cortesius). Syntypes (sex undet.), Veracruz,
Mexico, 1921 (Paris). New Comb.
=DobIa arcuata Navas 1926:428. Hoi. 9, San Jose, Costa Rica (Paris).
New Synonymy.
gracile Adams 1957:90. Hoi. 9, Riverside, California, VIII.31.1939 (CAS).
impluviatus (Gerstaecker) 1893:30 (as Glenurus). Hoi. 9, Locotal, Bolivia
(Greifswald), New Comb.
insipidus Adams 1957:88. Hoi. 5, 5 miles S. San Miguel, Lower California,
VII.20.1938 (MCZ).
Distribution: Nearctic (Mohave-Colorado Desert; Vizcaino Desert).
longior Banks 1938a:225. Syntypes 9 9 , Yucatan, Mexico (MCZ).
Taxonomy: Adams ( 1957:92) .
macer (Hagen) 1861 :236 (as Myrmeleon) . Hoi. (sex undet.), Mexico
(not located) .
=Segura v it reus Navas 1914c: 18. Hoi. <$ , Cuernavaca, Mexique, 1871
(Paris).
=Novulga mexicana Navas 1925:189. Hoi. $ , Veracruz, Mexique, 1921
(Paris).
=Hesperoleon atomarius Navas 1933b: 105. Hoi. (sex undet.) , Cuernavaca,
Mexico, 1929-30 (Hamburg, destroyed) . New Synonymy.
Taxonomy: Banks (1928:70) ; Adams (1957:86).
Distribution: Nearctic (Arizona); Neotropical (Mexico) .
nigri basis Banks 1920:329. Syntypes $ , 9 , St. George, Utah, VI. 5, 6 (MCZ)
=Eremeleon affine Banks 1942:144. Hoi. $ , Miraflores, Lower California,
VII.8.1938 (CAS).
Taxonomy: Banks (1928:71 ); Adams (1957:91).
Distribution: Nearctic (deserts).
ornatipennis (Alayo) 1968:69 (as Antilloleon) . Hoi. Soroa, Pinar del
Rio, Cuba, IV. 1963 (Havana). New Comb,
pallens Banks 1941c: 101. Syntypes, Picacho Peak, Arizona, VII. 23 (MCZ) .
Taxonomy: Adams (1957:92).
22
Contributions in Science
No. 186
peterseni (Banks) 1922:59 (as Glenopsis) . Lect. $, Chanchamayo, Peru, XI
(MCZ). New Comb.
Taxonomy: Stange (1961:674).
psilocerus (Gerstaecker) 1893:32 (as Glenurus) . Hoi. (sexundet.), Merida,
Venezuela (Greifswald) . New Comb.
Taxonomy: Banks (1922:59; 1943:172).
pulchra (Esben-Petersen) 1932:118 (as Joergenia). Hoi. 9, Paraguay, II.
1932 (Copenhagen). New Comb.
punctipennis (Banks) 1910:147 (as Psammoleon). Hoi. 9, San Antonio,
Colombia (MCZ). New Comb.
—Formicaleo tetrastictus Navas 1913:51. Types(?), Venezuela (not
located).
—Formicaleo stictopterus Navas 1916:232. New name for punctipennis
Banks 1910. (nomen nudum).
Taxonomy: Banks (1913a:299; 1943 : 169) ; Stange (1968:58).
Distribution: Neotropical (Colombia to Argentina).
sectoralis Adams 1958:7. Hoi. 9, 5 miles S. San Miguel, Lower California,
VII.20.1938 (CAS).
triguttatus (Navas) 1914c: 19 (as Formicaleo) . Hoi. 9, San Pedro de Sula,
Honduras (Vienna). New Comb.
Genus Glenurus Hagen
Glenurus Hagen 1866:372. Type-species: Formicaleo grata Say, design, by
Banks (1928:67).
=Lec/o5'c/w5’ Navas 1918:493. Type-species: Ledoscius penningtoni Navas,
orig. design.
SYNONYMY: Navas (1924:107).
gratus (Say) 1839:45 {as Formicaleo) .
=Myrmecoleon roseipennis Burmeister 1839:995. Hoi. (sex undet.), Nord-
Amerika (Halle).
Taxonomy: Walker (1853:392); Hagen (1861:225; 1866:405); Banks
(1892:360; 1928:67).
Distribution: Nearctic (East 100° Meridian).
heteropteryx GtTsXaQokQY 1885:17. Hoi. 9, Chiriqui (Greifswald).
^Glenurus discorsNawas 1920:202. Hoi. 9, La Trinite, 1916 (Paris).
Taxonomy: Banks (1922:58); Navas (1922b:187; 1935:362).
Distribution: Neotropical (Panama, Venezuela, Trinidad, Ecuador).
incalis Banks 1922:58. Syntypes, Chanchamayo, Peru (MCZ).
/i/n/ger Gerstaecker 1893:125. Hoi. 9, Chiriqui (Greifswald).
Taxonomy: Banks (1938:420).
Distribution: Nearctic (Arizona); Neotropical (Mexico to Panama).
peculiaris (Walker) 1859:194 (as Myrmeleon) . Hoi. 9, Brazil (BM).
1970
Catalog of the Western Hemisphere Glenurini
23
—Glenuriis brasiliensis Navas 1920:416. Lect. $, Sao Sebastiao, Brasil
(not located). New Synonymy.
Taxonomy: Hagen (1866:405); Banks (1922:58); Navas (1923:771).
penningtoni (Navas) 1918:493 (as Ledoscius) . Hoi. La Rioja, Argentina
(not located).
—Glenurus croesus Banks 1 922 : 59. Syntypes, Sara, Bolivia, 450 m. ( MCZ) .
Taxonomy: Navas (1922b: 187; 1924:107).
Distribution: Neotropical (Bolivia, Argentina),
pro/ Navas 1929:17. Hoi. $, Colima, Mexico (Paris).
snowii Banks 1907:100. Hoi. $, Baboquivari Mts., Arizona (MCZ).
Taxonomy: Banks (1928:67, 69).
Genus Navasoleon Banks
Navasoleon Banks 1943:168. Type-species: Gymnocnemia boliviana Banks,
orig. design.
Taxonomy: Stange (1963:810).
boliviana (Banks) 1929:330 (as Gymnocnemia) Hoi. $, Rio Longo, Bolivia
(MCZ).
Taxonomy: Banks (1943:168).
bosqiii (Navas) 1922:258 (as Gymnocnemia) . Hoi. 9, Santiago del Estero,
Argentina (not located).
Taxonomy: Stange (1968:58).
leptocera (Navas) 1915:125 (as Gymnocnemia) . Hoi. (sex undet.) Cata-
marca, Argentina, 1.21.1910 (Buenos Aires).
Taxonomy: Stange (1968:58).
Genus Pachyleon Stange
Pachyleon Stange 1970: Type-species: Pachyleon alvarengai Stange, orig.
design.
alvarengai Stange. Hoi. $ , Jacare, Mato Grosso, Brazil, XI. 1961 (LACM) .
Genus Psammoleon Banks
Psammoleon Banks 1899:69. Type-species: Myrmeleon ingeniosus Walker,
orig. design.
=Diazus Navas 1914:220. Type-species: Diazus clavatus Navas, orig. de-
sign.
Taxonomy: Banks (1913:226; 1928:60-67; 1942:166; 1943:169); Alayo
(1968:65).
albovaria (Banks) 1942:146 (as Puren) . Hoi. (sex undet.) , Venancio, Lower
California, VII.17.1938 (CAS). New Comb,
arizonensis Banks 1935:53. Syntypes : 19, Tucson, Arizona (MCZ) ; 2 9 9 ,
Phoenix, Arizona (MCZ).
24
Contributions in Science
No. 186
Distribution : Nearctic (Arizona-Sonora Desert; Mohave-Colorado Desert) .
banksi Esben-Petersen 1932:115. Hoi. Amapala, Honduras, XL 12. 1907
(Hamburg, destroyed).
bipunctatus (Navas) 1915:465 (as Formicaleo) . Hoi. $, Guyana Francesa
(Paris) .
Taxonomy: Banks (1943:170).
bistictus (Hagen) 1861:235 (as Myrmeleon) . Hoi. (sex undet.), Cuba
(Berlin?).
Taxonomy: Navas (1921 : 1 17) ; Banks (1928:61; 1941 : 177) ; Alayo
(1968:65).
Distribution: Neotropical (so. Florida, West Indies, Yucatan, Mex. ) .
cautus (Walker) 1853:349 (as Myrmeleon) . Hoi. $ , Brazil (BM) .
=Feinerus nebulosus Navas 1922:21. Hoi. 2, Peru 1920 (Paris). New
Synonymy.
Taxonomy: Banks (1943: 170).
clavatus (Navas) 1914:221 (as Diazus). Hoi. $ , Amazones 1860 (Vienna) .
New Comb.
connexus (Banks) 1920:329 (as Puren) . Hoi. S , San Jacinto Mts., California
VI.25 (MCZ).
Taxonomy: Banks (1928:61, 64) .
cubensis Alayo 1968:67. Hoi. (sex undet.), Tortuguilla, Guantanamo, Or-
iente, Cuba, VI. 1964 (Havana).
debilis (Gerstaecker) 1893:44 (as Formicaleo) . Hoi. 2, Chiriqui (Greifs-
wald) .
Taxonomy: Banks (1943:170).
Distribution: Neotropical (Panama; Colombia).
decipiens Banks 1935:54. Syntypes: IS, Georgia (MCZ) ; sex undet.,
Shreveport, Louisiana (MCZ).
giittipes Banks 1906:99. Hoi. $ , Tryon, North Carolina (MCZ).
Taxonomy: Banks (1928:61, 63).
im bell is (Banks) 1941 : 102 (as Puren) . Hoi. 2 , Port au Prince, Haiti, XI.20
(MCZ). New Comb.
ingeniosus (Walker) 1853:337 (as Myrmeleon) . Syntypes, Brazil & no data
(BM).
Taxonomy: Hagen (1861:236; 1866:404); Banks (1899:69; 1906:99;
1943:169).
Distribution: Neotropical (Colombia, Brazil) .
iniquus (Navas) 1914:208 (as Formicaleo) new name for Formicaleo in-
aequalis Navas 1913:51 (Preoccupied by F. inaequalis Navas 1912) : Hoi.
(sex undet.) , Amapola, Honduras, XI (not located) . New Comb,
inscriptus (Hagen) 1861 :230 (as Myrmeleon) . Hoi. S , Pecos River, Western
Texas (MCZ) .
Taxonomy: Hagen ( 1866:424) ; Banks ( 1904: 106; 1928:66).
1970
Catalog of the Western Hemisphere Glenurini
25
Distribution: Nearctic (Arizona-Sonora Desert).
leachii (Guilding) 1829:49 (as Formicaleo) . Types (?), Jamaica (not lo-
cated) .
minor Banks 1928:62. Lect. $, Dry Tortugas, Loggerhead Key, Florida,
VI.1917 (MCZ).
Taxonomy: Banks (1941:176, 177); Stange (1961:674); Alayo (1968:
66).
Distribution: Neotropical (So. Florida; West Indies).
normalis Banks 1942:145. Hoi. $, Venancio, Lower California, VII.17.1938
(CAS).
nubipennis (Navas) 1917:275 (as Formicaleo). Hoi. (sex undet.), Coachi,
Colombia, VI. 19 15 (not located). New Comb,
parallela Banks 1935:54. Syntypes, Honduras (MCZ).
Banks 1920:331. Hoi. 5, Chapada, Brazil (MCZ).
Taxonomy: Banks (1943:170).
posticatus Banks 1941:3. Hoi. (sex undet.), Colombia (AMNH).
Taxonomy: Banks (1943:169).
reductus Banks 1941:177. Hoi. 5 , Stakes Bay, Cayman Brae, Cayman Islands
(MCZ).
serrei (Navas) 1920:201 Formicaleo) . F\o\. 5 , La Trinite, 1914 (Paris).
New Comb.
—Formicaleo chaperi Navas 1922:256. Hoi. S, Venezuela, 1885 (Paris).
New Synonymy.
Taxonomy: Banks (1943:170).
sinuatus Currie 1903:275. Hoi. $ , Santa Rita Mts., Arizona, V.31 (USNM).
=Psammoleon serpentinus Navas 1922:185. Hoi. (sex undet.), Jemez
Springs, New Mexico, VII. 1916 (Zaragoza).
Taxonomy: Banks (1928:5, 64).
zay as i Alsiyo 1968:67. Hoi. 9 , Peninsula de Guanahacabibes, Pinar del Rio,
Cuba, 1956 (Private coll. Zayas, Habana).
Resumen
Revision Generica y Catalogo de la Tribu Glenurini del Hemisferio
Occidental con la Descripcion de un Nuevo Genero y Especies de Brasil.
Estudios sistematicos de la Glenurini del hemisferio occidental han
llevado a grandes cambios en su clasificacion. Ocho nombres genericos han
sido reducidos a sinominos y un genero y especie nueva, Pachyleon alvarengai
es descripto de Brasil. Estos sinonimos son: Elachyleon Esben-Petersen 1927
i=Sericoleon Esben-Petersen 1932); Eremoleon Banks 1901 {=Incamoleon
Banks 1913, =Sosa Navas 1914, —Cortesius Navas 1924, =Dobla Navas
1926, —Joergenia Esben-Petersen 1932, = Antilloleon Banks 1943); Psam-
moleon Banks 1899 {—Diazus Navas 1914). Se dan ahora ocho generos
validos. Se incluyen descripciones y observaciones sobre cada genero ademas
26
Contributions in Science
No. 186
de una clave para su identificacion. Se discute la posicion sistematica de la
tribu Glenurini que se pasa a la subfamilia Myrmeleontinae, con la inclusion
de los generos de la tribu Dimarellini. Se agrega un catalogo sinoptico de las
especies y generos de la tribu con datos sobre los tipos y se indican muchas
combinaciones nuevas ademas de cuatro sinonimos especificos nuevos. Estos
son: Psammoleon cautus (Walker) 1853 {—Feinerus nebulosus Navas 1922);
Psammoleon serrei (Navas) 1920 {—Formicaleo c/zapm Navas 1922); Glen-
urus peculiaris (Walker) 1859 {—Glenurus brasiliensis Navas 1920) ; Eremo-
leon macer (Hagen) 1861 {~Hesperoleon atomarius ^diwis 1933).
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. 1958. Further notes on the genus Eremoleon Banks, with a new species
(Neuroptera: Myrmeleontidae). Psyche, 64:6-8, figs. la-e.
Alayo, D. P. 1968. Los Neuropteros de Cuba. Poeyana (B), 2:5-127, 31 figs, 10 pis.
Banks, N. 1892. A synopsis, catalogue, and bibliography of the neuropteroid insects
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. 1901. A list of neuropteroid insects from Mexico. Amer. Ent. Soc., Trans.,
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. 1924. Descriptions of new neuropteroid insects. Harvard, Mus. Comp. Zool.,
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Bull., 68:1-84, 101 figs.
. 1935. A few new North American Neuroptera. Psyche, 42:53-57.
-. 1938a. A new myrmeleonid from Yucatan. In Pearse, A. S., et al.. Fauna of
the caves of Yucatan. Carnegie Inst. Wash. Publ., 491:225.
. 1938b. Two new genera of Myrmeleonidae. Ent. Soc. Wash., Proc., 40:125-
129, 4 figs.
. 1941a. Some new and interesting Neuroptera in the American Museum of
Natural History. Amer. Mus. Novitates, 1 143: 1-5.
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Catalog of the Western Hemisphere Glenurini
27
. 1941b. Results of the Oxford University Cayman Islands biological expedi-
tion, 1938 (Neuroptera). Entomol. Mon. Mag., 77:176-177.
. 1941c. Three new species of Myrmeleonidae (Neuroptera). Psyche, 48:101-
104.
. 1942. Contributions toward a knowledge of the insect fauna of Lower Cali-
fornia. No. 4. Myrmeleonidae. Calif. Acad. Sci., Proc., (4) 24:133-152, 17 figs.
. 1943. Neuroptera of northern South America. Part 2. Myrmeleonidae. Bol.
ent. Venezolana, 2:161-173.
Burmeister, H. 1839. Handbuch der Entomologie. Berlin. Gymnognatha. (2): 757-
1017.
Currie, R. P. 1903. Myrmeleonidae from Arizona. Ent. Soc. Wash., Proc., 5:272-
284.
Esben-Petersen, P. 1927. New and little-known species of Neuroptera in British
collections. Part III. Ann. Mag. Hist., (9) 20:343-350, 8 figs.
. 1932. New and little-known Neuroptera. Dansk naturh. Foren, Vidensk.
Medd. 94:109-123, 13 figs.
Gerstaecker, a. 1885. Vier Decaden von Neuropteren aus der Familie Megalop-
tera, Burm. Naturw. Ver. Greifswald, Mitt., 16: 1-49.
. 1888. Weitere Beitriige zur Artenkenntniss der Neuroptera Megaloptera.
Naturw. Ver. Greifswald, Mitt., 19:89-130.
. 1893. liber neue und weniger gekannte Neuropteren aus der Familie Megal-
optera, Burm. Naturw. Ver. Greifswald, Mitt., 25:93-173.
Guilding, L. 1829. The generic characters of Fonnicaleo\ with the description of two
new species. Linn. Soc. London, Trans., 16:47-51.
Hagen, H. 1860. Beitrag zur Kenntniss der Myrmeleon-Avitn. Stettin, ent. Ztg., 21:
359-369.
. 1861. Synopsis of the Neuroptera of North America. With a list of the South
American species. Smithson. Misc. Coll., 4: 1-347.
. 1866. Hemerobidarum Synopsis Synonymica. Stettin, ent. Ztg., 27:369-462.
Markl, W. 1954. Vergleichend-morphologische Studien zur Systematik und Klassi-
fikation der Myrmeleoniden. Naturf. Ges. Basel, Verb., 65: 178-263, 85 figs.
Navas, L. 1913. Neuropteros nuevos de America. Broteria, 11:45-53, 6 figs.
. 1914a. Neuropteros sudamericanos. Primera Serie. Broteria, 12:45-57, 215-
235, 11 figs.
. 1914b. Neuropteros nuevos o poco conocidos. Segunda & Tercera Serie.
Acad. Cienc. Barcelona, Mem., (3) 11(8, 13): 105-1 19, 193-215, 14 figs.
. 1914c. Neuropteres nouveaux de I’Amerique du Nord. II. Ent. Zeitschr., 28:
18-20, 4 figs.
— . 1915a. Neuropteros nuevos o poco conocidos. Quinta Serie. Acad. Cienc.
Barcelona, Mem., (3) 1 1(27) :455-480, 18 figs., pi. 1, 6 figs.
— . 1915b. Neuropteros sudamericanos. Segunda Serie. Broteria, 13:5-13, 6 figs.
. 1916. Neuropteros nuevos o poco conocidos. Acad. Cienc. Barcelona, Mem.,
(3) 12(13):219-243.
— — . 1917. Neue Neuropteren. Ent. Mitteil., 6:274-282, 5 figs.
. 1918a. Algunos insectos de la Republica Argentina. Acad. Cienc. Madrid,
Rev., 16:491-504, 10 figs.
. 1918b. Insecta nova. IV Serie. Apcad. Pont. Nuovi Lincei, Mem., (2) 4:1-
23, 5 figs.
. 1919. Algunos insectos Neuropteros de la Republica Argentina. Serie Ter-
cera. Acad. Cienc. Madrid, Rev., 17:287-305, 6 figs.
— — — . 1920. Sur des Nevropteres nouveaux ou critiques. Deuxieme Serie. Soc. sci.
Bruxelles, Ann., 39:189-203, 5 figs.
28
Contributions in Science
No. 186
. 1921. Insectos americanos nuevos o criticos. Broteria, 19:113-124, 5 figs.
— . 1922a. Insectos sudamericanos. Cuarta Serie. Acad. Cienc. Madrid, Rev.,
19:255-267.
-. 1922b. Estudis sobre Neuropters (Insectes). Inst. Cien. Barcelona, Arx., 7:
179-203, 2 figs.
. 1922c. Algunos insectos del museo de Paris. Acad. Cien. Zaragoza, Rev., 7:
15-27, 6 figs.
-. 1924. Algunos insectos del Museo de Paris. Broteria, 21:99-114, 5 figs.
-. 1925. Insectos exoticos nuevos o poco conocidos. Acad. Cienc. Barcelona,
Mem., (3) 19:181-200, 12 figs.
— -. 1926. Neue Insekten (Neuropt.). Deutsch Ent. Zeitschr., 1926:428-431, 2
figs-
. 1927. Veinticinco formas nuevas de insectos. Bol. Soc. iber. cien. nat., Zara-
goza, 26:48-75, 9 figs.
. 1929. Insectos neotropicos. 5a Serie. Rev. chil. Hist. Nat. 33: 17-24, 3 figs.
. 1930. Neuropteros del Ecuador. Rev. chil. Hist. Nat. 34:107-109, 1 fig.
— . 1932. Alcuni insetti del Museo di Zoologie della R. Universita di Torino.
Turin, Univ., Mus. ZooL, Boll., (3) 42 (26) : 1-38, figs.
— -. 1933a. Insectos de la Argentina. Rev. Acad. Cienc. Zaragoza, 16:87-120, 22
figs.
— . 1933b. Decadas de insectos nuevos. Broteria (Cienc. Nat.), 2: 101-110.
. 1935. Insectos sudamericanos. Decima Serie. Acad. Cienc. Madrid, Rev.,
32:360-375.
. 1936. Decadas de insectos nuevos. Broteria (Cienc. Nat.) 32:161-170.
Rambur, J. P. 1842. Histoire naturelle des insectes. Nevropteres. Paris. 534 p., tab.
col. i-xii.
Say, T. 1839. Descriptions of some N. American Neuropterous Insects, and observa-
tions on some already described. Acad. Nat. Sci. Philadelphia, Journ., 8:9-46.
Smith, R. C. 1931. The Neuroptera of Haiti, West Indies. Ent. Soc. Amer., Ann., 24:
798-823, 21 figs.
Stance, L. A. 1961. Lectotype designations in the New World Myrmeleontidae.
Canad. Ent., 93:674-677.
. 1963. The Dimarellini of Mexico with the descriptions of two new species of
Dimarella. Ent. Soc. Amer., Ann., 56:810-816, 16 figs.
. 1968. Catalogo de Neuroptera de Argentina y Uruguay. Acta Zool. Lill., 22
(1967):5-86.
— . 1970. Revision of the Ant-lion Tribe Brachynemurini of North America.
U.C. Publ. Ent. 55:1-192, 231 figs., 31 maps.
Walker, F. 1853. List of the specimens of neuropterous insects in the collection of
the British Museum. Part II (Sialidae — Nemopterides) . London: 193-476.
. 1859. Characters of undescribed Neuroptera in the collection of W. W.
Saunders. Ent. Soc. London, Trans., 5 : 176-199.
Accepted for publication February 18, 1970
I LOS
* ANGELES
I COUNTY
i MUSEUM
I
CONTRIBUTIONS
IN SCIENCE
MBER 187
C;i L ^ a
May 4, 1970
FURTHER STUDIES ON AUDIBLE VOCALIZATIONS
OF THE AMAZON FRESHWATER
DOLPHIN, INIA GEOFFRENSIS
By Melba C. Caldwell and David K. Caldwell
Los Angeles County Museum of Natural History • Exposition Park
Los Angeles, California 90007
( 1 IQ,:
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 sep-
arately, and numbers run consecutively regardless of subject matter. Number 1 was
issued January 23, 1957. The series is available to scientific institutions and scien-
tists on an exchange basis. Copies may also be purchased at a nominal price. Inquiries
should be directed to Virginia D. Miller, Los Angeles County Museum of Natural
History, 900 Exposition Boulevard, Los Angeles, California 90007.
Robert J. Lavenberg
Managing Editor
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Manuscripts for the LOS ANGELES COUNTY MUSEUM, CONTRIBU-
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papers will be determined by the amount and character of new information. Al-
though priority will be given to manuscripts by staff members, or to papers dealing
largely with specimens in the Museum’s collections, other technical papers will be
considered. All manuscripts must be recommended for consideration by the curator
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to those specifications listed below and will be examined for suitability by an Edi-
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side the Museum.
Authors proposing new taxa in a CONTRIBUTIONS IN SCIENCE must
indicate that the primary type has become the property of a scientific institution of
their choice and cited by name.
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published at the head of each paper. (7) A Spanish summary is required for all
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Virginia D. Miller
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FURTHER STUDIES ON AUDIBLE VOCALIZATIONS
OF THE AMAZON FRESHWATER
DOLPHIN, INIA GEOFFRENSIS
By Melba C. Caldwell^ and David K. Caldwell^
Abstract: Evidence is presented indicating the absence of a
quasi-pme tone or whistle type of sound production by the Ama-
zon freshwater dolphin. A discussion and sonagrams are given of
the audible squeal or screech often attributed to Inia geoffrensis.
Phonations of the toothed whales and dolphins fall roughly into three
categories. These include two types which generically are related (Norris,
1969) and probably are produced by the same mechanism, as they both con-
sist of discrete clicks. One type demonstrates a slower click repetition rate
and is usually associated with environmental investigation (echolocation),
whereas the other is made up of bursts of clicks of higher repetition rates of
150 per second or more (Caldwell, Caldwell and Evans, 1966b). The latter
create tonal sounds variously labeled by such names as barks, squawks,
screeches or squeaks. These are associated with behavioral situations wherein
we find a positive correlation between the time of their emission and the use-
fulness of a signal that could cue conspecifics to approach or withdraw (Cald-
well and Caldwell, 1967). The third type of sound, most frequently termed a
whistle, is a quasi-^mo, tone sound that cannot be resolved into individual
clicks. These are the harmonic emissions of Tavolga ( 1965: 9) and Evans
(1967). All odontocetes probably emit click sounds (Norris, 1968), but
some do not emit the so called pure tone.
No whistles have been reported in sound work on the primitive Amazon
freshwater dolphin, Inia geoffrensis (see Schevill and Watkins, 1962; Schevill,
1964; Caldwell, Caldwell and Evans, 1966a, 1966b; Caldwell and Caldwell,
1967; Evans, 1967; Poulter, 1968; Caldwell and Caldwell, 1969b).
All of these earlier works contain two deficiencies. First, on only one
brief occasion was work done with more than one or two animals in a tank
(Caldwell and Caldwell, 1967) . This lack of community tank sound studies
constitutes a potentially serious deficit, as vocalizations of odontocete ceta-
ceans of the marine species with which we have worked tend to diminish in
relative number in captivity if other animals are not present. Second, several
^Research Associate in Cetology, Los Angeles County Museum of Natural History;
Staff Research Associate, Communication Sciences Laboratory, University of Flor-
ida, Gainesville.
^Research Associate in Cetology, Los Angeles County Museum of Natural History;
Associate Professor, Communication Sciences Laboratory, and Staff Research Asso-
ciate, Florida State Museum, University of Florida, Gainesville.
1
2
Contributions in Science
No. 187
people handling Inia have reported a loud squeal or screech emitted by the
animal which, from the observers’ descriptions, might have been interpreted
as a whistle (Allen and Neill, 1957; Layne and Caldwell, 1964; Caldwell,
Caldwell and Evans, 1966b; Caldwell and Caldwell, 1969a, 1969b). This
sound also has been described to us in personal conversation with Earl S.
Herald of the Steinhart Aquarium, San Francisco; Lawrence Curtis, formerly
of the Fort Worth Zoo, Texas; Leo Baumer of Iquitos, Peru; and W. J.
LeBlanc and William C. Raulerson, Marineland of Florida. Until recently
we had heard the sound out of water on one occasion, but did not record it.
Further studies on this sound were thus considered necessary to determine
whether these sounds are broad-band pulsed sounds and not a narrow-band
whistle. Correction of these two deficiencies seemed mandatory, not only to
help solidify our basic knowledge of cetacean phonations, but also because we
have come to regard the absence or presence of the pure tone whistle as one
of the significant characters in the precariously understood familial relation-
ships between members of the order Cetacea. Evidence to date suggests that
the species of cetaceans considered to be more primitive also either lack the
quasi-^xxxQ tone whistle or that when present it is less clear-cut than in those
species considered to be more advanced.
We recorded Inia phonations from a community tank over a period of
two years for a total of some 25 hours. The dolphins were held in a large
display tank at Marineland of Florida and at various times the community
contained from four to eleven animals of mixed sizes and sexes. Sound record-
ings were made with the animals at rest or swimming slowly, feeding, fighting,
copulating, sick, dying, and semi-stranded. The community tank contained
eleven animals during one seven hour recording session of a semi-stranding.
The animals were recent captives which, in our experience, along with strand-
ing increases the possibility of eliciting at least a few cetacean phonations.
Additionally we have air recorded isolated individuals as they were being
handled and medicated, as well as during venipuncture and force feeding.
Within the range of our equipment (40 to 20,000 Hz), we have not
detected quasi-^mc tone sounds, nor did Evans {in Herald, et al., 1969; per-
sonal conversation, August, 1969) indicate the production of such sounds
in his studies of Inia phonations with equipment sensitive to frequencies up to
100,000 Hz. The loud squeals that we recorded are pulsed as shown when
they are subjected to detailed analysis, and the quasi-^mo, tone or whistle is
still unknown in the vocal repertoire of Inia and we believe now that it is not
to be expected.
An adult female emitted squeals both singly and in series 22 and 19
times on two successive days when she was removed from her tank for medi-
cation. On another occasion this same female, although not being handled,
intermittently emitted loud squeals for about an hour when the water level
in her tank was so low that she was partly stranded; but no recordings were
1970
Vocalizations of the Amazon Dolphin
3
0 0.2 0.4
TIME (SEC.)
0 0.2 0.4 0.6 0.8 1.0 1.2 1.4 1.6 1.8 2.0 2.2
TIME (SEC.)
Figure 1. Phonations of Inia geoffrensis. Squeals emitted in air in daylight by isolated
animals during application of medication. Upper: Adult female (MLF 165), Janu-
ary 27, 1969. Lower: Juvenile male (MLF 202), March 10, 1969. Effective filter
bandwidths 300 Hz. (Photographs courtesy Marineland of Florida).
made. A juvenile male emitted seven squeals on one occasion when removed
for medication and venipuncture. The four occasions are the only times of
perhaps 30 to 40 in-air recording or listening sessions that the squeal was
emitted by any individual of this species although all were subjected to the
same amount of handling.
Sound spectrograms (sonagrams) of the squeals of both animals (Fig. 1)
show them to be the burst-pulse type with a high click repetition rate (Wat-
kins, 1967), Although these are broad band clicks with some energy extend-
ing above 12 kHz (Fig. 2), a strong fundamental frequency at one to two
kHz is demonstrated on the sonagrams of the sounds of both animals (Fig. 1).
These sonagrams show several characteristics in common with the type 2
4
Contributions in Science
No. 187
0 0.1 0.2 0.3 0.4 0.5 0.6 0.7 0.8 0.9 |.0
TIME (SEC.)
Figure 2. Phonations of Inia geoffrensis. First two of the same three squeals illus-
trated in Fig. 1 (lower), but played at half recorded speed. Effective filter band-
width 600 Hz. (Photograph courtesy Marineland of Florida).
or complaint type of signal of the pilot whale, Globicephala melaena, (Busnel
and Dziedzic, 1966: 615, fig. 11), the tin horn sound emitted when excited
by a strong stimulus by the Pacific whitesided dolphin, Lagenorhynchus
obliquidens, (Caldwell and Caldwell, 1967 : 889, figs. 5-7), and the squawk
component of the whistle-squawk emitted in air by an excited Atlantic bottle-
nosed dolphin, Tursiops truncatus, (Caldwell and Caldwell, 1967: 897, fig.
22). Fundamental frequencies, depicted harmonics, and durations are similar
in the four species. However, to the human ear the sounds are not alike.
Sound Equipment
The recordings discussed in this paper were made with a Uher 4000
Report-L recorder operating at a tape speed giving it a frequency response
of 40 to 20,000 Hz with a compatible Uher microphone. Sound spectrograms
were prepared on a Kay Sona-Graph model 662A Sound Spectrograph Ana-
lyzer calibrated in two sections from 85 to 12,000 Hz. When the recorded tape
speed is reduced by half, and then fed into the analyzer, the response of the
latter is doubled to 24,000 Hz. The effective filter bandwidths used for the
illustrated analyses are indicated in the figure captions.
Acknowledgments
Financial support for certain phases of this work came from the National
Science Foundation (GB-1189), the National Institute of Mental Health
(MH-07509-01), the Office of Naval Research (N00014-67-C-0358, and
modifications POOl and P002), and Marineland, Inc. The photographs are by
William A. Huck. One of our associates, Nicholas R. Hall, made the air
recordings that included the squeals.
1970
Vocalizations of the Amazon Dolphin
5
Literature Cited
Allen, Ross, and Wilfred T. Neill. 1957. White whales of the Amazon. Nat.
Hist. 66:324-329.
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 K. S. Norris, editor. Whales, dolphins, and porpoises. Berkeley:
Univ. Calif. Press, pp. 607-646.
Caldwell, Melba C., and David K. Caldwell. 1967. Intraspecific transfer of
information via the pulsed sound in captive odontocete cetaceans. In R.-G.
Busnel, editor, Les Systemes Sonars Animaux, Biologie et Bionique. Jouy-en-
Josas, France: Laboratoire de Physiologie Acoustique. 2:879-936.
— AND — . 1969a. The ugly dolphin. Sea Frontiers, 15(5) :308-314.
AND . 1969b. More about the ugly dolphin. Sea Frontiers, 15(6) :
349-355.
— , AND William E. Evans. 1966a. Preliminary results of studies on
the sounds and associated behavior of captive Amazon freshwater dolphins,
Inia geoffrensis. In Conf. on Biol. Sonar and Diving Mammals, Stanford
Research Institute, California, Proc., 3:178-185.
, AND . 1966b. Sounds and behavior of captive Amazon fresh-
water dolphins, Inia geoffrensis. Los Angeles County Mus., Contr. Sci., No.
108:1-24.
Evans, William E. 1967. Vocalization among marine mammals. In W. N. Tavolga,
editor. Marine bio-acoustics. New York: Pergamon Press. 2:159-186.
Herald, Earl S., Robert L. Brownell, Jr., Frederic L. Frye, Elkan J. Morris,
William E. Evans, and Alan B. Scott. 1969. Blind river dolphin : first side-
swimming cetacean. Science, 166:1408-1410.
Layne, James N., and David K. Caldwell. 1964. Behavior of the Amazon dolphin,
Inia geoffrensis (Blainville), in captivity. Zoologica, 49(2) :81-108, pis. 1-4.
Norris, Kenneth S. 1968. The evolution of acoustic mechanisms in odontocete
cetaceans. In Ellen T. Drake, editor, Evolution and environment. New Haven,
Conn. : Yale Univ. Press, pp. 297-324.
— . 1969. The echolocation of marine mammals. In H. T. Andersen, editor.
The biology of marine mammals. New York: Academic Press, pp. 391-423.
Poulter, Thomas C. 1968. Marine mammals. In T. A. Sebeok, editor. Animal
communication; techniques of study and results of research. Bloomington :
Indiana Univ. Press, pp. 405-465.
ScHEViLL, William E. 1964. Underwater sounds of cetaceans. In W. N. Tavolga,
editor. Marine bio-acoustics. New York: Pergamon Press. 1:307-316.
— AND William A. Watkins. 1962. Whale and porpoise voices. Woods Hole,
Mass.: Woods Hole Oceanogr. Inst. Phonograph record and 24-page booklet.
Tavolga, William N. 1965. Review of marine bio-acoustics; state of the art: 1964.
Port Washington, N. Y.: U. S. Naval Training Device Center, Tech. Rpt.:
NAVTRADEVCEN 1212-1. 100 pp.
Watkins, William A. 1967. The harmonic interval : fact or artifact in spectral
analysis of pulse trains. In W. N. Tavolga, editor. Marine bio-acoustics. New
York: Pergamon Press. 2:15-43.
Accepted for publication February 3, 1970
LOS
CONTRIBUTIONS
IN SCIENCE
i ANGELES
i
COUNTY
MUSEUM
JMBER 188
May 4, 1970
o
C:;iL
FOUR NEW TEREBRID GASTROPODS FROM
EASTERN PACIFIC ISLANDS
By Twila Bratcher and R. D. Burch
Los Angeles County Museum OF Natural EiiSTORY • 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 sep-
arately, and numbers run consecutively regardless of subject matter. Number 1 was
issued January 23, 1957. The series is available to scientific institutions and scien-
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Robert J. Lavenberg
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Virginia D. Miller
Editor
FOUR NEW TEREBRID GASTROPODS FROM
EASTERN PACIFIC ISLANDS
By Twila Bratcher^ and R. D. Burch-
Abstract: Four new terebrids are described from Eastern
Pacific islands: Terebra hertleini, T. jacqiielinae, and T. purdyae
from the Galapagos Islands, Ecuador, and T. stohleri from So-
corro Island, Mexico,
While examining the terebrid collections dredged by the R/V Valero III
during the Allan Hancock Foundation Pacific Expeditions and by the ex-
peditions of California Academy of Sciences to the Eastern Pacific islands, we
discovered three news species of Terebra’, the fourth was brought to our atten-
tion by Mrs. Jacqueline DeRoy, who dredged specimens at Academy Bay in
the Galapagos Islands.
The following abbreviations are used in the text: AHF, Allan Hancock
Foundation (material on loan to LACM); AMNH, American Museum of
Natural History; ANSP, Academy of Natural Sciences of Philadelphia; B&B,
Bratcher and Burch collection; BM(NH), British Museum (Natural History) ;
CAS, California Academy of Sciences; LACM, Los Angeles County Museum
of Natural History; MCZ, Museum of Comparative Zoology, Harvard Uni-
versity; SB, Santa Barbara Museum of Natural History; SD, San Diego
Museum of Natural History; SU, Stanford University; USNM, United States
National Museum; WAM, Western Australia Museum.
Acknowledgments
We wish to express our appreciation to Drs. James H. McLean of the
Los Angeles County Museum of Natural History, Leo G. Hertlein of the
California Academy of Sciences, and Harald Rehder and Joseph Rosewater
of the U.S. National Museum for their cooperation and loan of materials. We
thank Ben and Ruth Purdy of San Diego, California, and Laura Shy of West-
minster, California, in whose collections additional specimens of some of the
new species were found, for making their collections available to us. We are
indebted to Mrs. Jacqueline DeRoy for sending material from the Galapagos
Islands for study. We also wish to thank Maurice Giles of the California
Academy for preparing several of the photographs and Mrs. Margaret Hanna
for retouching them.
Terebra hertleini, sp. nov.
Figures 1-2
Diagnosis: A small species distinguished from other small terebrids by
turreted whorls and large subsutural nodes.
’Research Assistant in Invertebrate Zoology, Los Angeles County Museum of
Natural History. 8121 Mulholland Terrace, Los Angeles, California 90046.
“Formerly Research Assistant in Invertebrate Zoology, Los Angeles County Museum
of Natural History. (Deceased, February 22, 1970.)
1
1
Contributions in Science
Description of holotype: Size small; color white, whorls flat, turreted, ten
in number plus two glassy, convex nuclear whorls; first postnuclear whorl con-
stricted; sculpture of three spiral cords per whorl and obsolete axial ribs be-
ginning in large nodes anterior to suture (nine on penultimate whorl); sculp-
ture on body whorl of three spiral cords crossing obsolete ribs that end in faint
nodes at periphery, these nodes being less prominent than those at suture;
anterior to periphery spiral cords cross minute axial striations; aperture semi-
quadrate; outer lip thin, white within; columella straight with one weak
plication; siphonal fasciole well developed with sharp keel; anterior canal of
medium length, recurved. Length, 1 1.8 mm; diameter, 5.0 mm.
Type locality ; Academy Bay, Santa Cruz (Indefatigable) Island, Gala-
pagos Islands, Ecuador, 08° 46' 16" S, 90° 19' 38" W, CAS loc. 27536, 3. 5-5.5
fms, dredged, 45 specimens. Most of the specimens are very small and appear
not to have been live taken as they have a chalky appearance.
Type material: Holotype, CAS 13222. Paratypes: AMNH 157281 ;
ANSP 316670; B&B 741; BM(NH); CAS 13223-13227; LACM-AHF 1288;
SB 27147; SD 51962; SU 9996; USNM 680212. The LACM paratype is from
AHF station 168-34, 15-25 fms, and was live collected. Academy Bay.
Discussion: This species shows variation in the spiral cords being well
developed in some specimens and almost obsolete in others. The nodes at the
periphery of the body whorl are inconspicuous in some individuals. T erebra
hertleini has a superficial resemblance to T. jacquelinae sp. nov., but is a much
smaller species. The holotype of the former with ten whorls measures 1 1.8 mm
in length while that of the latter with 13 whorls measures 33.2 mm. T erebra
jacquelinae has a row of large nodes posterior to the suture which are absent
in T. hertleini.
T erebra hertleini is named for Dr. Leo G. Hertlein of the California
Academy of Sciences in appreciation of his encouragement and assistance in
our work on the eastern Pacific Terebridae.
Terebra jacquelinae, sp. nov.
Figures 3-4
Diagnosis: A species differing from other west American terebrids in
having extremely concave whorls and axial ribs that fade at the center of the
whorl and become large nodes at each end.
Description of holotype: Size medium; color shiny cream; first four
postnuclear whorls flat, remainder of whorls very concave, 1 3 in number plus
1.5 shiny, opaque, somewhat bulbous nuclear whorls; sculpture on first five
Figures 1-8. 1. T erebra hertleini sp. nov. Holotype, CAS 13222. X 9. 2. T. hertleini,
early whorls of holotype. 3. T. jacquelinae sp. nov. Holotype, CAS 13215. X 2.5.
4. T. jacquelinae, early whorls of holotype. 5. T. purdayae sp. nov. Holotype, LACM-
AHF 1 182. X 5. 6. T. purdyae, early whorls of holotype. 7. T. st older i sp. nov. Holo-
type, LACM-AHF 1 180. X 5.5. 8. T. stohleri, early whorls of holotype. Photos 1-5
by Maurice Giles; 6-8 by Twila Bratcher.
1970
Four New Terebrid Gastropods
3
4
Contributions in Science
postnuclear whorls of weak axial ribs ending in small nodes at suture and of
obsolete spiral grooves that cross the ribs; apical angle increases after sixth
postnuclear whorl and ribs begin to fade at center of whorl while nodes at rib
endings become more prominent, those anterior to suture being slightly more
prominent than posterior ones; interspaces on later whorls contain minute
striae, body whorl of medium length with 12 obsolete ribs ending in nodes
at periphery; anterior to periphery weakly incised spiral grooves cross equally
weak axial grooves; aperture elongate, white within; columella white, straight,
with one rounded plication that continues to become keel of well developed
siphonal fasciole; anterior canal short, recurved. Length, 33.2 mm; diameter,
8.4 mm.
Type locality: Academy Bay, Santa Cruz (Indefatigable) Island, Gala-
pagos Islands, Ecuador, 0° 46' 16" S, 90° 19' 38" W, CAS loc. 39585, about 10
fathoms, on hard packed coralline sand bottom, collected by Allyn G. Smith
and Andre and Jacqueline DeRoy, February, 1964, holotype and four para-
types.
Type material: Holotype, CAS 13215; 4 paratypes, CAS 13216-13219.
Additional paratypes are all from Academy Bay. CAS Templeton Crocker
Expedition, May, 1932, loc. 27536, 3. 5-5. 5 fms., 11 specimens, distributed as
follows: 8 paratypes, CAS 13220; 1 paratype, BM(NH) ; 1 paratype, SU 9997;
1 paratype, USNM 680213. Hancock Expeditions, AHF sta. 807-38, 10-25
fms., January, 1938, 28 specimens, distributed as follows: 26 paratypes,
L ACM- AHF 1179; 1 paratype, MCZ 271946; 1 paratype, SD 51963. Thirteen
additional paratypes were dredged by the DeRoys, 5-6 fms., 6 remain in the
DeRoy collection, others distributed as follows: 1 paratype, AMNH 157282;
1 paratype, ANSP 316671; 1 paratype, SB 27148; 1 paratype, WAM; 3 para-
types, B&B.
Referred material: Numerous specimens of this species were among
material taken by the Hancock and California Academy of Sciences expedi-
tions and by the DeRoys at collecting stations in the vicinity of Santa Cruz
(Indefatigable), San Salvador (James), and Baltra (Seymour) Islands in the
Galapagos Islands at depths of 2-20 fathoms.
Discussion: Although there is little variation among mature individuals
of this species except that some are more slender than others, occasional im-
mature specimens have the peripheral nodes forming a sharp keel that differs
in appearance from that of adults. Most of the specimens examined are of a
light cream color, but a number of specimens, most of which were collected at
San Salvador Island, range in color from beige to light brown with cream
colored nodes. The largest specimen examined measures 36.3 mm in length and
10.8 mm in width and is in the DeRoy collection. This species should not be
confused with Terebra frigata Hinds, 1844, which has less concave whorls,
pronounced spiral sculpture, a more slender profile, and ribs that are con-
tinuous from suture to suture. The Gulf of Mexico species, T. concava (Say,
1827), is a smaller and much more slender species with definite spiral sculp-
1970
Four New Terebrid Gastropods
5
tore, having small sharp nodes at the subsutural band and periphery of the
body whorl.
Terebra jacqiielinae is named for Mrs. Jacqueline DeRoy of Isla Santa
Cruz. Galapagos Islands, who first brought this species to our attention.
Terebm purdyae, sp. nov.
Figures 5-6
Diagnosis: A small slender species that differs from other west American
species in having finely cancellate sculpture and a straight columella.
Description of holotype: Size small, slender; color shiny pale cream with
weak fulvous blotches; whorls almost flat, 1 1 in number plus four purple-beige
convex nuclear whorls; suture fairly well defined; barely evident subsutural
band of beading, slightly more prominent than beading on remainder of whorl;
sculpture finely cancellate, remarkably consistent from second postnuclear
whorl through body whorl, consisting of axial ribs (29 on penultimate whorl)
crossed by cords (four on penultimate whorl) giving a beaded effect; axial ribs
about equal to interspaces; body whorl of medium length with sculpture con-
tinuing anterior to periphery but less well defined; aperture elongate and
slender; columella straight with no plication; faint siphonal fasciole with
posterior keel; anterior canal short, recurved. Length, 13.9 mm; diameter,
3.3 mm.
Type locality: North of Santa Maria (Charles) Island, Galapagos Islands,
Ecuador, 0° 59' S, 90° 25' W, 70-80 fms., sand and rock bottom, AHF station
195-34, January 29, 1934, 2 specimens.
Type material: Holotype, LACM-AHF 1182; 1 paratype LACM-AHF
1183. An additional paratype, CAS 13221, is from Post Office Bay, of the
same island, 8-10 fms., sand and rock and algae bottom.
Referred material: LACM, Hill coll. 1363, Costa Rica, 2 specimens;
AHF 201-34, Panama, 4 specimens; USNM 192963, Panama, 4 specimens.
Discussion: The sculpture of this species resembles that of Terebra shyana
Bratcher and Burch, 1970, but T. purdyae is a smaller species having a nucleus
of four whorls and a straight columella with no plication. Terebra shyana has
a nucleus of three whorls and a slightly curved columella with a faint plication.
Terebra panamensis Dali, 1908, also has cancellate sculpture, but it is a
heavier, broader shell with coarse sculpture.
Terebra purdyae is named for Ruth Purdy of San Diego, California, in
recognition of her generosity in sharing specimens, not only with us in our
work on Terebridae, but with those studying other families of mollusks.
Terebm stohleri, sp. nov.
Figures 7-8
Diagnosis: A sturdy, medium-small, ivory colored species with little
resemblance to other species of Terebra.
6
Contributions in Science
Description of holotype: Size medium-small; color shiny ivory with
slightly darker blotches; whorls convex, 14 in number plus remaining 2.5 (part
missing) glassy convex nuclear whorls; suture deeply channeled, constricted;
subsutural band inconspicuous; whorls slightly shouldered anterior to suture;
sculpture of slightly curved axial ribs (28 on penultimate whorl), about equal
to interspaces and of evenly spaced spiral grooves (three including subsutural
groove), the two whorls posterior to body whorl having an additional groove
immediately posterior to suture; body whorl of medium length; sculpture
posterior to periphery remains constant; anterior to periphery axial ribs con-
tinue and spiral grooves become more numerous and close set (seven between
periphery and siphonal fasciole); aperture elongate; outer lip thin with sculp-
ture pattern showing through; columella straight with no plication; siphonal
fasciole striate; anterior canal short, recurved. Length, 21.4 mm; diameter,
5.1 mm.
Type locality: Braithwaite Bay, Socorro Island, Mexico, 18° 42' 20" N,
110° 56' 15" W, sand and red mud bottom, March 17, 1939, AHF station
922-39, 1 specimen.
Type material: Holotype, LACM-AHF 1180; 1 paratype, LACM-AHF
1181. The paratype is from Cape Rule, Socorro Island, AHF station 291-34,
4-10 fms.
Referred material: LACM A. 5498, Galapagos Islands, 1 specimen;
LACM, Hill coll. 1365, Chamela Bay, Mexico, 2 specimens; LACM A. 375,
Tres Marias Islands, Mexico, 1 specimen; Shy collection, Manzanillo, Mexico,
1 specimen.
Discussion: There is no other species with which this beautifully sculp-
tured, shiny species can be easily confused.
This species is named in honor of Dr. Rudolph Stohler who has given so
much of his time, his personal finances, and himself to the publishing of The
Veliger, for the advancement of conchology and malacology.
Resumen
Se describen cuatro nuevos terebridos procedentes de las Islas del Pacifico
Este: T ere bra hertleini, T. jacquelinae y T. purdyae procedentes de las Islas
Galapagos, Ecuador, y T. stohleri de la Isla Socorro, Mexico.
Literature Cited
Bratcher, Twila, and R. D. Burch. 1970. Five new species of Terebra from the
Eastern Pacific. Veliger 12(3) :295-300, pi. 14.
Dale, W. H. 1908. Reports on the dredging operations off the west coast of Mexico,
and in the Gulf of California. . . . XIV. The Mollusca and the Brachiopoda.
Mus. Comp. Zool. Harvard, Bull. 43(6): 205-487, pis. 1-22.
Say, Thomas. 1827. Descriptions of marine shells recently discovered on the coast
of the United States. Acad. Nat. Sci. Philad., Journ., ser. 1, 5:207-208.
Accepted for publication December 30, 1969
LOS
ANGELES
COUNTY
MUSEUM
CONTRIBUTIONS
IN SCIENCE
JMBER 189
May 4, 1970
REINSTATEMENT OF THE TURRID GENUS BELLASPIRA
CONRAD, 1868 (MOLLUSCA: GASTROPODA)
WITH A REVIEW OF THE KNOWN SPECIES
By James H. McLean and Leroy H. Poorman
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 sep-
arately, and numbers run consecutively regardless of subject matter. Number 1 was
issued January 23, 1957. The series is available to scientific institutions and scien-
tists on an exchange basis. Copies may also be purchased at a nominal price. Inquiries
should be directed to Virginia D. Miller, Los Angeles County Museum of Natural
History, 900 Exposition Boulevard, Los Angeles, California 90007.
Robert J. Lavenberg
Managing Editor
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Virginia D. Miller
Editor
REINSTATEMENT OF THE TURRID GENUS BELLASPIRA
CONRAD, 1868 (MOLLUSC A: GASTROPODA)
WITH A REVIEW OF THE KNOWN SPECIES
By James H. McLean^ and Leroy H. Poorman-
Abstract: The clavinid turrid genus Bellas pira Conrad is
redefined and its type species, from the Virginia Miocene, is
illustrated. Six known living species of the genus are treated. Two
are from the Caribbean and four are from the eastern Pacific.
Three new species are described: B. acclivicosta, from Guaymas
in the Gulf of California; B. clarionensis, from Clarion Island,
Mexico; and B. margaritensis from Venezuela.
The genus Bellaspira Conrad, 1868, type species Mangelia virginiana
Conrad, 1862, from the Virginia Miocene, had served a need for a small group
of American turrids showing rounded axial ribs, fine spiral sculpture and a
closely constricted anal canal, not deeply notched as in most other clavinid
genera. Conrad’s original description of the genus was adequate and his illus-
tration of the type species, although small, showed the diagnostic features in
suflficient detail to allow Dali (1919) to associate with it a Recent west Ameri-
can species. Unfortunately, a specimen of some other species was mislabeled
as the type of B. virginiana, causing Bartsch and Rehder (1939) to interpret it
incorrectly. As a result, three Recent species, all of which had been associated
with the genus Bellaspira at some time, have been without a satisfactory
generic allocation until now.
Abbreviations for institutions cited in the text are as follows: AHF, Allan
Hancock Foundation (on loan to LACM); AMNH, American Museum of
Natural History, New York; ANSP, Academy of Natural Sciences, Philadel-
phia; CAS, California Academy of Sciences, San Francisco; LACM, Los
Angeles County Museum of Natural History; SDNHM, San Diego Natural
History Museum; SU, Stanford University, California; USNM, United States
National Museum of Natural History.
Acknowledgments
We are greatly indebted to Mrs. Virginia Orr Maes of the Academy of
Natural Sciences, Philadelphia, for locating the type specimen of Bellaspira
virginiana and providing us with photographs and helpful information about
the western Atlantic species. Dr. Joseph Rosewater and Mr. Jack Byas of the
U.S. National Museum kindly located and arranged the loan of specimens in
^Curator of Invertebrate Zoology, Los Angeles County Museum of Natural History.
^Research Assistant, Los Angeles County Museum of Natural History.
1
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Contributions in Science
No. 189
that institution. Radula slides were prepared by Mrs. Ellen Brennan, Museum
research assistant.
Bellaspira Conrad, 1868
Bellaspira Conrad, 1868, p. 261; Dali, 1918, p. 323; Woodring, 1928, p. 162;
Grant and Gale, 1931, p. 584.
Type species, by monotypy, Mangelia virginiana Conrad, 1862, Miocene,
Yorktown, Virginia.
Non Bellaspira Conrad. Bartsch and Rehder, 1939, p. 133; Gardner, 1948, p.
271; Powell, 1966, p. 103.
Conrad (1868) described the genus as follows: “Subfusiform, longi-
tudinally ribbed; canal very short, labrum obsoletely and widely notched,
curved outwards, simple; labium reflexed, entire; columella straight.” With
further reference to the type species, he added, “It is a beautiful and graceful
shell with distant ribs and very minute, close revolving lines, which become
gradually more distinct on the body whorl. The labium is slightly callous near
the upper extremity.”
The shell figured as the type of Mangelia virginiana by Bartsch and
Rehder (1939), copied by Gardner (1948), and Powell (1966), in no way
agrees with Conrad’s description or illustration, and as Powell noted, the
afiinity of the specimen is “with the mangelias or eucitharids,” in the subfamily
Mangeliinae rather than the Clavinae.
Mrs. Virginia Maes has found that both the specimen figured by Bartsch
and Rehder, ANSP 1610, and another specimen, ANSP 30737, are labeled
"‘Mangelia virginiana'' in Conrad’s handwriting. There is no doubt that the
latter specimen is the one figured by Conrad in 1868. There may have been
another original specimen in lot 1610 which was subsequently lost. In order to
avoid further confusion, the only extant specimen, ANSP 30737, is here con-
sidered the holotype. This specimen has seven postnuclear whorls, not five as
stated in the original description.
Grant and Gale (1931) included two other genera in the synonymy of
Bellaspira, the European Atoma Bellardi, 1875, and Haedropleura Bucquoy,
Dautzenberg, and Dollfus, 1883, but these taxa, as treated by Powell, 1966,
lack the features of Bellaspira. Bellaspira is therefore strictly an American
genus, with Recent species in the Caribbean, Panamic, and Californian
provinces.
Radulae of two west American species, B. melea and B. acclivicosta, new
species, have been examined (Figs. 1,2). The radula is of the prototypic type
described by Powell (1966, p. 70) as occurring in many of the strongly ribbed,
glossy shelled genera of Clavininae. The radula consists of a vestigial rachi-
dian, comblike laterals and long, narrow marginals.
1970
Reinstatement of the Genus BELLASPIRA
3
Figures 1-2. Radulae of Bellaspira, greatly enlarged. 1. B. acclivicosta sp. nov., Sola-
dita Cove, Guaymas, Mexico. 2. B. melea Dali, San Carlos Bay, Guaymas, Mexico.
A new generic diagnosis, taking the features of the living species into
account, follows:
Shell 7-19 mm in height, solid, spire elevated; outline concave below
the suture, convex above; nuclear whorls two, smooth; axial sculpture of low
rounded ribs, spiral sculpture of fine incisions or threads; anterior canal broad,
only slightly notched, posterior notch constricted, mature specimens with a
small deposition of callus at top of inner lip, immature specimens lacking a
turrid sinus; lip of mature shell not greatly thickened, but penultimate axial
rib thickened, producing a swelling as in other clavinid genera; color white to
pink, some with darker spiral banding or markings on the axial ribs; oper-
culum leaf-shaped, nucleus terminal; radula prototypic.
Bellaspira virginiana (Conrad, 1862)
Fig. 3
Mangelia virginiana Conrad, 1862, p. 286.
Bellaspira virginiana, Conrad, 1868, p. 261, pi. 21, fig. 12.
Non Bellaspira virginiana, Bartsch and Rehder, 1939, p. 133, pi. 17, fig. 6;
Gardner, 1948, p. 271, pi. 37, fig. 14; Powell, 1966, p. 103, pi. 16, fig. 6.
Description: Postnuclear whorls seven, axial ribs eight, continuous from
whorl to whorl, spiral striae well spaced; height, 14 mm.
Type locality: Miocene of Yorktown, Virginia.
Type material: Holotype (here regarded as discussed above): ANSP
30737 (Fig. 3).
Discussion : We have no other information on the occurrence of the type
species. Additional material is not represented in the Philadelphia Academy.
4
Contributions in Science
No. 189
Figures 3-5. 3. Bellaspira virginiana (Conrad). Holotype, ANSP 30737. X 4. 4. B.
pentagonalis (Dali). Holotype, USNM 9300. X 8. 5. B. pentepleiira Schwengel
( = B. pentagonalis) . Holotype, ANSP 176450. X 5.
Bellaspira pentagonalis (Dali, 1889)
Figs. 4, 5
Drillia pentagonalis Dali, 1889, p. 90.
Bellaspira (?) pentepleura Schwengel, 1940, p. 51, pi. 3, figs. 3, 3a; Powell,
1966, p. 104.
Description: Postnuclear whorls six, axial ribs five, continuous from
whorl to whorl, color white with a peripheral row of squarish chestnut spots
on the axial ribs, spiral striae weakly developed; height, 8-10 mm.
Type material and type localities: Holotype, Drillia pentagonalis, USNM
9300 (Fig. 4); 1 paratype, USNM 679297. Type locality: USFC sta. 2596, 17
miles east of Cape Hatteras, North Carolina, 49 fathoms. Holotype, Bellaspira
pentapleura, ANSP 176450 (Fig. 5) ; 3 paratypes, ANSP 315786. Type local-
ity: Off Palm Beach Florida, 45 fathoms, T. L. McGinty.
Distribution: Cape Hatteras, North Carolina, to Key West and Saint
Petersburg, Florida, 20-60 fathoms. Approximately 80 lots are represented in
the National Museum collection.
Discussion: Dali’s early name Drillia pentagonalis has been overlooked,
evidently because the type was never illustrated. The holotype is worn and
faded, measuring 6.9 mm in height. This species has only five axial ribs and
thereby differs from all the other species having more numerous ribs.
1970
Reinstatement of the Genus BELLASPIRA
5
Bellaspira margaritensis sp. nov.
Fig. 6
Diagnosis: A relatively large species, having more numerous axial ribs
and more deeply incised spiral striae than B. pentagonalis.
Description of holotype: Shell relatively large for the genus; nucleus of
two smooth, rounded whorls; postnuclear whorls eight, axial ribs seven, con-
tinuous from whorl to whorl; suture simple, slightly undulating; axial ribs
evenly rounded, no subsutural channel; spiral sculpture of well defined, broad-
ly spaced spiral incisions, more crowded in the subsutural area, approximately
12 on the penultimate whorl. Color whitish except for a broad pinkish area
about the periphery, weakly demarked and not extending over the rib surfaces.
Parietal callus well defined, terminating in a small node in front of the anal
sulcus. Edge of outer lip thin, with a shallow stromboid notch near the base, a
minor axial rib slightly behind the edge; the penultimate rib more strongly
developed than the others. Dimensions (in mm) : height 15.0, diameter 5.5.
Type locality: Four to seven miles north of Margarita Island, Venezuela,
11° 08' N, 64° 17' W, 17-21 fathoms on sand, R/V Velero III Station A42-39,
21 April 1939, four specimens, two live-taken.
Type material: Holotype, LACM-AHF 1337 (Fig. 6); two paratypes,
LACM-AHF 1338; one paratype, USNM 675964.
Figures 6-8. 6. Bellaspira margaritensis sp. nov. Holotype, LACM-AHF 1337. X 4.2.
7. B. melea Dali. Holotype, USNM 15952. X 4.5. 8. B. melea Dali. Off San Carlos
Bay, Guaymas, Mexico. X 5.
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Contributions in Science
No. 189
Discussion: The only other species with roundly arched axial ribs is
B. pentagonalis, from which B. margaritensis differs in having two additional
axial ribs, more prominent and broadly spaced incised spiral sculpture, and a
color band not developed across the rib surface.
Beilaspira melea Dali, 1919
Figs. 7, 8
Beilaspira melea Dali, 1919, p. 29, pi. 19, fig. 8; Grant and Gale, 1931, p. 584.
Elaeocyma melea, Pilsbry and Lowe, 1932, p. 1 10.
Clavus {ICymatosyrinx) melea. Keen, 1958, p. 448, fig. 736.
Description: Postnuclear whorls 7-8, axial ribs 6-8, continuous from
whorl to whorl, first rib behind the lip greatly thickened; color whitish to flesh
pink, with traces of darker banding, rib surfaces lighter in color; height,
13-19 mm.
Type locality: Panama, collected by Col. E. Jewett.
Type material : Holotype, USNM 15952 (Fig. 7) .
Distribution: Tepoca Bay, Sonora, Mexico, to Port Utria, Colombia,
10-40 fathoms. Twenty-two lots are represented in the LACM collection.
Intermediate stations in Mexico include Concepcion Bay, Baja California;
Guaymas, Sonora; Cleopha Island, Tres Marias Islands; Sihauntanejo, Guer-
rero; Guatulco Bay, Oaxaca; and in Costa Rica, Port Culebra; indicating that
the species is widely distributed throughout its range.
Remarks: Dali (1919) correctly assigned this species to Beilaspira, not-
ing that it is “typical Beilaspira” It so resembles B. virginiana that it would
undoubtedly be considered conspecific but for the discrepancies of age and
distribution. The type specimen is a worn shell not showing spiral sculpture,
but Dali observed that spiral sculpture could have been worn away. In fresh
specimens (Fig. 8) the spiral sculpture is extremely minute and somewhat
undulating. Beilaspira melea varies considerably in color. Some specimens are
white and others have pink or flesh colored banding, the darkest band located
at the periphery.
Beilaspira acclivicosta sp. nov.
Figs. 9-11
Diagnosis: A relatively large species with a characteristic color pattern,
having more numerous and more strongly crested axial ribs than B. melea.
Description of holotype: Shell large for the genus, pinkish white to flesh
colored; nucleus of two smooth, rounded, glassy pink whorls; postnuclear
whorls nine, axial ribs 10, not continuous from whorl to whorl; suture simple,
slightly undulating at the base of the ribs; ribs weak on the shoulder, strong
and nodulous at the periphery, becoming obsolete on the base. Entire shell
1970
Reinstatement of the Genus BELLASPIRA
1
Figures 9-11. All Bellaspira acclivicosta sp. nov. 9. Holotype, LACM 1193. X 3.
10. Off Isla Candelero, Guaymas, Mexico, Poorman collection. X 3. 11. Pulmo Bay,
Baja California, Mexico, LACM 66-19. X 5.5.
covered by microscopic, raised spiral threads, most numerous above the
periphery and strongest on the base, crossed by incremental growth threads to
form a minutely reticulated surface. Peripheral crests of ribs white, surrounded
by red-brown, strongest above and below; a second and smaller row of dots on
the ribs between the periphery and the anterior tip. Columella straight, parallel
to axis of shell, twisting slightly to the left at the base. The parietal callus is
well defined and terminates in a domelike pad in front of the anal sulcus. Outer
lip thin, smooth within, nearly parallel to the columella, showing a shallow
stromboid notch near the base and thickened by the final rib, which is pre-
ceded by an unusually wide valley and the penultimate rib that is stronger
than all the others. Periostracum lacking; operculum with apical nucleus.
Dimensions (in mm) : height 18.1, diameter 6.8.
Type locality: One kilometer south of the east point at the entrance to
Bahia San Carlos, Guaymas, Sonora, Mexico; 21° 56' N, 111° 03' W; 15-20
fathoms on rock and broken shell bottom, December, 1965, to April, 1967,
Forrest and Roy Poorman, ten specimens.
Type material: Holotype, LACM 1193 (Fig. 9); one paratype, ANSP
315788, one paratype, AMNH 147999; one paratype, CAS 13159; one para-
type, SDNHM 50000; one paratype, SU 9964; one paratype; USNM 679183;
three paratypes, Poorman collection.
Referred material: Poorman collection, two specimens dredged at Isla
Candelero, Guaymas (Fig. 10); LACM 68-27, four specimens, diving at 60
feet, Soladita Cove, Guaymas; LACM 66-19, one specimen, diving at 5-20
8
Contributions in Science
No. 189
feet, Pulmo Bay, Baja California (Fig. 11); Shy collection (Westminster,
California) , one specimen dredged at Manzanillo, Colima, Mexico.
Discussion: Immature shells of B. acclivicosta do not show a turrid sinus
and lack the callus deposition defining the anal sinus. Figure 10 shows a speci-
men slightly larger and broader than the holotype that has an immature lip.
This specimen also is more distinctly colored than the holotype. The specimen
from Pulmo Bay (Fig. 1 1 ) has six postnuclear whorls and measures 9.1 mm
in height. It is evidently a miniature specimen of B. acclivicosta having a ma-
ture lip. It is similar to the type lot except for size.
The new species differs from B. melea in having about ten discontinuous
axial ribs rather than seven continuous ribs per whorl. The spiral sculpture
consists of fine raised spiral threads on B. acclivicosta, whereas the weak
grooves on B. melea appear to be scratched into the surface.
The name is derived from the Latin nouns acclivis, steep, and costa, rib,
suggestive of the steep, concave slope of the ribs below the suture. The
specific name is intended as a noun in apposition.
Bellaspira grippi (Dali, 1908)
Figs. 12-15
Bela grippi Dali, 1908, p. 137.
Lora grip pH, Dali, 1921, p. 74, pi. 3, fig. 8; Oldroyd, 1927, p. 100, pi. 7, fig. 6.
Figures 12-14. All Bellaspira grippi (Dali). 12. Holotype, USNM 200653. X 5.7.
13. San Martin Island, Baja California, Mexico, LACM 67-50. X 6.4. 14. Asuncion
Island, Baja California, LACM 67-66. X 7.4.
1970
Reinstatement of the Genus BELLASPIRA
9
Bellaspira grippi, Grant and Gale, 1931, pp. 514, 565; Burch, 1946, p. 19
rgrippiV’].
Description: Relatively small, postnuclear whorls six, subsutural area
markedly concave; axial ribs 12-14, not continuous from whorl to whorl, ribs
obsolete at the base and often over the final whorl; color white to flesh pink,
some with dark and light banding; height, 7-1 1 mm.
Type locality: Five miles south of the entrance to San Diego Bay, 15
fathoms, C. W. Gripp, collector.
Type material: Holotype, USNM 200653 (Fig. 12), one paratype,
USNM 697389.
Distribution: Redondo Beach, California, to Asuncion Island, Baja Cali-
fornia; Guadalupe Island, Mexico. Not uncommon on rocky bottoms under
kelp and dredged on gravel bottoms to depths of 25 fathoms.
Discussion : Grant and Gale (1931 ) were the first to associate this species
with the genus Bellaspira, a relationship missed by Dali, who had correctly
assigned his B. melea in 1919.
A wide range of variation is shown in this species. The typical southern
Californian form, represented by the holotype (Fig. 12) and a specimen from
San Martin Island, Baja California (Fig. 13) , is whitish or flesh colored and
usually shows a slightly darker band in the concave area of the shoulder. One
specimen from Asuncion Island, outer coast of Baja California (Fig. 14), 6.9
mm in height, is dark rose colored, except on the siphonal fasciole, and the
shoulder is more strongly tabulate, the axial ribs forming projecting nodes.
Incised spiral sculpture is unevenly spaced on the body whorl. The largest of
five specimens from Guadalupe Island (Fig. 15) is 7.5 mm in height. All are
strongly tabulate and the spiral incisions are evenly spaced; two are dark
flesh-colored and one is white.
Bellaspira clarionensis sp. nov.
Fig. 16
Diagnosis: A medium sized species larger than B. grippi and having more
numerous spiral striae.
Description of holotype: Shell of medium size for the genus, nuclear
whorls two, rounded; postnuclear whorls seven, axial ribs 10, alternating in
position on early whorls, but aligned from whorl to whorl on the final three
whorls. Suture simple, slightly undulating; shoulder concave; axial ribs well
defined across the shoulder, cresting at the periphery, extending across the
body whorl and becoming obsolete on the base. Spiral sculpture of closely
spaced spiral striae, more crowded upon the shoulder and overriding the axial
ribs. Color whitish with a narrow band of flesh pink upon the shoulder and
another uninterrupted narrow band across the base. Parietal callus projecting,
anal sinus narrow. Lip edge thin, stromboid notch barely perceptible, the
10
Contributions in Science
No. 189
Figures 15-16. 15. Bellaspira grippi (Dali). Guadalupe Island, Mexico, LACM 65-42.
X 9.5. 16. B. clarionensis sp. nov. Holotype, LACM-AHF 1366. X 4.6.
second and third ribs behind the aperture coalesced and thickened. Dimensions
(in mm) : height 13.5, diameter 5.9 mm.
Type locality: Off Sulphur Bay, Clarion Island, Revillagigedo Islands,
Mexico, 18° 20' N, 114° 44' W, 28-45 fathoms on sand, R/V Velero III
Station 917-39, 16 March 1939, one specimen. Two additional paratype
specimens were dredged at stations 136-34 and 921-39, Sulphur Bay, Clarion
Island, 30-35 fathoms.
Type material: Holotype, LACM-AHF 1366 (Fig. 16), one paratype,
LACM-AHF 1367; one paratype, USNM 675565.
Referred material: LACM A. 3 75, five specimens, dredged by George
Willett, 20-35 fathoms. Clarion Island, 24 March 1938.
Discussion: Bellaspira clarionensis differs from B. grippi in reaching a
larger size, having a narrower shoulder area, elongate rather than nodular axial
ribs, and more numerous spiral striae.
Resumen
El genero clavinido turrido Bellaspira Conrad se redifine y la especie tipo,
del Mioceno de Virginia, se ilustra. Seis especies vivientes conocidas en este
genero son aqui tratadas. Dos de ellas son del Caribe y cuatro del Pacifico
1970
Reinstatement of the Genus BELLASPIRA
11
Este. Se describen tres nuevas especies: B. acclivicosta, de Guaymas en el
Golfo de California; B. clarionensis, de la Isla Clarion, Mexico; y B. marga-
ritensis de Venezuela.
Literature Cited
Bartsch, P., and H. a. Rehder. 1939. New turritid mollusks from Florida. U.S. Nat.
Mus. Proc., 86:127-138.
Burch, J. Q. (ed.). 1946. Distributional list of the West American Marine Mollusks
from San Diego, California, to the Polar Sea. [Extracts from the] Minutes,
Conch. Club So. Calif., 62: 1-52.
Conrad, T. A. 1862. Descriptions of new genera, subgenera and species of Tertiary
and Recent shells. Acad. Nat. Sci. Phila., Proc., 14: 284-291.
. 1868. Descriptions of new genera and species of Miocene shells, with notes
on other fossil and Recent species. Amer. Jour. Conchology, 3: 257-270.
Dale, W. H. 1889. Reports on the results of dredging, under the supervision of
Alexander Agassiz, in the Gulf of Mexico (1877-78) and in the Caribbean Sea
(1879-80), by the U.S. Coast Survey steamer “Blake.” XXIV. Report on the
Mollusca. — Part II. Gastropoda and Scaphopoda. Mus. Comp. Zook, Bull.,
18: 1-492.
. 1908. Some new Californian shells. Nautilus, 21: 136-137.
. 1918. Notes on the nomenclature of the mollusks of the family Turritidae.
U.S. Nat. Mus., Proc., 54: 313-333.
. 1919. Descriptions of new species of mollusks of the family Turritidae from
the west coast of America and adjacent regions. U.S. Nat. Mus., Proc., 56: 1-86.
. 1921. Summary of the marine shellbearing 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. U.S. Nat. Mus., Bull., 112: 1-217.
Gardner, J. 1948. Mollusca from the Miocene and Lower Pliocene of Virginia and
North Carolina, Part. 2. Scaphopoda and Gastropoda. U.S. Geol. Surv., Prof.
Paper, 199B: 177-310.
Grant, U. S., IV, and H. R. Gale. 1931. Catalogue of the marine Pliocene and
Pleistocene Mollusca of California and adjacent regions. San Diego Soc. Nat.
Hist., Mem., 1: 1-1036.
Keen, A. M. 1958. Sea shells of tropical west America; marine mollusks from Lower
California to Colombia. Stanford Univ. Press, Stanford, California, xi -f 624 p.
Oldroyd, I. S. 1927. The marine shells of the West Coast of North America. 2,
Gastropoda, Scaphopoda, and Amphineura. Stanford Univ. Publ. Univ. Ser.,
Geol. Sci., 2(1): 1-297.
PiLSBRY, H. A., AND H. N. LowE. 1932. West Mexican and Central American mol-
lusks collected by H. N. Lowe, 1929-31. Acad. Nat. Sci., Phila., Proc., 84:
33-144.
Powell, A. W. B. 1966. The molluscan families Speightiidae and Turridae. Bull.
Auckland Inst, and Mus. 5: 1-184.
SCHWENGEL, J. S. 1940. New Mollusca from Florida. Nautilus, 54: 49-52.
Woodring, W. P. 1928. Contributions to the geology and paleontology of the West
Indies. Miocene mollusks from Bowden, Jamaica. Part II. Gastropods and
discussion of the results. Carnegie Inst. Wash., Publ. no. 385: 1-564.
Accepted for publication February 28, 1969
LOS
ANGELES
COUNTY
MUSEUM
CONTRIBUTIONS
ZYL, IN SCIENCE
UMBER 190
May 20, 1970
A NEW SPECIES OF FOSSIL KELLETIA (MOLLUSCA:
GASTROPODA) FROM THE LOMITA MARL,
LATE CENOZOIC OF SAN PEDRO, CALIFORNIA
By Leo G. Hertlein
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 sep-
arately, and numbers run consecutively regardless of subject matter. Number 1 was
issued January 23, 1957. The series is available to scientific institutions and scien-
tists on an exchange basis. Copies may also be purchased at a nominal price. Inquiries
should be directed to Virginia D. Miller, Los Angeles County Museum of Natural
History, 900 Exposition Boulevard, Los Angeles, California 90007.
Robert J. Lavenberg
Managing Editor
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. Al-
though priority will be given to manuscripts by staff members, or to papers dealing
largely with specimens in the Museum’s collections, other technical papers will be
considered. All manuscripts must be recommended for consideration by the curator
in charge of the proper section or by the editorial board. Manuscripts must conform
to those specifications listed below and will be examined for suitability by an Edi-
torial Committee. They may also be subject to review by competent specialists out-
side the Museum.
Authors proposing new taxa in a CONTRIBUTIONS IN SCIENCE must
indicate that the primary type has become the property of a scientific institution of
their choice and cited by name.
MANUSCRIPT FORM.— (1) The 1964 AIBS Style Manual for Biological
Journals is to be followed in preparation of copy. (2) Double space entire manu-
script. (3) Footnotes should be avoided if possible. Acknowledgments as footnotes
will not be accepted. (4) Place all tables on separate pages. (5) Figure legends and
unavoidable footnotes must be typed on separate sheets. Several of one kind may
be placed on a sheet. (6) An abstract must be included for all papers. This will be
published at the head of each paper. (7) A Spanish summary is required for all
manuscripts dealing with Latin American subjects. Summaries in other languages
are not required but are strongly recommended. Summaries will be published at
the end of the paper. (8) A diagnosis must accompany any newly proposed taxon.
(9) Submit two copies of manuscript.
ILLUSTRATIONS. — All illustrations, including maps and photographs, will
be referred to as figures. All illustrations should be of sufficient clarity and in the
proper proportions for reduction to CONTRIBUTIONS page size. Consult the 1964
AIBS Style Manual for Biological Journals in preparing illustration and legend copy
for style. Submit only illustrations made with permanent ink and glossy photo-
graphic prints of good contrast. Original illustrations and art work will be returned
after the manuscript has been published.
PROOF. — Authors will be sent galley proof which should be corrected and
returned promptly. Changes in the manuscript after galley proof will be billed to
the author. Unless otherwise requested, page proof also will be sent to the author.
One hundred copies of each paper will be given free to each author or divided
equally among multiple authors. Orders for additional copies must be sent to the
Editor at the time corrected galley proof is returned. Appropriate order forms will
be included with the galley proof.
Virginia D. Miller
Editor
A NEW SPECIES OF FOSSIL KELLETIA (MOLLUSCA:
GASTROPODA) FROM THE LOMITA MARL,
LATE CENOZOIC OF SAN PEDRO, CALIFORNIA
By Leo G. Hertlein^
Abstract: Kelletia kanakoffi, a new species of gastropod
belonging to the family Neptuneidae, is described from strata of
Late Pliocene or Early Pleistocene age in San Pedro, California.
Introduction
Numerous species of fossil mollusks have been reported from Hilltop
Quarry and nearby localities (see Hanna, 1923; Schenck, 1945: 513-514;
Woodring et ai, 1946: 49-52; Valentine, 1961: 411-413) in San Pedro, Los
Angeles County, California. Intensive collecting in this area by George P.
Kanakoff and his associates has yielded many interesting fossils now preserved
in the Los Angeles County Museum of Natural History.
Recently Mr. Kanakoff sent the author two specimens of a large gastro-
pod, referable to the genus Kelletia. Study of these fossils revealed that they
represent an undescribed species here described as new.
The strata in which they were found have been considered to be of Late
Pliocene age by some authors but as Early Pleistocene age by others. An age
of three million years given by Obradovich (1965), based upon Potassium-
Argon dating of glauconite, represents the most recent maximum age attributed
to these sediments. This date would certainly place the Lomita Marl, a mem-
ber of the lower San Pedro formation, in the Late Pliocene or very Early
Pleistocene. Several species of mollusks reported from the Lomita Marl occur
elsewhere only in strata believed by some authors to be of Pliocene age.
Acknowledgments
I am grateful to Mr. George P. Kanakoff, Curator Emeritus of the
Section of Invertebrate Paleontology, Los Angeles County Museum of Natural
History, who lent the specimens of the new species described herein for study
and description. Acknowledgment also is due Victor Zullo, Allyn G. Smith
and Barry Roth, California Academy of Sciences, whose advice and criticism
was particularly helpful during the preparation of the manuscript. Alan J.
Galloway of the same institution, and Warren O. Addicott, United States
Geological Survey, called my attention to certain literature concerning the
geochronology of the Lomita Marl. The illustrations in the text are from photo-
graphs made by Maurice Giles, Staff Photographer, California Academy of
Sciences.
^Curator (Emer.) of Invertebrate Paleontology, California Academy of Sciences;
and Research Associate, Los Angeles County Museum of Natural History.
1
2
Contributions in Science
No. 190
Class Gastropoda
Family Neptuneidae Tryon
Genus Kelletia Bayle, in Fischer
Kelletia Bayle, in Fischer, 1884: 625. Sole species, “5. Kelleti, Forbes.”
Wenz, 1938 (reprint, 1962): 1166
Type species: Siphonalia kelleti Forbes [=Fusus kelletii Forbes (1852:
274, pi. 9, fig. 10)] “taken on the Californian coast.”
This genus (and its type species) was named for Captain Kellett of the
British Royal Navy. Some authors have used the orthography Kellettia but
most recent authors have followed the original spelling given by Bayle for the
genus and kelletii for the species as given by Forbes.
West American authors have placed Kelletia in either the family Nep-
tuneidae or the Buccinidae. Powell ( 1929: 63) stated that the shell characters
as well as the radula of this genus are very similar to Austrosipho Cossmann,
and these genera he placed in the family Buccinulidae Finlay. Wenz ( 1962:
1151) placed both that family and the Neptuneidae as subfamilies under the
Buccinidae. He considered Austrosipho Cossmann, 1906, and Verconella
Iredale, 1914, to be synonyms of Penion Fischer, 1884.
Ruth ( 1942), in a monograph of Siphonalia in the Cenozoic of western
North America, considered Kelletia a subgenus which included four species
that he described and illustrated. The geologic range of the subgenus indicated
by the included species was from Paleocene (Martinez) to Recent. Sub-
sequent to Ruth’s paper, one species, Kelletia vladimiri Kanakoff, was described
from strata of Pliocene age in southern California. One species of Kelletia
now lives in the region between Santa Barbara, California, and San Quintin,
Baja California, from the intertidal zone to a depth of about 64 meters (35
fathoms). Another species, Kelletia lischkei Kuroda, lives in Japanese waters
where it has been reported to range from 32° to 39° North Latitude on the
Pacific coast and to 40° North on the Japan Sea coast.
In their catalogue of the Tertiary mollusks of Japan, 1952, Hatai and
Nisiyama did not allocate any species to Kelletia but species assigned to Kel-
letia (considered to be a subgenus of Siphonalia) have been reported by others
from strata of Late Tertiary age in Indonesia and in India.
Two species of Pliocene age from Ecuador, tentatively assigned to Kel-
letia by Olsson ( 1964: K. ecuadoriana, p. 165, pi. 24, figs. 2-2c and K. rugosa,
p. 165, pi. 24, figs. 1, la), do not closely resemble species of that genus
described from California.
Anderson ( 1964: 249) recently proposed a subgenus Boreokelletia, based
upon Kelletia (Boreokelletia) hosiusi (Beyrich, 1856), from strata of Miocene
age in Germany. The type species earlier was referred to the genus Strepto-
chetus Cossman, 1889, by Kautsky (1925: 1 17), and others.
1970
A New Species of Fossil Kelletia
3
Kelletia kanakoffi, new species
Figures 1 , 2, and 3
Diagnosis: A new species of Kelletia which differs from other west Ameri-
can species of this genus in its large size, thick shell, coarse spiral sculpture
and large umbilicus.
Description: Shell large, thick, fusiform, the spire elevated; whorls of spire
five (the apex lacking), angulated near the middle, posteriorly slightly con-
cave and extending over a little more than one half of the preceding whorl, the
suture appressed; axial sculpture consisting of well developed, axially elon-
gated nodes, seven on the uppermost, eight on the succeeding one, ten on the
penultimate whorl and eight on the body whorl, followed by a low, carinate
ridge at an angulation posterior to the middle, with vague vertical ridges out-
lined by lines of growth; the entire shell is sculptured with spiral ridges sepa-
rated by deep grooves, about twenty on the penultimate whorl and about
twenty-three on the body whorl, the anterior ones much coarser than the
posterior ones; some ridges bear one to three spiral grooves and where eroded
these separate small riblets; aperture ovate, outer lip not thickened, the inte-
rior lightly lirate corresponding to the exterior sculpture, pillar smooth, twisted,
inner lip smooth; canal well developed, narrow, twisted and recurved, a large
siphonal fasciole present, in front of which is a large, rather wide umbilicus.
Dimensions: The holotype, length (apex lacking), 181 mm, maximum
diameter, 90 mm, length of aperture and canal, 102 mm; paratype, length,
178 mm, maximum diameter, 91 mm, length of aperture and canal, 104 mm.
Type locality: Loc. 435 LACMIP, Lomita Marl about 50 feet below the
crossing of West Park Drive, Host Place and Coralmount Drive, San Pedro,
California. Late Pliocene or Early Pleistocene. John Sutherland collector,
July, 1966.
Type specimens: Holotype Catalog Number 2456, and Paratype Cata-
logue Number 2457, Los Angeles County Museum of Natural History, Inver-
tebrate Paleontology Collection.
Comments: This new species differs from the Recent Kelletia kelletii
(Forbes, 1852: 274, pi. 9, fig. 10) in the larger size, coarser sculpture, very
large umbilicus, the more angulated shoulder and slightly carinate body whorl,
and in that the whorls on the spire overlap the preceding ones to a less extent
posteriorly than do those of the well known living species (see Arnold, 1903:
pi. 4, fig. 5; Ruth, 1942: pi. 48, fig. 5; Abbott: 1954, pi. 24, fig. W).
A large specimen of Kelletia kelletii 161 mm long and 76 mm in maxi-
mum diameter, in the collections of the California Academy of Sciences, from
Locality 31642 (CAS), from 3.7 miles off Huntington Beach, California, in
37 to 38 meters (20 to 21 fathoms), has only a short, narrow, shallow, umbili-
cal groove in comparison to the large umbilicus of K. kanakoffi, new species.
The largest specimen of K. kelletii reported in the literature (Anon., 1959:
4
Contributions in Science
No. 190
Figure 1. Kelletia kanakoffi, new species. Holotype, LACMIP 2456, from Lomita
Marl, San Pedro, California; Late Pliocene or Early Pleistocene age. Approximately
X 5/6; true length (apex lacking), 181 mm, maximum diameter, 90 mm.
1970
A New Species of Fossil Kelletia
5
Figure 2. Kelletia kanakoffi, new species. Dorsal view of holotype shown in figure 1 .
6
Contributions in Science
No. 190
Figure 3. Kelletia kanakoffi, new species. Paratype, LACMIP 2457, from the same
locality as the holotype shown in figure 1. Approximately X 5/6; true length,
178 mm, maximum diameter, 91 mm. Note the large umbilicus on this specimen.
1970
A New Species of Fossil Kelletia
7
19), from southern California, is 170 mm long and 87 mm in maximum
diameter.
The same shell characters which serve to separate Kelletia kanakoffi,
new species, from K. kelletii also differentiate it from Kelletia vladimiri Kana-
koff (1954: 114, pis. 29, 30, 31), a smaller, more slender form described
from the Pico formation, of Pliocene age, in Los Angeles County, California.
This species is named for Mr. George P. Kanakoff, Curator Emeritus,
Los Angeles County Museum of Natural History, whose careful collecting has
brought to light many interesting specimens.
Literature Cited
Anonymous 1959. “Lost Operculum Club.” Min. Conch. Club South. California
(J. Q. Burch, Editor), No. 190: 17-23, July-August.
Abbott, R. T. 1954. American Seashells. (D. Van Nostrand Co., New York), pp.
I-XIV, 1-541, pis. 1-40, figs. 1-100 in text.
Anderson, H. J. 1964. Faunen aus dem Miocan Nordwestdeutschlands. Die Miocane
Reinbek-Stufe in Nord- und Westdeutschland und ihre Mollusken-Fauna. Fort-
schritte in der Geologie von Rheinland und Westfalen. Geologisches Landes-
amt Nordrhein-Westfalen, 14: 31-368, pis. 1-52, figs. 1-18 in text.
Arnold, R. 1903. The Paleontology and Stratigraphy of the Marine Pliocene and
Pleistocene of San Pedro, California. Mem. Calif. Acad. Sci., 3: 1-420, pis.
1-37, June 27.
Fischer, P. 1884. Manuel de Conchyliologie et de Paleontologie Conchyliologique
ou Histoire Naturelle des Mollusques Vivants et Fossiles. (Librairie F. Savy,
Paris), Fasc. 7; 609-688, figs. 370-456 in text, June 30.
Forbes, E. 1852. On the marine mollusca discovered during the voyages of the
Herald and Pandora, by Capt. Kellett, R. N., and Lieut Wood, R. N. Proc.
Zool. Soc. London for 1850: 270-274, pis. 9, 1 1, January 24.
Hanna, G D. 1923. Results of preliminary examination of seven samples of sedi-
ments from near Lomita. Bull. South. Calif. Acad. Sci., 22 (2) : 64-65, July.
(“List of the fossil mollusks found in the Lomita beds, as identified by Mrs.
Ida S. Oldryod. . . .”, p. 65).
Hatai, K., and S. Nisiyama, 1952. Checklist of Japanese Tertiary marine mollusca.
Sci. Repts. Tohoku Univ., Sendai, Japan, Second Ser. (Geology), Special vol.
3: 1-449; supplement, pp. 452-464.
Kanakoff, G. P. 1954. A new Kelletia from the Pliocene of California. Bull. South.
Calif. Acad. Sci., 53 (2): 114-117, pis. 29, 30, 31, May-August (issued
August 1).
Kautsky, F. 1925. Das Miocan von Hemmoor und Basbeck-Osten. Abh. d. Preuss.
Geol. Landesanstalt, Neue Folge, Heft 97, pp. 1-255, pis. 1-12.
8
Contributions in Science
No. 190
Obradovich, J. D. 1965. The Potential use of Glauconite for Late-Cenozoic Geo-
chronology. International Association for Quaternary Research, 7th Congress
(United States), Proc., 8: 267-279.
Olsson, a. a. 1964. Neogene Mollusks from Northwestern Ecuador. Paleo.
Research Inst., Ithaca, New York, pp. 1-256, pis. 1-38, October 28.
Powell, A. W. B. 1929. The Recent and Tertiary species of the genus Buccinulum
in New Zealand, with a review of related genera and families. Trans, and Proc.
New Zealand Inst., 60 (Quarterly issue), Pt. 1 : 57-101, pis. 1-4, figs. 83-132
in text. May 31.
Ruth, J. W. 1942. The molluscan genus Siphonalia of the Pacific coast Tertiary.
Univ. Calif. Publ. Bull. Depart. Geol. Sci., 26(3) : 287-306, pis. 47, 48,
March 24.
SCHENCK, H. G. 1945. Geologic Application of Biometrical Analysis of Molluscan
Assemblages. Jour. Paleo., 19(5) : 504-521, pis. 66, 67, September.
Valentine, J. W. 1961. Paleoecologic Molluscan Geography of the Californian
Pleistocene. Univ. Calif. Publ. Geol. Sci., 34(7) : 309-442, figs. 1-16 in text,
May 17. ( See especially pp. 4 1 1 -4 1 3 ) .
Wenz, W. 1938 (1962). Handbuch der Palaozoologie herausgegeben von O. H.
Schindewolf. Gastropoda von W. Wenz. (Gebriider Borntraeger: Berlin. Foto-
mechanischer Nachdruck, 1962), Bd. 6, Teil 1, Gastropoda, Teil 1, Lieferung
5: 961-1200, Abbildungen 2788-3416.
Woodring, W. P., M. N. Bramlette, and W. S. W. Kew. 1946. Geology and
Paleontology of Palos Verdes Hills, California. U.S. Geol. Surv., Prof. Paper
207: I-V, 1-145, pis. 1-37, figs. 1-16 in text. (See especially “Hilltop quarry
and nearby localities,” pp. 49-52).
Accepted for publication March 16, 1970
i LOS
; ANGELES
1 COUNTY
' MUSEUM
CONTRIBUTIONS
cllsfa IN SCIENCE
llMBER 191
May 20, 1970
STUDIES ON THE FROGS OF THE GENUS LEPTODACTYLUS
(AMPHIBIA: LEPTODACTYLIDAE). VI.
BIOSYSTEMATICS OF THE MELANONOTUS GROUP
By W. Ronald Heyer
ii
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 sep-
arately, and numbers run consecutively regardless of subject matter. Number 1 was
issued January 23, 1957. The series is available to scientific institutions and scien-
tists on an exchange basis. Copies may also be purchased at a nominal price. Inquiries
should be directed to Virginia D. Miller, Los Angeles County Museum of Natural
History, 900 Exposition Boulevard, Los Angeles, California 90007.
Robert J. Lavenberg
Managing Editor
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. Al-
though priority will be given to manuscripts by staff members, or to papers dealing
largely with specimens in the Museum’s collections, other technical papers will be
considered. All manuscripts must be recommended for consideration by the curator
in charge of the proper section or by the editorial board. Manuscripts must conform
to those specifications listed below and will be examined for suitability by an Edi-
torial Committee. They may also be subject to review by competent specialists out-
side the Museum.
Authors proposing new taxa in a CONTRIBUTIONS IN SCIENCE must
indicate that the primary type has become the property of a scientific institution of
their choice and cited by name.
MANUSCRIPT FORM.— (1) The 1964 AIBS Style Manual for Biological
Journals is to be followed in preparation of copy. (2) Double space entire manu-
script. (3) Footnotes should be avoided if possible. Acknowledgments as footnotes
will not be accepted. (4) Place all tables on separate pages. (5) Figure legends and
unavoidable footnotes must be typed on separate sheets. Several of one kind may
be placed on a sheet. (6) An abstract must be included for all papers. This will be
published at the head of each paper. (7) A Spanish summary is required for all
manuscripts dealing with Latin American subjects. Summaries in other languages
are not required but are strongly recommended. Summaries will be published at
the end of the paper. (8) A diagnosis must accompany any newly proposed taxon.
(9) Submit two copies of manuscript.
ILLUSTRATIONS. — All illustrations, including maps and photographs, will
be referred to as figures. All illustrations should be of sufficient clarity and in the
proper proportions for reduction to CONTRIBUTIONS page size. Consult the 1964
AIBS Style Manual for Biological Journals in preparing illustration and legend copy
for style. Submit only illustrations made with permanent ink and glossy photo-
graphic prints of good contrast. Original illustrations and art work will be returned
after the manuscript has been published.
PROOF. — Authors will be sent galley proof which should be corrected and
returned promptly. Changes in the manuscript after galley proof will be billed to
the author. Unless otherwise requested, page proof also will be sent to the author.
One hundred copies of each paper will be given free to each author or divided
equally among multiple authors. Orders for additional copies must be sent to the
Editor at the time corrected galley proof is returned. Appropriate order forms will
be included with the galley proof.
Virginia D. Miller
Editor
STUDIES ON THE FROGS OF THE GENUS LEPTODACTYLUS
(AMPHIBIA: LEPTODACTYLIDAE) . VI.
BIOSYSTEMATICS OF THE MELANONOTUS GROUP
By W. Ronald Heyer^
Abstract: Six species are recognized in the Melanonotus
group: dantasi, discodactyliis, melanonotus, podicipinus, pustu-
latiis, and wagneri. A synonymy, diagnosis, summary of charac-
teristics, and distributional summary are presented for each
species. Series of Leptodactyliis were analyzed from broad geo-
graphic and ecologic areas and situations for standard length,
ventral pattern, ventrolateral gland development, posterior thigh
pattern, and toe disk development. Correlations are evident
between 1 ) larger size and more mesic habitats in wagneri,
2) darker ventral pattern and greater annual rainfall in mela-
nonotus, 3) lighter ventral pattern and greater ventrolateral-
gland extent in wagneri, and 4) greater ventrolateral-gland extent
and greater annual rainfall in wagneri. The hypothesis is pre-
sented that character displacement is involved in the sharp
differences in size and pattern between populations of melanono-
tus and wagneri as well as podicipinus and wagneri in sympatry.
Leptodactyliis melanonotus is the most primitive of the
species, while discodactyliis and piistiilatus are the most ad-
vanced.
Leptodactyliis melanonotus, podicipinus, and wagneri are
associated with old land masses and are species adapted for xeric
conditions. Leptodactyliis dantasi and discodactyliis are limited
to the Tropical Rainforest of the western Amazonian Basin. Lep-
todactyliis piistiilatus is distributed in the xeric regions of eastern
Brasil.
Introduction
Since the frog genus Leptodactylus was proposed by Fitzinger in 1826,
it has had an unstable systematic history. No one since Boulenger, in 1882, has
treated the group as an entity. The present paper is the first of a projected series
attempting to elucidate the biosystematics of each of the species groups within
the genus. In the Melanonotus species group, characters of gross morphology
and distribution are used to analyze the interspecific relationships, ecological
distribution patterns, and geographical distribution patterns of the species.
Methods
All adult specimens of the Melanonotus species group were examined for
size, sex, ventral color pattern, ventrolateral gland development, posterior
1 Research Associate in Herpetology, Los Angeles County Museum of Natural
History; and Biology Department, Pacific Lutheran University, Tacoma, Washing-
ton 98447.
1
2
Contributions in Science
No. 191
thigh pattern, and degree of toe-tip expansion. Analysis of these characters
is the basis for the taxonomic conclusions. For convenience, taxonomic con-
clusions are presented first before discussing the variation of characters an-
alyzed in detail.
Ail available adult individuals of L. dantasi, discodactylus and pustulatus
were studied for variation in 34 characters. The same 34 characters were ex-
amined in a series of 15 males and 15 females for each of the other three
species. Thirty specimens of melanonotus, podicipinus, and wagneri were
chosen to represent the extremes in geographic origin and morphological vari-
ation. Details of the methods of examining individuals are the same as used
previously (Heyer, in press). Percentages of standard length (SL) are pre-
sented as follows, 5-6. 1-7, where the first number is the minimum, the second
the mean, and the third the maximum percentage. The terminology follows
Peters (1964) and Elias and Shapiro (1957). A dissecting microscope was
used to examine the pattern of melanophore distribution on the ventral surfaces
since melanophores in a contracted state are difficult to distinguish with the
naked eye.
Prior to this study, five characters appeared adequate to define popula-
tions of the Melanonotus species group. Male and female sizes were recorded,
and standardized forms were drawn which encompassed the range of variation
for the ventral pattern, ventrolateral gland development, posterior thigh pat-
tern, and toe-disk development. If a selected character appeared intermediate
in any single frog, it was arbitrarily placed in the category it more closely
resembled.
One to 100 individuals have been collected from each of 454 localities in
Mexico and Middle America. As the status of the frogs in this geographic area
was reasonably clear, samples for analysis were selected every 100 km along
both coasts and in any inland area. In regions where faunal changes might be
expected on the basis of other anuran distribution patterns more samples were
analyzed. The samples, usually 10 frogs per locality, from 56 localities from
Mexico through Panama and ail available South American specimens of the
Melanonotus species group were analyzed. The frogs were examined to deter-
mine how many distinctive morphotypes were present. Examples demonstrat-
ing the range of variation for individuals from each population were then
recorded. Analyses of only the largest specimens of each distinct morphotype
from each locality were recorded. Thus in the following analysis, size refers
to the maximum for males and females for a given locality.
A series of mating call recordings were made in western Mexico in July
1967 on an Uher 4000 L portable tape recorder at 7.5 ips. The tapes were
analyzed on a Kay Sonagraph 6061 B. Information on number of notes per
call group, dominant frequency, harmonics, and frequency shifts was re-
corded from the sonagrams. The tapes are deposited af the University of
Southern California, Department of Biological Sciences. Localities, tapes
1970 Studies on the Frogs of the Genus LEPTODACTYLUS
3
(one individual per tape), and specimens on deposit at the Los Angeles County
Museum of Natural History, respectively, are: COLIMA, Colima, WRH
67-12, LACM 37037; JALISCO, 5 km W Acatlan, 1400 m, WRH 67-20,
LACM 37038, WRH 67-21, WRH 67-22, WRH 67-23, LACM 37039, WRH
67-24, WRH 67-25, LACM 37040; 7.7 km E La Huerta, 340 m, WRH 67-26,
LACM 37041; 37.5 km SW Tecalitlan, 910 m, WRH 67-15, WRH 67-16,
WRH 67-17; 0.5 km NE Tonila, 1300 m, WRH 67-18, LACM 37428;
MICHOACAN, 2.6 km N Capirio, 300 m, WRH 67-13, LACM 37427; NA-
YARIT, 13.2 km NE San Bias, WRH 67-31; Santa Cruz, 15 m, WRH 67-30,
LACM 37043; Tepic, 950 m, WRH 67-27, LACM 37042, WRH 67-28;
SINALOA, 36 km S Los Mochis turnoff on Mexican Highway 15, 10 m, WRH
67-2, LACM 37426, WRH 67-3, LACM 37036; SONORA, 3.1 km E Hermo-
sillo, WRH 67-32.
Certain environmental parameters were chosen to evaluate as possible
correlates with SL, gland development, ventral pattern, and posterior thigh
pattern.
Total annual rainfall, number of months with less than 100 mm of rain-
fall (dry months), and elevation were recorded for each locality. The climatic
data were taken from Espinal and Montenegro (1963) for Colombia; Hold-
ridge (1964) for Middle America; Tosi (1960) for Peru; Vivo Escoto (1964)
for Mexico; Walter and Lieth (1960-67) for South America and Lesser An-
tillean Islands. Elevational data were taken from museum catalogues or from
the Millionth Map series (American Geographical Society).
The correlation coefficient used to compare categories was the product
moment method for ungrouped data, as presented by Arkin and Colton (1966,
pp. 80-82). Mary Nafpaktitis wrote a Fortran program for this formula suited
to my data. Robert J. Lavenberg wrote a Basic program for a Ltest, as pre-
sented by Fisher (1948, pp. 193-196). The correlation coefficients and /-tests
were run on a General Electric Time Share Computer terminal, provided by
the Los Angeles County Museum of Natural History. The t values were
located on a table (Fisher, 1948, p. 174) to determine the level of statistical
significance (F value) of the correlation coefficients. A P value of .05 or less
was considered statistically significant.
Acknowledgments
This report is a modified section from my doctoral dissertation on file at
the University of Southern California. My dissertation committee has been a
constant source of encouragement — Jay M. Savage, Chairman, Robert M.
Chew, John S. Garth, John L. Mohr, Basil G. Nafpaktitis, and John D. Soule.
My fellow graduate students have aided me considerably in helping to define
my ideas through stimulating discussions. Of particular help have been John
R. Meyer, Roy W. McDiarmid, Norman J. Scott, Jr., Philip A. Silverstone
and David B. Wake.
4
Contributions in Science
No. 191
My studies of the genus Leptodactylus would have been lessened without
the help of the following field companions: James R. Dixon, Miriam Heyer,
Roy W. McDiarmid, Marco Tulio Pacheco, Norman J. Scott, Jr,, and Charles
F. Walker.
The curators of collections with which I worked were particularly gra-
cious in providing information, lending large amounts of material and
allowing dissections and skeletal preparations in certain cases ( Museum
abbreviations as used in the text in parentheses) : James E. Bohlke, Academy
of Natural Sciences, Philadelphia (ANSP) ; Werner C. A. Bokermann, Sao
Paulo (WCAB) ; Antenor Leitao de Carvalho, Museu Nacional, Rio de
Janeiro; Javier Castroviejo, Museo Nacional de Ciencias Naturales, Madrid;
Doris M. Cochran and James A. Peters, United States National Museum
(USNM and field series GOV and JAP) ; James R. Dixon, Texas A. & M.
University (TCWC) ; William E. Duellman, Museum of Natural History,
University of Kansas (KU) ; Josef Eiselt, N aturhistorisches Museum, Vienna;
Alice G. C. Grandison, British Museum (Natural History) (BMNH) ; W.
Hellmich, Zoologische Sammlung des Bayerischen Staates, Munich; B. Huben-
dick, Naturhistoriska Museet, Goteborg; Robert F. Inger and Hymen Marx,
Field Museum of Natural History (FMNH); Alan E. Leviton, California
Academy of Sciences (CAS) ; Clarence J. McCoy, Carnegie Museum (CM) ;
Gunther Peters, Institut fiir Spezielle Zoologie und Zoologisches Museum,
Berlin; Hobart M. Smith, formerly of Museum of Natural History, University
of Illinois (UIMNH) ; Robert C. Stebbins, Museum of Vertebrate Zoology,
University of California at Berkeley (MVZ) ; Paulo E. Vanzolini, Departa-
mento de Zoologia, Sao Paulo (DZ); Greta Vestergren, Naturhistoriska
Riksmuseet, Stockholm; Charles F. Walker, Museum of Zoology, University
of Michigan (UMMZ) ; Ernest E. Williams, Museum of Comparative Zoology,
Harvard University (MCZ) ; John W. Wright, Los Angeles County Museum
of Natural History (LACM) ; Richard G. Zweifel, American Museum of
Natural History (AMNH) . Field series from the University of Southern
California collections are designated CRE (Costa Rica), JRM (Honduras), ^
and PAS (Colombia).-
Robert J. Lavenberg and Mary Nafpaktitis facilitated my use of the
General Electric Time Share Computer service provided by the Los Angeles
County Museum of Natural History.
I acknowledge the support from two National Science Foundation Sum-
mer Fellowships for Teaching Assistants (Summers of 1964 and 1965), an
NDEA Dissertation Travel Fellowship administered by the University of
Southern California for travel in the summer of 1967, and a National Science
-To be deposited in the herpetological collection of the Los Angeles County Museum
of Natural History.
1970 Studies on the Frogs of the Genus LEPTODACTYLUS
5
Foundation Graduate Traineeship for full academic support for the past two
years, 1966-1968.
For her constant encouragement and secretarial help, I thank my wife,
Miriam.
Species Accounts
Adult members of the Melanonotus species group may be confused only
with those of the Ocellatus species group because members of both groups
have extensive toe fringes as adults. The Ocellatus group members always
have smooth dorsolateral folds, while members of the Melanonotus group
usually do not have any indication of a dorsolateral fold, or, if such are present,
they are weak and warty, never smooth.
Only brief synonymies are presented, including synonyms and generic
reallocations of the valid species, as Gorham (1966) has recently published
a bibliographic synonymy of the genus Leptodactylus.
Leptodactyhis dantasi Bokermann
Leptodactylus dantasi Bokermann, 1959; 5-8, figs. 1-5 (Type locality, Brasil:
Acre; Feijo. Holotype WCAB 1240, female) .
Diagnostic characters: The only other species in the Melanonotus group
with dark bellies with discrete light spots are podicipinus and pustulatus. L.
dantasi has large light spots on the belly and lacks a metatarsal fold; podici-
pinus has small light spots on the belly and has a well developed metatarsal
fold. Leptodactylus dantasi lacks light spots on the posterior face of the thigh;
pustulatus has large, well defined light spots on the posterior face of the thigh.
Summary of characteristics: Snout rounded from above, rounded acute
in profile; canthus rostralis indistinct; loreal slightly concave in cross section;
tympanum distinct, horizontal diameter 0.5 eye diameter; vomerine teeth in
slightly arched series, posterior to choanae; head length greater than width, 43
per cent SL; head width 39 per cent SL; interorbital distance 6 per cent SL;
first finger much longer than second, first just shorter than third, second longer
than fourth; no distinct ulnar ridge; back warty-pustulose, tibia with many,
pronounced warts; supratympanic fold does not reach shoulder; ventrolateral
glands present but not distinct; toe tips very slightly expanded, dorsal surfaces
not grooved; toes with well developed lateral fringes; subarticular tubercles
moderately well developed; metatarsal fold absent; tarsal fold distinct along
distal two-thirds of tarsus, weakly continuous with toe fringe; tarsus and foot
with many horny spicules, spicules dark on scattered warts; standard length
of female holotype 68 mm; femur shorter than tibia, 37 per cent SL; tibia
shorter than foot, 39 per cent SL; foot longer than femur, 48 per cent SL; upper
lip barred; light outlined dark band from interorbit to sacrum; rest of back
uniform; upper surfaces of limbs faintly barred; venter with large light spots on
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Figure 1. Geographic distribution of Leptodactylus dantasi, discodactyliis, piistii-
latus, and podicipinus. Dashed line indicates 2000 meter contour. Open symbols
indicate sites of sympatry with wagneri.
1970 Studies on the Frogs of the Genus LEPTODACTYLUS
1
a dark ground, spots not always regular, spots anastomosing on limbs; posterior
thigh uniformly dark.
Distribution: Known only from the type locality (Fig. 1 ), 249 m.
Remarks: L. dantasi is at present known only from the holotype.
Leptodactylus discodactylus Boulenger
Leptodactylus discodactyliis Boulenger, 1883 : 637, pi. 58, fig. 3 (Type locality,
Peru: Loreto; Yurimaguas. Holotype BMNH 84.2.18.44, female). Gorham,
1966: 128 (Literature list, synonymy) .
Leptodactylus nigrescens Andersson, 1945: 57-58 (Type locality, Ecuador:
Rio Pastaza and Rio Napo, Watershed. Type series, Naturhistoriska Riks-
museet, Stockholm, unnumbered. Gorham, 1966: 132 (Literature list,
synonymy) .
Diagnostic characters: The only other species in the Melanonotus species
group that has disk-like swellings of the toe tips is wagneri. The upper disk
surfaces of discodactylus have longitudinal grooves; if disks are present in
wagneri, the upper surfaces are never grooved.
Summary of characteristics : Snout rounded, subovoid, or subelliptical
from above, usually rounded or rounded-vertical in profile; canthus rostralis
indistinct; loreal slightly concave in cross section; tympanum distinct, hori-
zontal diameter 0.5-0.66 eye diameter; male vocal slits elongate, arise lateral
to posterior 0.25-0.33 tongue and parallel jaw almost to the angle of the jaw;
single internal vocal sac in males; vomerine teeth in transverse or very slightly
arched series, posterior to choanae; head length greater than width, 37-39.1-43
per cent SL; head width 32-35.2-39 per cent SL; interorbital distance 8-9. 5-1 1
per cent SL; finger tips swollen or not; first finger longer than second, first
shorter than third, second longer than or rarely equal to fourth; no spines on
thumb of male; arm of male not hypertrophied; no distinct ulnar ridge; back
rarely smooth to scattered with coni apicales; tibia especially scattered with
coni apicales; supratympanic fold extends to shoulder; gland present at corner
of mouth or not; post-tympanic gland present or absent; ventrolateral glands if
present, not extensive; glands usually not present on posterior face of thigh;
no chest spines on males; toe tips expanded into disks, dorsal disk surfaces
grooved; toes with well developed lateral fringes; subarticular tubercles well
developed; metatarsal fold present, weakly to strongly developed; tarsal fold
distinct along distal 0.5-0.66 tarsus, not continuous with toe fringe; tarsus
scattered with coni apicales; foot smooth, or outer sole scattered with coni
apicales; standard length of males to 35 mm, females to 35 mm; femur shorter
than tibia, 38-43.6-48 per cent SL; tibia shorter than foot, 43-45.8-49 per cent
SL; foot longer than femur, 48-52.0-56 per cent SL; upper lip uniform or
barred; interorbital region usually with a light stripe bordered behind by a dark
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triangle, the dark triangle may extend to the sacral region, or uniformly dark;
rest of back usually uniform, may have faint large splotches; upper surfaces of
limbs barred to uniform; venter with a profusion of melanophores anteriorly
only, to profuse over the entire venter; posterior thigh mottled.
Distribution : Known elevational range : 150-1000 m.
Known from a few localities of the upper Amazonian drainage in Ecuador
and Peru (Fig. 1).
Nomenclature: Boulenger described L. discodactylus from Peru in 1883.
The subsequent use of the name in the literature has been only a citation of the
original description. Dr. James A. Peters allowed me to send one of his
specimens, JAP 6197, to the British Museum for comparison with the holo-
type of L. discodactylus. Specimen 6197 is from the upper known elevational
range of the species, and its characteristics agree with those of other high
elevation populations discussed in the remarks section. Dr. Alice G. C. Grandi-
son of the British Museum (Natural History) compared specimen 6197 with
the holotype of L. discodactylus. She noted the departures from the holotype
characters associated with elevation (see remarks section, below), but con-
cluded that in all other aspects, including the diagnostic toe fringe, toe disks,
and upper toe disk surface grooves, the specimens were identical.
Andersson described Leptodactylus nigrescens on the basis of three speci-
mens from east Ecuador in 1945. His description is based upon the largest
specimen. I was able to examine the largest syntype of L. nigrescens at the
United States National Museum where Dr. Peters’ private collection was
available for comparison. Mr. Werner C. A. Bokermann had previously bor-
rowed the two smaller specimens and allowed me to examine them at the
National Museum. The type series is composite. The largest syntype of L.
nigrescens possesses the diagnostic combination of toe fringe and toe disks
with grooved upper surfaces of L. discodactylus. The other two specimens
have well developed toe disks, no finger disks, no toe fringe, no tarsal fold, and
two rows (1 pair) of dorsolaterally arranged coni apicales. I consider the
smaller two syntypes of L. discodactylus to represent Eleutherodactylus
nigrovittatus Andersson, described in the same paper as L. nigrescens. In a
cursory literature review, I was unable to find a senior synonym for this distinct
little frog, but one may exist. To avoid future confusion, I hereby designate the
largest (35 mm) specimen (the specimen has no museum number) the lecto-
type of L. nigrescens. Leptodactylus nigrescens Andersson is a junior synonym
of L. discodactylus Boulenger.
Remarks: The few individuals available have certain differences which
correlate with elevation.
Contrasting the highland samples from Ecuador (1000 m) with the low-
land samples from Ecuador and Peru (150-300 m), one finds the following
consistent differences (lowland population characteristics in parentheses) :
1 ) the finger tips are not swollen (finger tips swollen, forming small disks) ;
1970 Studies on the Frogs of the Genus LEPTODACTYLUS
9
2) the tarsal fold is usually distinct along the distal 0.66 of the tarsus (tarsal
fold usually distinct on distal 0.5 of tarsus); and 3) smaller adult size, males
to 30 mm, females to 32 mm (both males and females to 35 mm). I assume
that the differences noted are responses to different climatic parameters
associated with differences in elevation.
Leptodactylus discodactylus has been taken in sympatry with L. wagneri
at two localities, Ecuador: Pastaza; Puyo, 1000 m and Peru: Loreto; Rio
Tamaya, Sobral, 150 m.
Leptodactylus melanonotus (Hallowell)
Cystignathus melanonotus Hallowell, 1860: 485 (Type locality, Nicaragua.
Type apparently lost).
Cystignathus echinatus Brocchi, 1877: 181-2 (Type locality, Guatemala: Rio-
madre Nieja. Syntypes Paris Museum 6322-3) .
Cystignathus microtis Cope, 1879: 265 (Type locality, Mexico: Guanajuato;
Guanajuato. Syntypes USNM 9906, 9908, 9909).
Cystignathus perlaev is CopQ, 1879: 269-270 (Type locality, Mexico: Oaxaca;
Japana. Holotype USNM 10041, female).
Leptodactylus melanonotus, Brocchi, 1881: 20 (Cites Hallowell’s record).
Gorham, 1966: 131 (Literature list, synonymy).
Leptodactylus occidentalis Taylor, 1937: 349-52, pi. 1, figs. 1, 2, 7 (Type
locality, Mexico: Nayarit; Tepic. Holotype FMNH 100015, female). Gorham,
1966: 133 (Literature list, synonymy).
Diagnostic characters: There is no one character that immediately dis-
tinguishes L. melanonotus from the other members of the Melanonotus species
group. The toe disks of L. discodactylus distinguish it from L. melanonotus
(toe tips not expanded into disks). Leptodactylus dantasi, L. podicipinus, and
L. pustulatus have dark bellies with discrete light spots; L. melanonotus may
have a dark belly, but it is mottled and never distinctly spotted. Certain few
individuals of L. melanonotus are difficult to distinguish consistently from
L. wagneri (L. melanonotus characters in parentheses) : L. wagneri may have
the toe tips expanded into distinct disks (toe tips never disk-like); L. wagneri
may have a light longitudinal stripe on the posterior face of the thigh (no
distinct stripe) ; L. wagneri reaches a larger adult size, standard length of
males to 63 mm, females to 81 mm (males to 46 mm, females to 50 mm). The
only way to distinguish consistently L. melanonotus from L. wagneri is by
geography. Leptodactylus melanonotus is distributed from Mexico through
Middle America, and west of the Andes in South America. Leptodactylus
wagneri is found east of the Andes in South America.
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Summary of characteristics : Snout rounded-nearly semicircular, round-
ed, rounded-subelliptical, or rarely subovoid from above, rounded to rounded-
vertical in profile; canthus rostralis indistinct; loreal slightly concave in cross
section; tympanum distinct, horizontal diameter 0.5 to 0.75 eye diameter; male
vocal slits elongate, arise lateral to mid-point of tongue to almost the angle of
the jaw, usually parallel to jaw, rarely slightly oblique to the jaw; single internal
vocal sac in males; vomerine teeth usually in transverse series, rarely very
slightly arched, always posterior to the choanae; head usually longer than
wide, rarely equal, or rarely head length shorter than wide, head length SO-
SO.3-41 per cent SL; head width 30-34.1-38 per cent SL; interorbital distance
5-7. 2-9 per cent SL; finger tips not noticeably swollen; first finger just longer
than or about equal to second, first shorter than third, second longer than
fourth; finger ridges present, especially on second and third fingers; 2 spines on
male thumb; male arm not hypertrophied; ulnar ridge not developed; head
smooth or scattered with coni apicales; upper eyelids warty, glandular or
smooth, usually lacking coni apicales; rest of back with scattered coni apicales,
the coni may be on warts or not; upper femur and tibia scattered with coni
apicales; supratympanic fold extends to shoulder, rarely indications of several
warty dorsolateral folds; brown to orange ventrolateral glands poorly to ex-
tensively developed, same glandular material may or may not be on jaw angle,
post-tympanic region, groin, posterior thigh, or inner tibia and along the tarsal
fold; no chest spines on male; toe tips usually not expanded, sometimes slightly
expanded, never disk-like, never with dorsal surfaces grooved; toes with well
developed lateral fringes; subarticular tubercles well developed; metatarsal
fold present, weakly to well developed; tarsal fold distinct along distal 2/3-
5/6 tarsus, not continuous with toe fringe; tarsus with scattered coni apicales;
foot smooth, with scattered coni apicales, or coni apicales on outer sole only;
standard length of males to 46 mm, females to 50 mm; femur longer than,
equal to, or usually shorter than tibia, 36-40.0-45 per cent SL; tibia shorter
than foot, 37-42.7-47 per cent SL; foot longer than femur, 43-49.3-53 per
cent SL; upper lip barred to uniform; interorbit with dark, light outlined tri-
angle, a light and dark bar, or a light triangle reaching tip of snout anteriorly,
bordered posteriorly by a dark triangle; rest of back brown to gray with
darker indistinct spots, blotches, bands, stripes, or without pattern; upper limb
surfaces barred to uniform; venter with few melanophores, appearing light to
many melanophores profused especially anteriorly to rarely heavily profused
over entire venter; posterior thigh mottled.
Distribution : Elevational range : sea level — 1440 m. The species is known
from both coasts of Mexico — Hermosillo, Sonora and the Gomez Farias
region, Tamaulipas in the north, extending southward throughout Middle
America. The species is known only from the western lowlands of South
America to mid-Ecuador. (Figs. 2-4.)
Nomenclature: The holotype of Cystignathus melanonotus Hallowell is
1970 Studies on the Frogs of the Genus LEPTODACTYLUS
11
Figure 2. Geographic distribution of Leptodactylus melanonotus in Mexico.
apparently lost. The late E. R. Dunn noted (Dr. Dunn’s notes presently on
deposit in the library of Dr. Jay M. Savage) that the type was originally
deposited in the National Museum as specimen number 6264 and that it was
missing. The specimen is not listed in the catalog of type specimens of reptiles
and amphibians in the U.S. National Museum (Cochran, 1961). In recent
visits to the U.S. National Museum, although aided by the late Dr. Cochran
and by Dr. Peters, I was unable to locate the type. The evidence indicates that
the original holotype is lost.
Leptodactylus melanonotus is very closely related to both L. wagneri and
L. podicipinus. L. melanonotus is found in Mexico and Middle America, where
there is relatively little question as to what the species is. L. melanonotus is
found west of the Andes and some individuals are very difficult to differentiate
consistently from some individuals of L. wagneri, an Amazonian species. For
nomenclature stability, the name L. melanonotus should be applied to a single
specimen from the population found in Nicaragua, the type locality of the ori-
ginal, now lost, holotype. I therefore designate specimen number 84848 in the
collection at the University of Kansas, an adult male, from Nicaragua, Zelaya,
Bonanza, as the neotype of Cystignathus melanonotus Hallowell.
I have examined the holotype and two paratypes of Cystignathus microtis
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Figure 3. Geographic distribution of Leptodactylus melanonotus in Middle America.
Cope. Only two species of Leptodactylus (labialis and melanonotus) occur in
Mexico; they are quite distinct from each other. Members of the type series
of C. microtis agree with L. melanonotus in having fringed toes and mottled
thighs; both characters differentiate L. melanonotus from L. labialis. There is
no morphological reason for retaining C. microtis. The status of C. microtis
has always been questioned, because the type locality was given as Mexico :
Guanajuato; Guanajuato. Guanajuato is well above the elevational limits of
L. melanonotus. No other specimens of L. melanonotus have since been taken
from Guanajuato. I think the locality data are in error and probably represent
the shipping point from which the specimens were sent to the U.S. National
Museum rather than the collecting site.
I have examined numerous paratypes of Leptodactylus occidentalis
Taylor and have collected topotypic specimens which correspond closely with
the type series of L. occidentalis. As is shown in the character analysis portion,
L. occidentalis does not differ in any consistent way from L. melanonotus and
is therefore considered a junior synonym of the latter.
I have examined the holotype of Cystignathus perlaevis Cope and concur
with the numerous workers who have placed it in synonymy with L. mela-
nonotus.
1970 Studies on the Frogs of the Genus LEPTODA CTYL US
13
Figure 4. Geographic distribution of Leptodactylus melanonotus in South America,
Dashed line indicates 2000 meter contour. Open symbol indicates site of sympatry
with wagneri.
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No. 191
Remarks: A single specimen of L. melanonotus is recorded from Starr
County, Texas, in the United States. I have examined the specimen (UIMNH
29817); it is a juvenile L. melanonotus. This single record is about 400 km
north of the next recorded locality, from which several samples are known.
The locality data for the Texas specimen is possibly in error, and for that
reason I have not included the record in the range of L. melanonotus.
Leptodactylus melanonotus has been taken in sympatry with L. wagneri
at one locality in western Colombia (Narino, La Guayacana) . This instance of
sympatry is not too surprising, as the sources of the rivers Cauca and Patia
interdigitate at about 1600-1800 m within the range of L. wagneri. Lepto-
dactylus wagneri is found throughout the Valle de Cauca. Apparently L.
wagneri follows the Rio Patia into the west-coast lowlands of Colombia.
Leptodactylus podicipinus (Cope)
Cystignathus podicipinus Cope, 1862; 156 (Type locality, Paraguay. Holotype
ANSP 14539).
Leptodactylus podicipinus, Boulenger, 1882: 248 (Catalogue, synonymy).
Gorham, 1966: 135-6 (Literature list, synonymy).
Leptodactylus nattereri Lutz, 1926: 1011-2 (Type locality, Brasil: Sao Paulo;
Itapura, Ilha Seca. Syntypes AL 1314-1315).
Leptodactylus podicipinus podicipinus, Gans, 1960: 305-6 (List) . Gorham,
1966: 136 (Literature list, synonymy) .
Diagnostic characters : The only other species in the Melanonotus species
group with a dark belly with discrete light spots are L. dantasi and L. pustu-
latus. L. dantasi has large light spots on the belly and lacks a metatarsal fold.
L. podicipinus has small light spots on the belly and has a well developed
metatarsal fold. L. pustulatus has large, light discrete spots not only on the
belly, but on the entire ventral surface of the leg and on the posterior surface
of the thigh. L. podicipinus has small spots which, if on the leg, are only on the
ventral surface of the thigh. The males of L. podicipinus have thumb spines,
the males of L. pustulatus lack thumb spines.
Summary of characteristics : Snout rounded, subelliptical to subovoid
from above, rounded in profile; canthus rostralis indistinct; loreal slightly con-
cave in cross section; tympanum distinct, horizontal diameter eye
diameter; male vocal slits elongate, arise lateral to middle of tongue, parallel
jaw almost to angle of jaw; single internal vocal sac in males; vomerine teeth
usually in transverse series, rarely in very slightly arched series, always
posterior to choanae; head length greater than width, 34-37.0-41 per cent SL;
head width 32-34.6-38 per cent SL; interorbital distance 7-8.2-10 per cent SL;
finger tips not swollen; first finger usually longer than second, sometimes
1970 Studies on the Frogs of the Genus LEPTODACTYLUS
15
equal, first finger much shorter than, shorter than, or about equal to third,
second longer than fourth; finger ridges present, especially on second and
third fingers; 2 spines on male thumb; male arm not hypertrophied; ulnar
ridge usually not developed, sometimes a glandular ridge or 4 coni apicales in
a row; dorsum smooth, glandular, or warty, scattered with coni apicales, es-
pecially on posterior dorsum and upper leg surfaces; supratympanic fold
extends to shoulder, may or may not be faint indication of several warty
dorsolateral folds; ventrolateral glands orange-brown, small to large or absent,
glands present or absent on angle of jaw, dorsolateral folds, posterior thigh,
inner tibia, and along tarsal fold; no chest spines on male; toe tips not ex-
panded; toes with well developed lateral fringes; subarticular tubercles well
developed; metatarsal fold well developed; tarsal fold distinct along distal
2/ 3-5/ 6 tarsus, usually not continuous with toe fringe, rarely weakly con-
tinuous with toe fringe; tarsus scattered with coni apicales; foot smooth or outer
sole with scattered coni apicales; standard length of males to 38 mm, females
to 44 mm; femur usually shorter but sometimes longer than tibia, 35-40.9-48
per cent SL; tibia shorter than foot, 38-42.9-48 per cent SL; foot longer than
femur, 44-50.7-56 per cent SL; upper lip barred or uniform; interorbital
region with light and dark stripe, or light outlined dark triangle extending to
sacrum or not; rest of back uniform or with indistinct darker bands; upper
limb surfaces uniform or with indistinct dark bars; throat, chest, belly dark
with small light spots, rest of venter dark, but not with distinct light spots,
posterior portion of belly occasionally with limb pattern rather than light-
spotted; posterior thigh usually mottled, rarely with a moderately distinct light
longitudinal line.
Distribution: Elevational range: 18-550 m.
Leptodactylus podicipinus is found south of the Amazonian drainage
region in the following river drainages: Sao Francisco, Parana, Paraguay. The
species is absent from the narrow coastal lowland strip along eastern Brazil
from the States of Rio Grande do Norte to Rio de Janeiro. The species follows
the Serras de San Jose, Santiago, and Sunsas in Bolivia, and the Serras dos
Parecis and Pacaas Novos in western Brasil. (Fig. 1 ) .
Nomenclature: In his description of Cystignathus podicipinus, Cope
designated as type(s) “Mus. Smithsonian, (No. 5831) Philada. Acad.” Dr.
James A. Peters informed me that in the catalog under 5831 is a later remark
stating that the specimen had been sent to the Academy of Natural Sciences in
Philadelphia. Apparently there was only one specimen upon which Cope based
C. podicipinus. I have examined the holotype of C. podicipinus from the
Academy of Natural Sciences. The specimens I have examined from Paraguay
are morphologically similar to the holotype in all diagnostic features.
Lutz described Leptodactylus nattereri on the basis of specimens collected
in the state of Sao Paulo, Brasil, and included the specimens figured by Stein-
dachner (1864) from the collection of Natterer. The figures in Steindachner
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No. 191
(Plate XI, figs. 1 a-d) show the two spines per thumb and dark belly with
small light spots characteristic of L. podicipinus.
Remarks: Leptodactylus podicipinus has been taken in sympatry with
L. wagneri at two localities: Bolivia (Santa Cruz: Buenavista, 500 m) and
Brasil (Rondonia: Forte Principe da Beira, 100-200 m). At both localities
the species are very distinct, markedly differing in size, belly pattern, and
posterior thigh pattern. There is apparently a narrow band of sympatry
between L. podicipinus and L. wagneri, with character displacement occurring
in the band of sympatry.
Leptodactylus pustulatus (Peters)
Entomoglossus pustulatus W. Peters, 1870: 647, PI. 2, fig. 1 (Type locality,
Brasil: Ceara. Type apparently lost).
Leptodactylus pustulatus, Boulenger, 1882: 239 (Catalogue, synonymy) ;
Gorham, 1966: 137 (Literature list, synonymy).
Diagnostic characters : The only other species in the Melanonotus species
group with a dark belly with discrete light spots are L. dantasi and L. podici-
pinus. Leptodactylus pustulatus has large, light discrete spots on the posterior
surface of the thigh. Leptodactylus dantasi and L. podicipinus do not have
spots on posterior surface of the thigh. The males of L. pustulatus lack thumb
spines, the males of L. podicipinus have thumb spines.
Summary of characteristics: Snout rounded to subovoid from above,
rounded to rounded-obtuse in profile; canthus rostralis indistinct; loreal slightly
concave in profile; tympanum distinct, horizontal diameter 2/3 eye diameter;
male vocal slits elongate, parallel to jaw, extend from mid-tongue almost to
angle of jaw; single internal vocal sac in male; vomerine teeth usually in
transverse series, rarely very slightly arched, always posterior to choanae; head
length greater than width, 35-36.4-38 per cent SL; head width 33-34.0-35 per
cent SL; interorbital distance 5-6. 7-8 per cent SL; finger tips not expanded;
first finger longer than second, first shorter than third, second greater than
fourth; finger ridges present, especially on fingers two and three; no spines on
male thumb; male arm not hypertrophied; ulnar ridge not developed; dorsal
texture pustulose, with scattered coni apicales; supratympanic fold extends to
shoulder, may or may not be indications of 6-8 warty dorsolateral folds; brown
ventrolateral glands extend onto belly, or entire frog appears glandular; no
chest spines on males; toe tips not expanded; toes with well developed lateral
fringes; subarticular tubercles well developed; metatarsal fold present; tarsal
fold distinct along distal Vi-Va- tarsus, usually continuous with toe fringe,
occasionally weakly continuous; tarsus with scattered coni apicales; foot
usually with scattered coni apicales, rarely smooth; standard length of males
to 38 mm, females to 51 mm; femur shorter than tibia, 37-39.0-41 per cent SL;
tibia shorter than foot, 39-41.8-44 per cent SL; foot longer than femur, 45-
1970 Studies on the Frogs of the Genus LEPTODACTYLUS
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51.3-55 per cent SL; dorsum uniform or patterned, if patterned, a light inter-
orbital triangle originating at tip of snout bordered behind by a dark triangle,
rest of back with darker longitudinal lines; upper limb surfaces faintly barred
to uniform; venter dark with large light spots extending onto lower leg surfaces
and groin; posterior thigh with light spots as on venter.
Distribution: Known elevational range: 100-200 m.
The few localities from which L. pustulatus have been collected are all
low Amazonian. The major river drainages for which accurate locality data are
available include the Paranaiba, Araguaia, and Tocantins. (Fig. 1).
Nomenclature: Entomoglossus pustulatus was described by W. Peters on
the basis of a single female from Ceara, Brasil, in the collection of the Royal
Zoological Museum, Berlin. Dr. Peter Beurton of the Berlin Museum states
that he was unable to locate the type in the collections there. Dr. Hellmich of
the Munich Museum informed me that the type of Entomoglossus pustulatus
is not and has never been in the Munich collection, the likely alternative
depository for the type. This suggests that the type is lost. The identity of
L. pustulatus has been very unclear because of the rarity of specimens in
collections. Because of this confusion and because there appears to be no other
Ceara (topotypic) material in collections, I have selected an individual repre-
sentative of the population that I consider to be L. pustulatus. The nearest
locality to the type locality is the Rio Poti in Piaui. The specimen from this
locality is only in a fair state of preservation, but it still retains the pattern
diagnostic of L. pustulatus. I designate this specimen, MCZ 373, from the
collection of the Museum of Comparative Zoology, Harvard University, as
the neotype of Entomoglossus pustulatus Peters.
Remarks: L. pustulatus has been taken in sympatry with L. wagneri at
one locality (Brasil: Mato Grosso: Sao Domingoes, Rio das Mortes, 200 m).
Leptodactylus wagneri (Peters)
Plectromantis wagneri W. Peters, 1862: 232-3 (Type locality, Ecuador, west
side of the Andes. Type formerly at Munich Museum, destroyed in World
War II).
Platy mantis petersii Steindachner, 1864: 254-6, PI. 16, fig. 2, 2 a-c (Type
locality, Brasil: Amazonas; Marabitanas. Type lost).
Leptodactylus brevipes CopQ, 1887: 51-2 (Type locality, Brasil: Mato Grosso,
Chupada, 30 mi. NE of Cuyata, near the headwaters of the Xingu. Holotype
ANSP 11270, female).
Leptodactylus validus Garman, 1887: 14 (Type locality. West Indies: St.
Vincent; Kingston. Syntypes ANSP 26108, MCZ 2185).
Leptodactylus wagneri Nieden, 1923: 479 (Literature list, synonymy). Gor-
ham, 1966: 140 (Literature list, synonymy).
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Leptodactylus pallidirostris A. Lutz, 1930: 25-6, PL 1, fig. 3 (Type locality,
British Guiana: Essequibo; Kartabo).
Leptodactylus natalensis A. Lutz, 1930: 26-7, PI. 1, figs. 7-7a, PI. 3, figs. 1-2
(Type locality, Brasil: Rio Grande do Norte; Rio Bahu. Syntype USNM
81130, male).
Leptodactylus intermedius A. Lutz, 1930: 27-8, PI. 3, fig. 6 (Type locality,
Brasil: Amazonas; Manacapuri near Manaos).
Eleutherodactylus leptodactyloides Andersson, 1945: 43-4, fig. 15 (Type
locality, Ecuador: Pastaza; Rio Pastaza. Holotype Stockholm Royal Museum,
male).
Leptodactylus podicipinus petersii, Rivero, 1961: 47 (Redescription, syn-
onymy). Gorham, 1966: 136 (Literature list, synonymy).
Diagnostic characters: Leptodactylus wagneri may have expanded toe
tips, but the upper surface of the toe tip is never with longitudinal grooves,
distinguishing wagneri from discodactylus, which has well developed toe disks
with the upper disk surfaces grooved. Leptodactylus wagneri may have a dark
belly, but it is mottled, and not with distinct light spots, distinguishing it from
the dark-bellied, light-spotted dantasi, podicipinus, and pustulatus. Some in-
dividuals of wagneri are difficult to distinguish consistently from melanonotus
(melanonotus characters in parentheses); wagneri may have the toe tips ex-
panded into distinct disks (toe tips never disk-like) ; wagneri may have a light
longitudinal stripe on the posterior face of the thigh (no distinct stripe);
wagneri reaches a larger adult size, standard length of males to 63 mm, females
to 81 mm (males to 46 mm, females to 50 mm). The only consistent way to
distinguish melanonotus from wagneri is by geography. Leptodactylus mela-
nonotus is distributed from Mexico through Middle America, and west of the
Andes in South America. Leptodactylus wagneri is found east of the Andes
in South America.
Summary of characteristics: Snout usually rounded, rarely subelliptical
or subovoid from above, rounded to rounded-vertical in profile; canthus
rostralis indistinct; loreal slightly concave in cross section; tympanum dis-
tinct, horizontal diameter ’/2-% eye diameter; male vocal slits elongate, arise
lateral to tongue, parallel jaw and extend almost to the angle of the jaw or
slightly oblique to jaw; single internal vocal sac in male; vomerine teeth in
transverse to arched series, posterior to choanae; head length greater than,
equal to, or less than width; head length 33-36.8-42 per cent SL; head width
32-34.9-39 per cent SL; interorbital diameter 6-7. 8-9 per cent SL; finger tips
not noticeably swollen; first finger longer than second, first shorter than or
equal to third, second longer than or equal to fourth; fingers with lateral ridges,
especially on second and third fingers; two spines on male thumb; male arm
1970 Studies on the Frogs of the Genus LEPTODACTYLUS
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slightly hypertrophied only in largest specimens; no ulnar ridge; dorsal texture
smooth to glandular, posterior portion of back and legs with scattered coni
apicales; supratympanic fold extends to shoulder, may or may not be indica-
tion of 2-4 faint dorsolateral folds; orange-brown ventrolateral glands moder-
ately developed or absent; gland at angle of jaw developed or not; post-
tympanic gland developed or not; posterior thigh gland present or absent; no
chest spines on male; toe tips not expanded to moderately expanded and
disk-like, never with longitudinal grooves on dorsal surface; toes with well
developed lateral fringes; subarticular tubercles well developed; metatarsal
fold present; tarsal fold distinct along distal 4/7-3/4 tarsus, not continuous
with toe fringe; tarsus with scattered coni apicales; foot smooth, with scat-
tered coni apicales, or outer sole with scattered coni apicales; standard length
of males to 63 mm, females to 81 mm; femur shorter than tibia, 35-42.4-48 per
cent SL; tibia shorter than foot, 41-46.9-51 per cent SL; foot longer than
femur, 49-52.4-57 per cent SL; upper lip usually barred, sometimes uniformly
patterned; light and dark interorbital bar, dark bar may extend posteriorly as a
triangle or band as far as sacrum; rest of back uniformly patterned or indis-
tinctly spotted or striped, warts on sides may be darker than ground color;
upper limb surfaces barred to uniform; ventral pattern of scattered melano-
phores, melanophores may be profuse anteriorly only, or profuse over entire
venter; posterior thigh with distinct light longitudinal stripe to mottled.
Distribution: Elevational range: sea level — 1900 m.
The species is widely distributed throughout northern and central South
America east of the Andes. There is a single record west of the Andes in
Colombia. The species occurs in the Cauca and Magdalena valleys of west-
central Colombia, and throughout the greater Amazonian Basin; the species is
also distributed along the narrow eastern coastal strip of Brasil from the States
of Rio Grande do Norte to Rio de Janeiro. The species occurs on the islands
of Bequia, Granada, St. Vincent, Tobago, and Trinidad. (Fig. 5) .
Nomenclature: Wilhelm Peters (1862) described Plectromantis wagneri
from Ecuador on the basis of a single male specimen. Peters listed the type as
“von Dr. Moritz Wagner an den Westseite der Anden in Ecuador . . . im
zoologischen Cabinet du Munchen.” Dr. Richard Etheridge kindly looked for
the type when he was at the Munich museum. He writes (personal communi-
cation), “. . . the old museum had one specimen. No. 1080/0, from ‘Pastassa’
collected by Wagner. This may or may not have been the type, there is no
indication in the record that it was, but in any event the specimen was de-
stroyed during World War II.” It is likely that specimen number 1080/0
was the unique type of Plectromantis wagneri. Pastassa is a geographically
more reasonable locality than the west side of the Andes. Peters’ description is
quite good, and the secondary sexual characteristics of spines on the thumb of
the male leave no doubt as to the identity of the frog described by Peters. It is
the population herein called L. wagneri. The only Leptodactylus west of the
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Figure 5. Geographic distribution of Leptodactyliis wagneri. Dashed line indicates
2000 meter contour. Open symbols indicate sites of sympatry with other species of
the Melanonotus group.
1970 Studies on the Frogs of the Genus LEPTODACTYLUS
21
Andes in Ecuador with thumb spines on the male is L. melanonotus. The males
of L. melanonotus reach 40 mm SL in Ecuador; the type of P. wagneri was 68
mm SL. I consider that Peters’ locality is probably incorrect, and the data as
recorded for Munich specimen 1080/0 are correct. The name L. wagneri has
never been used in association with a Leptodactylus population in the literature
since the original description. The status of the northern South American
population of the Melanonotus group has always been uncertain. To stabilize
the nomenclature I hereby designate the holotype of Eleutherodactylus lepto-
dacty hides Andersson in the Stockholm Royal Museum of Natural History
(no number), an adult male, from the Rio Pastaza, as the neotype of
Plectromantis wagneri Peters.
Steindachner described Platymantis petersii on the basis of a single male
specimen from Marabitanas, Amazonas, Brasil. Dr. Josef Eiselt of the Vienna
Museum has been unable to find the unique type of Platymantis petersii. The
type has been searched for several times and is most likely lost. Leptodactylus
petersii has been used in various combinations for various populations of frogs
of the Melanonotus species group. I consider P. petersii to be a junior synonym
of L. wagneri, and that nomenclatural stability would be served by designating
a neotype which conforms with my analysis. I have not examined any material
from Marabitanas, nor do I know of any material from there. The closest
record that I know of to Marabitanas is in Venezuela. This specimen, AMNH
23182, is in very good condition. The locality is Venezuela: Amazonas; Rio
Pescado, approximately 150 m. The specimen is a female. I hereby designate
AMNH 23182 as neotype of Platymantis petersii Steindachner.
Cope described Leptodactylus brevipes on the basis of a single specimen
from Chupada, Mato Grosso, Brasil. The name has been cited in literature lists
based upon the original description but not upon the basis of recent material. I
have examined the type, and find it the same as wagneri. The belly has very
few melanophores, so that no confusion is possible with either podicipinus or
pustulatus, the only other members of the Melanonotus species group with
which it could possibly occur or be confused. Leptodactylus brevipes is re-
garded as a junior synonym of L. wagneri.
Leptodactylus validus was described by Garman from a series of speci-
mens from the island of St. Vincent. I have found no differences between any
of the island populations and the mainland population of wagneri, as shown
below in the character analysis section. No holotype was designated by Gar-
man. I hereby designate MCZ 71920, an adult male, from Kingston, St. Vin-
cent, as the lectotype of Leptodactylus validus Garman.
Lutz described three species of Leptodactylus of the Melanonotus group
in the same paper in 1930. The first, pallidirostris, was described from British
Guiana: Essequibo; Kartabo. I have not been able to examine the type, but
have examined numerous examples of the Melanonotus group from the type
locality. All specimens represent a single species and conform with the type
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description of pallidirostris. The specimens from Kartabo are not distinct from
wagneri. Thus, L. pallidirostris is considered a junior synonym of L. wagneri.
Leptodactylus natalensis is the second species described by Lutz. I have
examined one syntype of the type series and find it the same as other specimens
examined in this study from the type locality (vicinity of Natal). The syntype
has a lightly pigmented venter and marbled posterior thigh surfaces. I con-
sider L. natalensis Lutz to be a junior synonym of L. wagneri. No holotype was
designated. I designate the specimen I have examined, USNM 81130, a male,
as the lectotype of Leptodactylus natalensis Lutz.
The third species of the Melanonotus group described by Lutz is L. inter-
medins. As in the case of pallidirostris, I have not seen the types, but have
examined material from the type locality. The topotypic material I have ex-
amined from Manaus agrees with the description of intermedins. I consider the
material from Manaus to be wagneri', hence intermedins is a junior synonym
of L. wagneri.
Andersson described Eleutherodactylus leptodactyloides on the basis of a
single specimen from Ecuador. I have been able to examine the type specimen.
It is a Leptodactylus as suggested by the specific name, rather than an Eleu-
therodactylus. I have designated this specimen, above, as the neotype of
Plectromantis wagneri, thereby making Eleutherodactylus leptodactyloides a
junior synonym of L. wagneri.
Remarks'. Leptodactylus wagneri has been taken in sympatry with dis-
codactylus, melanonotus, podicipinus, and pustulatus.
Key to Adult Members of the Melanonotus Species Group
la. Toe tips expanded into disks, dorsal surfaces with longitudinal grooves
discodactylus
lb. Toe tips usually not expanded into distinct disks, dorsal surfaces never
grooved 2
2a. Ventral surfaces of legs and posterior surface of thigh with large, light,
distinct spots pustulatus
2b. Ventral surfaces of legs light or dark, if large spots present, spots not
distinct; posterior surface of thigh light or dark, never with large, light,
distinct spots 3
3a. Belly dark with distinct, light spots, spots usually distinct over entire belly,
but sometimes anastomosing on posterior belly 4
3b. Belly light or dark; if dark, no distinct light spots 5
4a. Belly with large light spots; no metatarsal fold dantasi
4b. Belly with small light spots; well developed metatarsal fold podicipinus
5a. Moderate sized frogs, males to 46 mm SL, females to 50 mm; posterior
thigh never with light longitudinal stripe; toe tips never noticeably ex-
panded; Mexico, Middle America, South America west of the Andes to
Ecuador .melanonotus
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23
5b. Moderate to large frogs, males to 63 mm SL, females to 81 mm; posterior
thigh may have a light longitudinal stripe; toe tips often noticeably ex-
panded; South America east of the Andes wagneri
Character Analysis
Standard length: Two broad geographic patterns are evident in male
L. melanonotus. The coastal populations vary between 30 and 40 mm. Modal
sized individuals (36 mm Hermosillo, Sonora, Mexico) or smaller specimens
(34 mm Gomez Farias region, Tamaulipas, Mexico; 33 mm Ecuador) are
associated with the extreme northern and southern geographic range. The
largest specimens are associated with two lowland pass regions in the montane
backbone of the Americas: the lowland pass at Tehuantepec, Mexico; and the
Arenal pass in Costa Rica. The geographic pattern of female L. melanonotus
size parallels the male geographic pattern, but the trends are not as pro-
nounced. In addition to the Tehuantepec and Arenal passes harboring the
largest females (47-50 mm), the Yucatan Peninsula and eastern lowlands of
Costa Rica also have females of large size (46-48 mm). The range of adult
female size is approximately 35-50 mm.
Leptodactylus wagneri males attain greatest size along the eastern slopes
of the Andes in Colombia, Ecuador, and Peru, ranging from about 40-60 mm.
Along the northern coast of South America, on the islands, and in the Ama-
zonian basin, the sizes range from 30-40 mm. The geographic pattern of
L. wagneri female size exactly parallels the male adult size with the exception
of the ranges. The east Andean slope populations most frequently range in
size from 50-80 mm; the north coast, island, and Amazonian populations
usually range from 40-60 mm.
Adult male L. podicipinus range from about 28-38 mm in length, with a
trend towards smaller size at the southern end of the geographic range. Adult
female L. podicipinus range in size from 36-44 mm also with a trend towards
smaller size at the southern end of the geographic range.
The geographic pattern of size variation is basically the same in males
and females, the only difference being that the females average larger than
the males.
The relationship between size and rainfall, number of dry months, and
elevation are statistically significant only in the case of L. wagneri. In L. wag-
neri males, a statistically significant correlation is present between greatest size
of adults and mesic habitats (r = .28, P = <.01 ). This correlation appears to
be biologically valid for both males and females, as a statistically significant
correlation was found with smaller size in more xeric habitats as measured by
number of dry months (males r = —.50, P — <.01, females r = — .27,
P — <.01). A statistically significant correlation was found for greatest size
and increasing elevation for both males and females in L. wagneri (males
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Figure 6. Ventral color pattern standards. Note that pattern F is restricted to
Leptodactylus piistulatiis.
1970 Studies on the Frogs of the Genus LEPTODACTYLUS
25
T = .38, P = <.01, females r = .42, P = <.01 ) . The elevational effects pro-
ducing size differences in these frogs are probably associated with rainfall
differences also.
Character displacement apparently occurs in the narrow bands of sym-
patry between L. melanonotus, podicipinus, and wagneri. At La Guayacana in
Colombia, a female melanonotus (mature) measures 38 mm, and a female
wagneri measures 59 mm. Two instances of sympatry are known for wagneri
and podicipinus. At Buenavista, Bolivia, a female podicipinus measures 40
mm, and a female wagneri 46 mm. At Forte Principe da Beira, Brasil, a female
podicipinus measures 34 mm, a female wagneri 43 mm, and a male podicipinus
29 mm, and a male wagneri 38 mm.
Ventral pattern : Five categories were used in scoring the ventral pattern.
These ranged from A, in which the throat and belly do not have any melano-
phores, to E, in which the throat and belly are heavily covered with mela-
nophores, leaving light spots (Fig. 6).
The distribution of ventral pattern is variable throughout most of the
geographic range of L. melanonotus. In many localities, conditions ranging
from B to D, encompassing the variation encountered in this species, are
found. The populations at the northern and southern extremes of the geo-
graphic range lack much of the variability as in other populations, and are
lighter in ventral coloration. Northwestern Mexican and South American
populations are average in pattern density. There is a weak trend of darker
pattern in Atlantic lowland populations.
No clear geographic trend in ventral pattern in L. wagneri is evident. The
usual range for the species is B to D, with a few populations having an inter-
mediate condition between D and E. Most adequate samples (10 or more
individuals) show a variation through only two categories, either B-C or C-D.
The occurrence of individuals with pattern B seems to be random with per-
haps more B category individuals throughout the northern part of the species
range, including the populations of islands off northern South America.
There is no geographic trend of ventral color pattern in L. podicipinus.
Almost all populations fall in the E category, with a few individuals in D or
between D and E.
A correlation between a darker ventral color pattern and greater annual
rainfall is present in L. melanonotus (r = .17, P — .05). There is no correlation
between ventral pattern and glandular extent, number of dry months, or ele-
vation in melanonotus. No correlations between ventral color pattern and
glandular extent, annual rainfall, number of dry months, or elevation are
present in podicipinus. A correlation is found between a lighter ventral color
pattern and greater ventrolateral gland extent in wagneri (r = —.14, P = .05).
This is most certainly not a cause-effect relationship, but it may indicate that
the genes responsible for these two characters are on the same chromosome or
are co-functional in some other way. No correlations between ventral color
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Figure 7. Ventrolateral gland standards.
1970 Studies on the Frogs of the Genus LEPTODACTYLUS
27
pattern and annual rainfall, number of dry months, or elevation are found for
wagneri.
Ventral color pattern is a character that has apparently undergone charac-
ter displacement in the area of sympatry between L. podicipinus and wagneri.
Certain individuals of wagneri closely approach the pattern of podicipinus. In
the two known instances of sympatry of podicipinus and wagneri, the ventral
color patterns are clearly distinct; wagneri individuals have a light belly;
podicipinus individuals have a dark, light spotted belly. Leptodactylus dis-
codactylus and wagneri have basically the same ventral color patterns. Lepto-
dactylus pustulatus is very distinct from wagneri with respect to ventral pattern.
Leptodactylus pustulatus has a distribution allopatric to those of dantasi and
podicipinus, the species it most closely resembles in ventral color pattern.
Ventrolateral glands: Five categories of ventrolateral gland extent were
established, ranging from no glands. A, to the entire belly, and most other
ventral surfaces covered with glands, E (Fig. 7). In addition, five categories
of gland color were noted in preserved animals: 1 — brown; 2 — gray; 3 —
black; 4— orange-brown; and 5 — -yellow-orange.
The geographic distribution of ventrolateral gland extent and coloration
in L. melanonotus is a mosaic. The ventrolateral glands are very compact and
brownish to orange-brown (B — 1,4) in populations from northwest Mexico
and Panama. The greater degree of glandular extent is found in populations
from the Isthmus of Tehuantepec, Yucatan Peninsula, Atlantic coastal British
Honduras and Honduras, and the Arenal pass region in Costa Rica. These
generalities are complicated in that individuals from a single locality 1 ) have
or lack glands and 2) demonstrate a range of variation of glandular extent
which encompasses the range of the entire species.
Most populations of L. wagneri either have no ventrolateral glands or
have orange-brown diffuse glands of moderate extent (C — 4). A few wagneri
specimens from the southern part of their geographic range have more ex-
tensive glandular extent. Where moderate samples (10 or more individuals)
are available, the specimens at any single locality demonstrate categories A
and C — -4. There is no geographic trend.
Many individuals of L. podicipinus lack ventrolateral glands. When
ventrolateral glands are present, they are of moderate extent and orange-brown
in preservative (C — 4, D — 4). Individuals from a single locality usually en-
compass the range of variation found within the species. No clear geographic
trend is evident.
There are no correlations between ventrolateral gland extent and annual
rainfall, number of dry months, or altitude in L. melanonotus and podicipinus.
A correlation is present between greater ventrolateral gland extent and greater
annual rainfall in wagneri (r = .13, P = .05). Such a correlation is more sug-
gestive of a causal relationship than the correlation between ventral color
pattern and ventrolateral gland extent in wagneri. There are no correlations
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Figure 8. Posterior surface of thigh color pattern standards. Note that pattern D is
restricted to Leptodactylus pustulatus.
1970 Studies on the Frogs of the Genus LEPTODACTYLUS
29
between ventrolateral gland extent and number of dry months or elevation in
wagneri.
Changes of extent of ventrolateral glands may occur seasonally, but there
are no collections made at all seasons from a single locality. In the summer of
1967 when breeding was just beginning, Dr. James R. Dixon and I collected
several series of L. melanonotus from western Mexico. We noticed that no
glands were evident on living animals. After a few days in preservative, how-
ever, the ventrolateral glands on the animals were dark brown in color and
distinct. The function of ventrolateral glands is not known.
Color pattern of posterior surface of thigh'. Four categories were dis-
tinguished: A) mottled; C) with a light longitudinal stripe; B) intermediate
between A and C; D) with distinct light spots on a dark background (Fig. 8).
Almost all L. melanonotus have a mottled posterior thigh, although a few
individuals from scattered localities show some indication of a light stripe
(category B).
In large series of L. wagneri from one locality, character states A, B, and
C are represented. This is true throughout the geographic range of wagneri,
with the exception of the Cauca Valley (Colombia) population.
Few large samples from L. podicipinus demes were available for this
study, but, on the basis of the series available, the distribution of the thigh
pattern appears to parallel the situation in wagneri. That is, categories A, B,
and C are represented within any deme.
There is no correlation between the posterior thigh pattern and the
climatic and elevational parameters used. It appears likely that the light stripe
in L. wagneri and podicipinus is genetically controlled by very few genes,
perhaps one. The scattered B categories in melanonotus populations indicate
that the gene potential for a posterior thigh light stripe is present. The D
category is found only in pustulatus.
The light stripe on the posterior thigh is another feature that shows
character displacement in the instances of sympatry between L. podicipinus
and wagneri. At both sites of known sympatry, wagneri individuals have well
developed stripes. The podicipinus at the sites of sympatry either have a mot-
tled thigh (Buenavista, Bolivia), or only an indication of a light stripe (Forte
Principe da Beira, Brasil). At this latter locality, the differences in thigh pat-
tern are very striking. The light stripe in wagneri individuals is completely
bordered with black and is very distinct from the thigh pattern of sympatric
podicipinus.
Toe-disk size: Three categories were used (Fig. 9): A) toe tips not
expanded at all; C) Toe tips expanded into definite disks; B) intermediate
condition.
There is a gradient of toe-tip size in L. melanonotus. In the northern part
of the range, Mexico to Costa Rica, most are in category A, only 3 demes
among those examined within this area had individuals with B condition. Many
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Figure 9. Toe tip standards. Pattern D is the upper disk surface of Leptodactylus
discodactylus showing longitudinal grooves.
individuals from Costa Rica have intermediate disks, B. In the South, from
Panama through Ecuador, all populations have B disks. In no case is the toe
tip expansion developed into a distinct disk as found in L. discodactylus.
A distinct geographic gradient in toe tip expansion is also seen in L.
wagneri. The Cauca Valley (Colombia) population is marked by non-ex-
panded toe tips. The rest of the Andean slope and upper Amazonian popula-
tions of Colombia, Ecuador, Peru, and Bolivia have categories A and B, with
B predominant. The frogs associated with the rivers draining northward, those
upon the islands, and probably those of the eastern Amazon region are either
B or C; that is, many individuals from these areas have distinct toe disks such
as are found in L. discodactylus.
No distinct geographic gradient in toe tip expansion is evident in L. podi-
cipinus. Most specimens lack expanded toe tips. A, but a few individuals from
scattered demes have slightly expanded toe disks, B.
The possible role of character displacement in toe-tip expansion is evident
at the two sites of sympatry between L. podicipinus and wagneri. At both sites,
the toe tips of podicipinus are not expanded. A, and the toe tips of wagneri are
slightly, but noticeably, expanded, B.
Mating call: Mating call samples were chosen to answer the specific status
concerning L. occidentalis. Demes were sampled in lowlands throughout the
latitudinal geographic range of occidentalis from Sonora through Nayarit.
Lowland samples of melanonotus from Jalisco, Colima, and Michoacan were
1970 Studies on the Frogs of the Genus LEPTODACTYLUS
31
Figure 10. Mating calls of Leptodactylus melanonotus in western Mexico. Vertical
scale in kilocycles per second, horizontal scale in seconds. Arrows indicate dominant
frequency band. A — Cali specimen from Sinaloa, 36 km S Los Mochis turnoff on
Mexico Highway 15. B-— Call specimen from Jalisco, 0.5 km NE Tonila. C — Call
specimen from Jalisco, 7.7 km E La Huerta.
A
8—1
also taken. Demes along elevational transects were sampled in the states of
Jalisco and Nayarit.
All of the calls have a basic similarity when heard and when viewed on
sonagrams. All calls consist of quite short notes, with a very wide frequency
range. The call components are difficult to compare with calls of other species
in which the components are very distinct. I consider the dominant frequency
the darkest, widest portion of the sonagram of the call, as substantiated in
sections, measuring the call duration at that point (Fig. lOA). Any other
noticeably prominent frequency below the dominant is taken as the funda-
mental frequency (Fig. lOB). Any periodic emphasized frequencies above
the dominant are considered harmonics, even though they may not be sharply
differentiated (Fig. IOC). The duration of the call is from .04 to .09 seconds,
the range .05 to .08 seconds being common in the call of a single individual.
Call duration does not correlate with temperature in the samples at hand. The
most striking differences among the calls are: 1) one or two notes per call
group; 2) the fundamental frequency is dominant or not; 3) the dominant
frequency values; 4) whether harmonics above the dominant are distinct or
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not; 5) whether the note is straight on the paper, or components are displaced
to the right.
Most of the calls are characterized by having a single note per call group.
Only in the state of Nayarit did individuals have calls consisting of two notes
per call group. One of the calls recorded at Tepic has a call consisting of both
one and two notes, as does the individual recorded from near San Bias. The
specimen recorded from Santa Cruz has a call consisting of two notes.
The calls recorded from near Acatlan, Jalisco, Tepic, Nayarit, and Her-
mosillo, Sonora, show the fundamental to be the same as the dominant. One
call from near Los Mochis, Sinaloa, has a fundamental distinct from the
dominant; another call has the fundamental equal to the dominant. All other
calls have fundamentals distinct from the dominant. Where fundamental fre-
quencies are present, they approximate one half the frequency of the dominant.
The dominant frequencies usually range from 2000 to 2500 cycles per
second. A range of 2000 to 2400 cycles per second is recorded in the deme
from Tepic, Nayarit. The only dominant differing from the range is from a call
recorded from Santa Cruz, Nayarit, which has a dominant of 3000 cycles per
second. The higher frequency can not be accounted for by differences in
temperature or elevation.
Three categories are used to distinguish whether harmonics above the
dominant are evident: 1) not present; 2) some indication of presence; 3)
present. Calls from Colima, Michoacan, and near La Huerta and Tecalitlan,
Jalisco, have harmonics. The call from near Tonila, Jalisco, has some indica-
tion of harmonics. The calls from near Acatlan, Jalisco, show all three states.
The calls from Tepic, Nayarit, either lack harmonics, or show some indica-
tions of harmonics. The calls from Santa Cruz, Nayarit, show harmonics
present or absent. All calls recorded north of Santa Cruz lack harmonics above
the dominant frequency.
Notes with certain components displaced to the right on the sonagram
are found in calls from localities that also have straight notes on the sonagram.
These localities demonstrating both types of notes are near Los Mochis, Sina-
loa, Santa Cruz, Nayarit, and near Acatlan, Jalisco.
Two of the call characteristics are gradually clinal in nature; the relation
of the fundamental to the dominant, and the presence or absence of harmonics
above the dominant. Two of the call characteristics are largely variable within
demes, and not constant on any geographic basis: number of notes per call
group; and whether the call is displaced to the right on the sonagram or not.
A call from a single deme shows a higher dominant frequency than the others.
Discussion : Physical size appears to be a major, pre-mating isolating
mechanism between L. melanonotus and wagneri. Additional collecting around
the northern end of the Andes in Colombia is necessary to determine the
interactions or lack of interactions between these two species.
Character displacement is most noticeable in sympatric demes of L. podi-
1970 Studies on the Frogs of the Genus LEPTODACTYLUS
33
cipinus and wagneri, where respective adult sizes, ventral color patterns,
posterior thigh color patterns, and toe tip expansions are very distinct. Ex-
pression of certain of these characters are similar in other populations of
podicipinus and wagneri. Ventral color pattern in particular shows the effects
of character displacement. Certain demes along the slopes of the Andes in
Bolivia and southern Peru have ventral color patterns that closely approach
the pattern of some podicipinus. The ventral patterns of wagneri and podici-
pinus are distinct at points of sympatry; the specimens do not exhibit gradation
of ventral pattern D to E.
The status of the population of frogs in northwest Mexico (L. occidentalis
Taylor) has been particularly uncertain. The main characters used to dis-
tinguish the population are adult size and extent and color of the ventrolateral
glands. The size of the population fits into a larger pattern discussed above and
is not peculiar to a series of demes in northwest Mexico. The ventrolateral
gland condition (B) is characteristic of the population in northwest Mexico,
but the same condition is found in certain demes in Panama. The mating call
data indicate that the extreme northwestern demes are differentiated from the
lowland demes in Colima and Michoacan, but that the differences are con-
nected by a series of demes demonstrating intermediate call types. The low-
land demes in Colima and Michoacan are separated geographically from the
highland demes of Jalisco and Nayarit by terrain with virtually no standing
water; water is either cascading or nonexistent. The population from north-
west Mexico appears to be isolated at present from the coastal demes of Colima
and southward. Morphological differentiation of the northwest Mexican pop-
ulation is slight, and the same differentiation is repeated in other demes of
L. melanonotiis throughout its geographic range.
The island demes of L. wagneri are in no way distinguishable from main-
land wagneri', in fact, the presence of expanded toe disks and the adult size
indicate that the demes originated from the population found in northern
Venezuela and Guyana.
I follow an earlier statement on the value of the subspecies concept
(Savage and Heyer, 1967). I believe the methods treating variation within a
species, as here presented, reveal more of the populational trends than does
the arbitrary practice of designating subspecies. It is nearly impossible to
fragment the three widespread species on the basis of any combination of
characters.
It is appropriate to interject a comment on my evaluation of the tax-
onomic decisions at this point. Most of the study was based on morphology; I
did not have the benefit of any biological data for most of the specimens; I
regard the taxonomic decisions as conservative. With the evidence available
for this study, my best judgment is that the Melanonotus species group is
composed of six species.
Relationships'. Four characters shared by members of the Melanonotus
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No. 191
group are useful in determining the degree of evolutionary advancement of
the species. More adequate justification for determining primitive and ad-
vanced character states is presented elsewhere (Heyer, 1968). The presence of
toe disks is considered an advanced condition, and the presence of grooves
on the upper disk surfaces is considered a further advancement. Absence of
thumb spines in males, a striped or distinctly spotted pattern on the posterior
thigh surface, and a ventral pattern of light spots on a dark background are
considered advanced conditions.
Certain trends among the species are clear. Leptodactylus melanonotus
is primitive in all four characters, and is probably most like the ancestral
Melanonotus group stock which gave rise to the six extant species. Lepto-
dactylus discodactylus and pustulatus are the most specialized. Leptodactylus
discodactylus has the most advanced disks, and the males lack thumb spines.
Leptodactylus pustulatus has an advanced ventral and thigh pattern, and the
males also lack thumb spines. Leptodactylus dantasi, podicipinus, and wagneri
are intermediate. Leptodactylus dantasi has the advanced ventral pattern.
The male is unknown for this species. Leptodactylus podicipinus and wagneri
have individuals with the advanced thigh stripe. Leptodactylus podicipinus
has a spotted belly and wagneri has individuals with toe disks. Based on only
these four characters, a graphic representation is very difficult as evolution
appears to have produced a mosaic pattern. Leptodactylus melanonotus,
podicipinus, and wagneri are the most closely related. I think dantasi and
pustulatus are more closely related to these three species than they are to
discodactylus. Leptodactylus dantasi, pustulatus, and podicipinus resemble
each other in general appearance, and the distinctive patterns of dantasi and
pustulatus could have been derived from a species very much like that of
podicipinus. For the present, I regard the loss of thumb spines to be a case of
parallel loss in discodactylus and pustulatus. The relationships are presented as
a dendrogram (Fig. 11).
Distributional Analysis
Although some areas of Latin America have been studied moderately
intensively, the geology, climate, soil types, and vegetation of many parts of
Middle and South America are known poorly or not at all. Savage (1966) has
recently given an excellent synthesis on the origins and history of the Central
American herpetofauna, but no such synthesis has been attempted for South
America. Only the broadest generalizations can be made with respect to the
distribution of the components of the Latin American herpetofauna and the
following remarks must be considered as provisional.
To discuss the distribution of the Melanonotus group, it is necessary to
review briefly the broad geographic, paleogeographic, climatic, and vegeta-
tional patterns of Latin America.
Geography: The great American Cordillera forms a distributional barrier
DISCODACTYLUS
1970 Studies on the Frogs of the Genus LEPTODACTYLUS
35
Figure 11. Proposed phylogeny of members of the Melanonotus group.
MELANONOTUS
36
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No. 191
to frogs of the Melanonotus species group; the upper elevational limit for any
species is about 1800 meters. Breaks in the montane chain where faunal inter-
change occurs are: the Isthmus of Tehuantepec in Mexico; the region around
Lake Managua in Nicaragua; the Arenal pass in Costa Rica; and an extensive
area including most of Panama and northernmost Colombia. The Andes of
South America pass along the western outline of the continent, abut against the
sea, or leave a very narrow lowland coastal strip to the west.
In South America, there are three major upland regions in addition to
the Andes. These are the Guiana shield, the central Brasilian shield, and the
coastal Brasilian shield. These upland regions do not form an elevational
barrier to the distribution of frogs of the Melanonotus group. The Guiana
shield is somewhat continuous from eastern Colombia, extending through
central and southern Venezuela, to and including almost all of the Guianas
and part of Brasil just below the Guianas. The central Brasilian shield is of
irregular outline bordering Bolivia on the west and extending as a band
eastward along the southern edge of the Amazon basin. The coastal Brasilian
shield extends from the easternmost tip of South America in the state of Rio
Grande do Norte along the coast southward to the state of Sao Paulo.
Paleogeography : The Cordillera of Latin America is geologically young,
its present form attained in the very vigorous orogenesis from Mid to Late
Tertiary, especially in Miocene and Pliocene times. Orogenesis continues in
some parts of the Cordillera. (For a more detailed account and further refer-
ences, see Childs and Beebe, 1963; Harrington, 1962; and Savage, 1966).
The Guiana, central Brasilian, and coastal Brasilian shields date from the
Early Precambrian and have probably been land positive since then (Harring-
ton, 1962) . The history of the present land areas amidst the shields is poorly
known, particularly since Mid-Tertiary to present, comprising, probably, the
whole of the history of the Melanonotus species group. The Amazon basin has
had a recurrent history of continental sediment deposition and marine embay-
ment. Continental deposition has definitely occurred in the Late Tertiary, but
whether any marine embayment alternated with continental deposition during
or following this time is not known (Harrington, 1962) . No precise informa-
tion on the isolation or continuity of the shields with relation to each other
exists, and it is impossible to comment on the isolation of the ancestral stocks
of the frogs of the Melanonotus species group.
Climate: Comments are limited to climates affecting the distribution of
members of the Melanonotus group. The coasts of Mexico and Middle Amer-
ica are characterized by seasonal rainfall patterns. A very pronounced hot and
dry season is found along the western Mexican coast southward to the middle
of Costa Rica. Similar conditions are found along the east coast of Mexico
south to Nicaragua. The Atlantic coastal area of Costa Rica is wet the year
around, but a seasonality in rainfall occurs. The only place in Middle America
with a nearly continuously wet tropical rainforest is the Osa peninsula region
1970 Studies on the Frogs of the Genus LEPTODACTYLUS
37
of Pacific Costa Rica. A wet climate with no cool season and basically no dry
season occurs in four restricted areas in South America. The first such area is
the Choco region along the western coast of Colombia extending into north-
west coastal Ecuador. The second and largest region is amidst the upper
tributaries of the Amazon in southern Colombia, eastern Ecuador, most of
eastern Peru, and northwestern Brasil. The third extends from coastal Guianas
to the Brasilian coast just south of the mouths of the Amazon. The fourth area
extends along a sometimes very narrow coastal strip of eastern Brasil from
the states of Bahia south to Rio de Janeiro. All the other regions in South
America occupied by the Melanonotus group have a pronounced dry season
from one to nine months in duration. The driest regions in South America, also
four, appear to limit the distribution of the Melanonotus group. The first runs
along the western lowlands of middle Ecuador southward the length of the
western lowlands of South America. The second lies along the north coast of
Colombia and Venezuela, penetrating inland various distances. The third is in
northeastern Brasil including the states of Maranhao, Piaui, and Ceara. The
fourth is the Gran Chaco of northern Argentina, southeastern Bolivia, and
southwestern Brasil (James, 1959; see also Walter and Lieth, 1960-1967).
Vegetation: The vegetation of Latin America is very complex and very
inadequately studied. Many different systems have been used to define and
describe the vegetation of Latin America. For the present I use the broad
vegetational maps found in James (1959) and Wagner (1964) as the basis for
discussion.
True tropical rainforests are found in the Osa peninsula region of Costa
Rica; along western coastal Colombia and northwestern coastal Ecuador; along
upper drainages of the Amazon characterized by high, continuous rainfall;
and along the narrow coastal strip of Brasil from the states of Rio Grande do
Norte to Rio de Janeiro. Tropical and subtropical rainforests which have a
definite seasonal rainfall distribution are found in a continuous belt along the
east coast of Mexico through Panama. Coastal patches of deciduous forest are
found within the tropical and subtropical rainforests along the Carribean
coast of Mexico, the tip of the Yucatan peninsula, easternmost British Hon-
duras, easternmost Honduras and northeastern Nicaragua. Seasonal tropical
rainforest is continuous with the tropical rainforest of the upper Amazon
and covers almost all of the Amazon basin not covered by the tropical rain-
forest. Tropical deciduous forests are found along the west coast of Mexico
from the state of Jalisco southeast to the middle of the Pacific coast of Costa
Rica; along the periphery of the Guiana shield; in eastern Brasil from the
north coast of the state of Rio Grande do Norte to middle Bahia, excluding the
narrow eastern coastal strip; and in the Gran Chaco. Thorn forests are found
along the northern coasts of northeast Colombia and Venezuela. Savannas
cover the Guianan and central Brasilian shields, and most of the coasts ad-
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No. 191
jacent to the Brasilian shields. A palm forest is found in northeast Brasil in the
states of Piaui and Maranhao.
Distribution of the Melanonotus Species Group: Most of the species are
known from scattered localities. Leptodactylus melanonotus has been most
thoroughly sampled, and moderately accurate distributional data are available
for at least Mexico and Costa Rica. Records for dantasi, discodactylus, and
pustulatus are very spotty, and more distributional data are needed to define the
geographic limits of the species. Leptodactylus podicipinus has been moder-
ately well collected in Argentina, but inadequate distributional data are avail-
able for the rest of its geographic range. Distributional data of wagneri are most
adequate for Ecuador, parts of Colombia and Guyana. Throughout the rest of
the geographic range of wagneri, records are so spotty that it is impossible to
determine whether it has a broadly continuous range or if it is composed of
many isolated populations.
The limited information gives the following patterns of occurrence :
Leptodactylus melanonotus, wagneri, and podicipinus form a series of
closely related, basically allopatric species ranging throughout low and mod-
erate elevations of tropical Latin America. All three species are found in areas
having a definite dry season. Judged by collections, melanonotus is not common
along the continuously wet lowlands of Colombia and Ecuador. Only wagneri
is commonly associated with the tropical rainforests of eastern Colombia,
Ecuador, and Peru. The foam nest formed by these species appears to be an
adaptation to a definite dry season.
In some more xeric portions of its range (southern Sonora and northern
Sinaloa, Mexico), melanonotus is found only in association with permanent
bodies of water, such as persisting springs or irrigation water. The northern-
most locality for melanonotus on the west coast of Mexico is a dam seepage by
Hermosillo, Sonora. In more mesic areas, it is often associated with tem-
porary bodies of water, such as rain filled ditches along roadsides.
Leptodactylus melanonotus, podicipinus, and wagneri are found in re-
gions with old land masses in South America: melanonotus with an old land
mass in western Colombia and Ecuador (land positive at least during Late
Mesozoic and Tertiary time, Harrington, 1962:1780) ; podicipinus with the
Brasilian shields; and wagneri with the Guiana shield. If one assumes that the
three species had a common, widespread ancestor, isolation of the land masses
in the Cenozoic could account for the fragmentation of the stock that gave
rise to melanonotus, podicipinus, and wagneri.
Individuals of melanonotus in the region from the Isthmus of Tehuante-
pec to the Arenal Pass of Costa Rica are characterized by having a larger size,
a more variable ventral pattern, and a greater ventrolateral gland extent than
individuals to the north or south. Such character gradients suggest that the
major dispersal point of melanonotus has been from this land bloc in Middle
America. Secondary dispersal in Middle America by a species which had its
1970 Studies on the Frogs of the Genus LEPTODACTYLUS
39
evolutionary origin in South America has also been hypothesized for the hylid
frog Phrynohyas venulosa by McDiarmid (1968). Dispersal into Middle
America by a species from western South America is consistent with Savage’s
(1966) scheme of historical herpetofaunal dispersal. He suggested that an
assemblage of species, termed the Western South American Herpetofaunal
Complex, evolved during the separation of nuclear Middle America from
South America from Eocene through Miocene. When Middle America and
South America were joined again at the beginning of the Pliocene, members
of the Western South American Complex invaded Middle America.
Leptodactylus wagneri today is found in a variety of climates and
associated vegetation types, but is found consistently in more mesic situations
than either melanonotus or podicipinus. Throughout most of its range, wag-
neri is found in tropical or subtropical rainforest with either a short or no dry
season. An exception occurs in the northern part of the range, particularly
along the extreme north coasts of Colombia and Venezuela. Leptodactylus
wagneri is not recorded from the palm forest or tropical deciduous forest of
eastern Brasil in the states of Maranhao through Rio Grande do Norte. Rec-
ords are so scattered for eastern Brasil that it is hard to tell whether its absence
in any region is a collecting artifact.
The islands of Trinidad and Tobago are on the continental shelf and were
probably connected to the mainland during glacial maxima. The presence of
wagneri on these islands is to be expected. The islands of Grenada, Bequia, and
St. Vincent are not on the continental shelf; they probably have been always
isolated from the continent of South America. Grenada and Bequia are on the
same bank. Currently, Bequia lies only 10 km from St. Vincent and possibly
was separated by no more than 5 km of water during glacial maxima. The
distance from the 100 fathom contour outside of Tobago to the 100 fathom
line surrounding Grenada is at most 100 km, and these two points are directly
connected by westerly surface ocean currents. All island populations are
indistinguishable from mainland populations of northeast Venezuela, suggest-
ing either that dispersal to the Lesser Antilles has been a rather recent event
or that Lesser Antillean populations have been populated repeatedly from
the mainland stock. A single dispersion of wagneri from South America (in-
cluding Trinidad and Tobago) could account for all of the populations on the
Lesser Antilles. Other members of the genus Leptodactylus are more success-
ful island colonizers, especially mystaceus which has apparently hopped from
island to island along the Lesser Antilles to as far as Haiti. As judged from the
similarity of the island and mainland populations, it is reasoned that wagneri
dispersed to the Lesser Antilles during the glacial maxima of the Pleistocene.
Leptodactylus podicipinus is associated with the southern river drainages
and the Sao Lrancisco drainage system. All of the regions, such as the Gran
Chaco, where podicipinus has been found have a very distinct dry season.
L. podicipinus is generally adapted to more xeric conditions than wagneri.
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No. 191
This may be seen where the two species occur close together, most notably
with wagneri along the thin mesic coastal strip of east Brasil, and podicipinus
close by in the dry parts of the Sao Francisco drainage system.
Leptodactylus dantasi, discodactylus, and pustulatus are known from
spotty distributional records, which suggest a distribution associated with
climatic and vegetational factors. Leptodactylus dantasi has been taken only
from the tropical rainforest in western Brasil. Leptodactylus discodactylus
is apparently limited to the tropical rainforest of southeastern Colombia,
eastern Ecuador, most of eastern Peru, and western Brasil. Leptodactylus
pustulatus is apparently limited to the more xeric conditions in the palm
forest, tropical deciduous forest, and savannas of eastern Brasil.
The total distribution of members of the Melanonotus species group
indicates that three species have broad ecologic tolerances and are distributed
in extensive geographic regions and that three of the species have much
narrower ecologic tolerances and have restricted geographic distributions.
Specimens Examined
Leptodactylus dantasi (N = 1): BRASIL. TERRITORIO DO ACRE:
Feijo, 249 m — WCAB 1240 (Holotype).
Leptodactylus discodactylus (N = 15): ECUADOR. NAPO: Payamino,
304 m— GOV 7238; PASTAZA: Puyo, 1000 m— GOV 2062, JAP 6197-8;
6333; Veracruz, 1000 m—JAP 6151, 6153-6, 6164. PERU. LORETO: Es-
tiron, Rio Ampiacu, 200 m — CAS 93316, 7; Rio Itaya, 150 m — AMNH
43217; Rio Tamaya, Sobral, 150 m— AMNH 42680.
Leptodactylus melanonotus (N = 520) : BRITISH HONDURAS.
STANN CREEK: Mango Creek, 120 m— MCZ 37884; 4.8 km S Waha Leaf
Creek, 120 m— MCZ 37875-6. COLOMBIA. CHOCO: Rio San Juan, Playa
de Oro, 400 m — FMNH 54374; NARINO: La Guayacana, 230 m — PAS
44; Rio Mataje, 200 m — USNM 147393; Valle de Cauca: Rio Raposo Vir-
ology Field Station, 20 m— USNM 151463. COSTA RICA. Alajuela: Cari-
blanco, 800 m — CRE 2928 (10); 0.5 km S Cariblanco, 820 m— CRE 694;
Los Chiles, 70 m— AMNH 54638, 54641-3, CRE 7217, 7219; Punta Cortez,
70 m— CRE 7218; CARTAGO: Cervantes, 1441 m— KU 28189; IICA, Tur-
rialba, 600 m— CRE 575, 2808 (2), 7074 (2), 7079 (7); GUANACASTE:
Arenal, 520 m— CRE 6251, 6258 (4), 6259, 6260 (2) ; 4.8 km NW Arenal—
CRE 2901 (3) ; Bebedero, 6 m— CRE 7162; Canas, 88 m— CRE 7164; 9.6 km
S La Cruz — CRE 8091 ; Rio Lagarto at Inter. Am. Hiway, 100 m—CRE 7122
(2) ; 2 km W Liberia — CRE 728 (4) ; 9 km N and 4 km E Liberia on Inter.
Am. Hiway on Rio Colorado — CRE 714 (2); 14.8 km S Liberia, 90 m — CRE
8168; 45 km SW Liberia, Hwy #21— CRE 8216; Hacienda La Norma, 5 km
N and 4.5 km W Liberia on Rio Colorado, 150 m—CRE 105 (2), 106 (6),
107, 251 (2); 3-11 km E Playa del Coco, 45 m—CRE 8012 (2); Finca San
Bosco de Tilaran, 640 m — CRE 6273, 6278; 3.2 km W Santa Cruz on the
1970 Studies on the Frogs of the Genus LEPTODACTYLUS
41
Playa del Tamarindo rd. — CRE 8233; Santa Cruz, 50 m — CRE 7128, 8232
(3); Silencio de Tilaran, 825-850 m— CRE 6217 (8), 6225 (2), 6228 (2);
0.5 km NW Tilaran, 530 m— CRE 520 (7); 3 km NE Tilaran— CRE 524; 5
km NE Tilaran, 600 m — CRE 8021; 5.5 km NE Tilaran, 560 m — CRE 8022;
6 km NE Tilaran, 550 m — CRE 523 (3), 8020 (9) ; Finca Taboga, 9.6 km S
and 8.0 km W Canas, 4 m — CRE 2902; HEREDIA: Puerto Viejo, 100 m —
KU 65683; 6.5 km W Puerto Viejo— KU 91798; 7.5 km W Puerto Viejo— KU
86293; LIMON: Batan, 15 m — KU 34146-51 ; 2.4 km E Los Diamantes,
260 m— CRE 8049; El Tigre, 680 m— CRE 290 (2); PUNTARENAS: 9.6
km ESE Golfito, 10 m — CRE 7105; junction of road and rail line (Base of
Peninsula), 5 m— CRE 238 (2), 253-4. ECUADOR. ESMERALDAS: San
Javier, 200 m— JAP 9077-82; Cacnaui, 20 m— JAP 2889-9; Rio Bogota, 200
m— GOV 6398; LOS RIOS: Vinces, 150 m— UMMZ 55587 (7). EL SAL-
VADOR. SAN SALVADOR: San Salvador, 620 m— FMNH 65071-86.
GUATEMALA. EL PETEN : N of La Libertad— UMMZ 75367 (12); 5.2
km S La Libertad — MCZ 21454; Toocog, 15 km SE La Libertad, 170-200 m
— KU 58959; ESCUINTLA: Rio Guacalate, nr. Masagua, 100 m— USNM
125246-51, 125253, 125261-3; ZACAPA: 8 km ENE Rio Hondo, 175 m—
KU 58960; 23 km NE Zacapa— ANSP 22165-6; 23 km W Zacapa,.240 m—
TCWC 16403, 16406. HONDURAS. ATLANTIDA: Corozal, 15 km E La
Ceiba, 10 m— JRM 2316, 2328-33, 2335, 2337-8, 2341, 2343; CHOLU-
TECA: 1.5 km NW Choluteca, 170 m— KU 65676; 5 km S Choluteca, 80 m
—MCZ 26460-2; CORTES: Agua Azul, Laguna Yogoa, 700 m— AMNH
54778; FRANCISCO-MORAZAN: 31 km S Guaimaca, Santa Clara, 1000
m— AMNH 54884-5, 54887; ISLAS DE LA BAHIA: Isla de Guanaja, SE
shore opposite Guanaja, 10 m — JRM 1938, 1961-6; Isla de Roatan, .5-4 km
NRoatan, 10-30 m— JRM 1828-9, 1896, 1916; OLANCHO: Escuela Nacional
de Agricultura, 4.5 km SE Catacamas, 400 m — JRM 1660-2. MEXICO. CAM-
PECHE: Chuina, 46 km S Champoton, 50 m— KU 75017-26; CHIAPAS:
4 km NW La Esperanza, 100-300 m— USNM 114372-6, 114381-2, 114385,
114388; Tonala, 55 m— USNM 114338-44; Tuxtla Gutierrez— UMMZ
115530; 9.8 km W Tuxtla Gutierrez, 520 m— UMMZ 119147 (2); COLIMA:
25.9 km N Manzanillo, 10 m— CAS 97111-8, 97155-7; GUERRERO: 12.9
km SW Tierra Colorada, 300 m— TCWC 8323-6, 8333-9; JALISCO: 5.2 km
W Acatlan, 1400 m— LACM 37381-408; MICHOACAN: Apatzingan, 540
m— FMNH 38835, 38837, 38853-5, 38859-60, 38862, 38865-6, 38873; NA-
YARIT: San Bias, 3 m— UMMZ 110892 (7), 115543 (3); Tepic, 1000 m—
LACM 37042-3; OAXACA: Cacahuatepec at river, 150 m— UIMNH 52854-
8; 8.3 km N Pochutla, 100 m— UMMZ 123982 (8) ; 5.2 km N Matias Romero,
150 m— UMMZ 115522 (3); 7.8 km S Matias Romero, 150 m— UMMZ
113804 (4); Tehuantepec, 35 m— FMNH 123248-50, 123253, 123256,
123264, 123267-9; Tolosa, 50 m— AMNH 53610; QUINTANA ROO: 3 km
E Caobas, 5-10 m— KU 75033; 4 km WSW Puerto Juarez, 5 m— KU 71117-
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No. 191
8; SAN LUIS POTOSf: 5.1 km NE Tamazunchale, 150 m— TCWC 7285-91;
5.1 km SW Tamazunchale, 150 m— UIMNH 16802-3; SINALOA; Culiacan,
45 m— AMNH 58382-91; 18.9 km SW Mazatlan, 48 m— KU 78259-61; 5.2
km W Villa Union— KU 29901-7; SONORA; Alamos, 300 m— AMNH
51356-65; Hermosillo, 28 m— AMNH 62737, MVZ 26066; TABASCO;
Teapa, 40 m— UMMZ 119148 (2); 20.7 km N Teapa— UMMZ 119901 (2);
33.7 km N Teapa— UMMZ 119903 (6); 57 km N Villahermosa, 30 m—
AMNH 69007-8; TAMAULIPAS; Pano Ayuctle, 12.9 km NE Gomez Farias,
100 m— UMMZ 98947 (2), 101185 (5), 102263, 102912; VERACRUZ; 5
km SW Boca del Rio— KU 23700; Cuatotolapam, 13 m— UMMZ 41640-3,
41646-7, 41649; 1 km E Mecayucan, 30 m— KU 24084, 24086, 24091-4;
25.9 km WNW Temapache, 50 m— KU 60225-8; YUCATAN; 31 km N
Tizimin, 5 m — CM 40108-10; 16.2 km S Yaxcopoil, Aguada Xcamal, 5 m — -
CM 45233-43; Yuncu, 25-50 m— UMMZ 73155, 73158 (2). NICARAGUA,
MANAGUA; 15.5 km NW Managua, 55 m— KU 43030-39; RIVAS; San
Jorge, Shore of Lake Nicaragua, 50 m — MCZ 28980-89; ZELAYA; Bonanza,
260 m— KU 84839-41, 84843-48; 10.4 km SW Bonanza, 300 m— KU 84842;
25.9 km above Recreo, Rio Mico, 30 m— UMMZ 79734, 79735 (4). PAN-
AMA. CHIRIQUI; Las Lagunas, ± 6 km W El Volcan, ± 1200 m— KU
76522; Finca Palosanto, 6 km WNW El Volcan, 1230 m — KU 76520-1;
COCLE; El Valle, 560 m— KU 76523-32; DARIEN; Rio Canclon, 60 m—
UMMZ 125008, 125009 (10). UNITED STATES, TEXAS; Starr County,
5.1 km SE Rio Grande City— UIMNH 29817.
Leptodactylus podicipinus (N = 154) ; ARGENTINA. CHACO; Resis-
tencia, 60 m— KU 84732-4; CORRIENTES; 30 km W Itati, 60 m— CAS
100504; Manantiales, 50 m — MCZ 35589-90; FORMOSA; Esteros Laguna
Oca, 65 m — MCZ 32775-6; SANTA FE; Banados Monteira, 15 km N Santa
Fe, 20 m — CM 38011; Banados del Rincon, 20 m — CM 37981-3, 38003,
38005; Santa Fe, 18 m— CM 39181, 39194. BOLIVIA. BENI; Ivon, 150-200
m— UMMZ 64099 (7) ; Puerto Almacen, 260 m— AMNH 72245-53; SANTA
CRUZ; Buenavista, 500 m— AMNH 39538; El Carmen, 212 m— CM 36234-
7; Naranjal, 5 km from Montero, 350 m — USNM 146551 ; El Porton, 550 m
—CM 36248; Robore, 330 m— MCZ 30032-6; 3 km S Robore, 330 m— CM
36238-41; San Jose de Chiquitos, 350 m— CM 36164, MCZ 30028-9. BRA-
SIL. BAHIA; Barreiras, 500 m — UMMZ 109986; Rio Grande at Sao Jose,
450 m— UMMZ 109989; MATO GROSSO; Agua Clara, 304 m— FMNH
67088; Conceicao, Paraguay River, 150 m — FMNH 9161; Corumba, 115 m
—CM 2429, 36165, UMMZ 104225 (2); 40 km from Corumba, Urucum
Mts., 200-300 m— CM 2430; Descalvados, 142 m— FMNH 9097, 9100-14;
1 day’s run below Descalvados, 150 m — USNM 132736-7; Maracaju, 385 m
—USNM 107705-8, 107710-11; Porto Esperan9a, 120 m— UMMZ 104232
(11), USNM 133005-8; Salobra, 190 m— UMMZ 104227 (2); MINAS
GERAIS; Januaria, 454 m— UMMZ 109984; Pirapora, 472 m— UMMZ
1970 Studies on the Frogs of the Genus LEPTODACTYLUS
43
109982 (10), USNM 98535, 99801-3; PARANA: Andira, 480 m— MCZ
32725-6; RONDONIA: Forte Principe da Beira, 100-200 m— WCAB 8330,
8982; Porto Velho, 60 m— KU 92933-37; SAG PAULO: Anhembi, 500 m—
WCAB 30880-1; Bauru, 499 m — DZ 53-4; Cachoeira de Emas, Pirassununga,
650 m — DZ 4636-42; Lins, 396 m — DZ 9024; Porto Marcondes, Parana-
panema, 250 m— =DZ 19979, 19999 (10 total). PARAGUAY. CAAGUAZJU:
Pastorea nr. Caaguazu, 250 m — MCZ 17902-11; CORDILLERA: Villeta,
110 m— AMNH 50654; PRESIDENTS HAYES: Rio Pilcomayo, 38.8 km
W Rio Paraguay, 80 m--»MCZ 25827-33.
Leptodactylus pustulatus (N = 7) : BRASIL. GOIAS: Aruana, 200 m—
DZ 4992, 8109; MATO GROSSO: Sao Domingoes, Rio das Mortes, 200 m
-^DZ 986-7, 996, 1069; PIAUf: Rio Poti^MCZ 373.
Leptodactylus wagneri (N = 711): BOLIVIA. BENI: Cachuela Es-
peranza, 150 m — UMMZ 64102 (2); nr. Guauara-mirim, Rio Marmori,
160 m — CM 2666; Swamp along Rio Guapore, San Antonio de Guapore,
150 m—USNM 115973; Huachi, 250 m"'— UMMZ 64103 (3), 64104;
Puerto Almacen, 260 m— AMNH 72251, 72254, 72398-402; Lake
Rogoagua, 200 m-^UMMZ 64100 (7); Rurrenabaque, 227 m— UMMZ
64097 (3); Villa Bella, 120 m~CM 2659; COCHABAMBA: Puerto
Chipiriri, 300 m“”AMNH 72242-3; PANDO: in front of Placido de Castro,
Acre, Brasil—DZ 6529-31; SANTA CRUZ: Buenavista, 500 m— CM 4331,
4353, UMMZ 64028-29, 64030 (2), USNM 118688-9, 146523-4. BRASIL.
ALAGOAS: Fazenda Canoas, Rio Largo, 60 m™— DZ 9279; Mangabeiras,
50 m--DZ 11982-3; Sao Miguel, 20 m---DZ 9250; AMAZONAS: Manaca-
puro, 50 m—USNM 103621-2; BAHIA: Salvador, 15 m— DZ 9131, 9550-1;
GOIAS: km 47 Estrada Goias- Aruana, 350 m-^DZ 1518; MARANHAO:
Carolina, 168 m— DZ 21664-5; MATO GROSSO: Dumba, 200 m--DZ 1448;
Banks of center of Rio Guapore, 250 m — CM 2508-9; Confluence of Rio
Araguaia and Rio Tapirape, 190 m- — AMNH 68093; Sao Domingoes, Rio das
Mortes, 200 m— DZ 10891-2, 10894-900, USNM 148677-91; PARA: Belem,
20 m-~-FMNH 83264-5, MCZ 36010-1, USNM 154068; Cachimbo, 500-600
m — DZ 21656-61; Rio Mapuera at the Equator, 80 m — AMNH 46181; San-
tarem, betwn Rio Tapajos and Rio Amazonas, 16 m — CM 2641; RIO DE
JANEIRO: km 31 on Estrada Mage-Estrada Rio Petropolis, 100 m — DZ
12034-5; RIO GRANDE DO NORTE: Natal, 120 m— AMNH 36261, MCZ
15847, UMMZ 68791 ; TERRITORIO DO ACRE: Iquiri, 150 m— DZ 6588-
92; TERRITORIO DO RIO BRANCO: Rio Urariquera, 100-200 m— WCAB
18792; TERRITORIO DO RONDONIA: Forte Principe da Beira, 100-200
m~-^WCAB 10690, 10703. COLOMBIA. AMAZONAS: Falls of Engano—
MCZ 28063; Leticia, 275 m— CM 36081-2, USNM 142179-86, 142187-94,
146252, 147031; ANTIOQUIA: Envigado, 1600 m— AMNH 39265, 39274,
39281; Medellin, 1500 m—AMNH 38785, 38808-15, 39465; Sonson, 2500
m-^AMNH 39631; BOYACA: Muzo, 1240 m— FMNH 69744, MCZ 24920;
44
Contributions in Science
No. 191
CAUCA: Popayan, 1700 m— FMNH 54377-83; CUNDINAMARCA: Ano-
laima, 1500 m— USNM 147054-5; Bogota, 2600 m~AMNH 13471; Fusa-
gasuga, 1700 m— AMNH 71581; La Mesa, 1000-1300 m-— USNM 144892-3;
META: Buenavista, 1200 m — MVZ 63067-73; Menegua, upper Rio Meta,
260 m— USNM 147273-4; 46 km S, 22 km W San Martin, 490 m— -MVZ
63076; Villavicencio, 500 m— ANSP 25765, FMNH 30571, 81788-9, MCZ
16277, MVZ 63074-5, 63738-9, UMMZ 74811, USNM 144848-66, 147056-
62; 7 km NE Villavicencio, 500-700 m— USNM 146382-4; NARINO: La
Guayacana, 100 m— FMNH 61754; NORTE DE SANTANDER: Astillero,
100-200 m— USNM 147063; E of Cucuta 200-500 m— ANSP 25758; PUTU-
MAYO: Rio Putumayo, 5 km N Puerto Asis, 260 m — PAS 47; Rumiyaco,
1000 m— FMNH 54376; SANTANDER: nr. Lebrija, 1086 m— USNM
144883-8, 146250; San Gil, 1095 m— CM 7938, UMMZ 74799; TOLIMA:
Guindio Mtns.— MCZ 8217; VALLE DE CAUCA: Buenaventura— USNM
147076; 2 km S Cali, 950 m— USNM 148800-23; VICHADA: Puerto Car-
reno— CM 17122. ECUADOR. NAPO: Lagarto Cocha, 200 m— GOV 9208;
Loreto, 400 m — GOV 9205, WCAB 36541 ; Rio Napo, Avila, 300 m —
UMMZ 92145-6; PASTAZA: Upper Bobonaza— JAP 8732; Canelos, 530 m
—MCZ 17950; Cerros de Abitagua, 1100 m— FMNH 25789, 26899, UMMZ
92147; Chichirota, 300 m— GOV 9207; Upper Rio Curaray — GOV 9210;
Upper Rio Oglan, 500 m— GOV 9206; Rio Pindo, 190 m— GOV 9195-7;
Puyo, 940 m— JAP 1958-71, 1973-2005, 2036, 2038, 6182, 6186-7, 6191-5,
6289-99, 6303-5, 6329-32, 6334, GOV 8021-2; Shell Mira, 1064 m— GOV
8503, JAP 2177; Don Tomas, 340 m— GOV 9209; Rio Villano, 300-400 m—
GOV 7731-6, JAP 3776; ZAMORA-CHINCHIPE: Copal, 910 m— JAP
6747-8; Limon, 1550 m— JAP 6937, 6964-5; Rio Llushin— GOV 9198-201,
9203, JAP 3751; Mendez, 580 m— JAP 2037, 6805, 6870, 6872-5, 6880,
6832-4, 6837, 6845-6, 6855-6, 6936; Plan de Milagro, 1700 m— JAP 7008,
7014-6, 7018-23, 7027, 7057; Plan Grande, 880 m— JAP 6903; NE of Rio-
bamba— FMNH 23509 (8) ; Sucua— JAP 2186, 2225. FRENCH GUIANA.
Sophie, on trail leading S from St. Elie Gold Pits, 200 m — MCZ 44564.
GUYANA. DEMARARA: Atkinson, McKenzie Trail, 50 m— USNM
162872-3; Dunoon, Demerara Riv., 20 m — UMMZ 50183-9; Georgetown, 5
m— UMMZ 80497; Wismar, 50 m— AMNH 45750, UMMZ 77517, 80417;
ESSEQUIBO: Isheraton, 250 m— AMNH 53435-7; Karanambo, 100 m—
AMNH 53438; Kartabo, 10 m— AMNH 10377, 10379-83, 11659, 13518-9,
25233, CM 5443-4, USNM 118059-62; Kuyuwini Landing, 250 m— AMNH
46281, 49352, 49355 (5); Lethem, 100 m— MCZ 50708; Marudi, 250 m—
AMNH 46238 (2); Moku Moku, 100-200 m— USNM 146366-7; Oko Mts.,
115 m — FMNH 26691 ; Santa Rosa Is., Monica Riv., 7 m — UMMZ 55833
(2), 80417; Shudikarwan, 200-300 m— AMNH 46274 (9). PERU. AYA-
CUCHO: Candalosa, ± 1400 m— FMNH 39727-32, 39734-46; La Mar, ±
760 m— FMNH 39725; La Mar, Ayna ± 1900 m— FMNH 39726; HUAN-
1970 Studies on the Frogs of the Genus LEPTODACTYLUS
45
UCO; Ganzo Azul, 150 m=-~FMNH 45144; JUNIN: Satipo, 500 m~^MCZ
24424-8, UMMZ 89476 (7); Tarma, Palmapata, 1200-1400 m-^FMNH
36827-30; Tarma, Valley of Vitoc River, 1200-1400 m—FMNH 36817-26;
LORETO: Rio Maranon, mo. of Rio Pastaza, 150 m- — AMNH 42221-3,
42225-7; Pampa Hermosa, 176 m— AMNH 42142-4, 42661; Roaboya, 100
m----AMNH 42823-4; Rio Tamaya, Sobral, 150 m-— AMNH 43238; Rio Alto
Tapiche, mo. R. Contaya, 200 m— AMNH 42988; Tibi Playa, 150 m— -
AMNH 42783; Rio Ucayali, Pacapa, 150 m---FMNH 56322; Upper Uto-
quina, 200 m — ^AMNH 42599; PASCO: Iscozazin Valley, Chontiila, 780 m —
LACM 40660-5; Rio Pachitea, Monte Alegra, 150 m — AMNH 43013; SAN
MARTIN: Tocache, 450-750 m-™AMNH 42627. SURINAM. MARO-
WIJNE: Kaiserberg airstrip, Zuid R., 200-300 m— FMNH 128925, 128927-
8, 128933-7, 128939-40. VENEZUELA. AMAZONAS. Rio Pescado—
AMNH 23182; ARAGUA: Colonia Tuvar, 1830 m — CM 7729; Cumboto,
150 m— UMMZ 113975 (2) ; Rancho Grande, nr. Maracay, 450 m — AMNH
70687; BOLIVAR: Arabopo, 1280 m-— UMMZ 85197 (5); Mt. Roraima,
Paulo, 1216 m^AMNH 39753, 39758-9; DELTA AMACURO: 140 km. NE
Barrancas, 5-10 m^LACM 31382-97; DISTRITO FEDERAL; Cotiza,
Camino de Calipan, 900 m-— USNM 117526, 128837; El Limon, 577 m~
CM 7676-7, 7697, 7699, USNM 121146; FALCON: Cerro Cosme, 100-200
m— MCZ 26144-6; Lagoon, Palma Sola, 50 m — -UMMZ 55551; MERIDA:
Merida, 1641 m—MCZ 2640, USNM 118176; Rio Albirregas nr. Merida,
1640 m-^AMNH 10517-9; MIRANDA: Petare, 885 m^CM 7769-71, 7791;
USNM 121147; MONAGAS: Caripito, dz 50 m-™AMNH 70669-86, USNM
117088-9; SUCRE: Cumanocoa, 300 m-^CM 9055-76; Latal, Hacienda
Mirasol, 975 m— --CM 9099, 9101; San Rafael, nr. Cumanocoa, 200-400 m--
CM 8972; El Yaque, 800 m— CM 9108-9; TaCHIRA: La Fria, Rt. Fork
Rio Oropito, 110 m— -UMMZ 55552-3; ZULIA: Chama, 10 m — -AMNH
10685-8. WEST INDIES: BEQUIA: north end~-USNM 103976-8; GRE-
NADA: Grand Etang-=-MCZ 2963-71, 2974-6; no specific locality— -AMNH
18961, 18970-2, 18992, USNM 67183-4, 67186-94; ST. VINCENT: Botani-
cal Garden-^USNM 144246; Brighton^USNM 79068-75; Mt. St. Andrews
“—USNM 79076-7; TOBAGO: Bacolet River“-AMNH 55873; Buccoo Bay
^AMNH 55863-5; nr. Speyside, 180 m-=-=MCZ 27788-9; TRINIDAD:
Nariva: Brickfield^FMNH 49656-60; Upper Tucker Valley-— USNM
119055-60; St. Andrew: Mount Harris—FMNH 49654-5; Sangre Grande—
MCZ 3295-8; San Rafael—FMNH 49664-5; St. George: St. Augustine-— -CM
5466.
Resumen
En el grupo Melanonotus se reconocen las siguentes seis especies: dantasi,
discodactylus, melanonotus, podicipinus, pustuiatus, y wagneri.
Para cada especie se ofrecen: sinonimia, diagnostico, sumario de carac-
46
Contributions in Science
No. 191
teristicas y distribucion. Series de Leptodactylus procedentes de amplias areas
geograficas y ecologicas fueron analizadas para estudiar largo normal, apa-
riencia ventral, desarrollo de las glandulas ventrolaterales, apariencia de las
extremidades posteriores, y desarrollo de los discos digitales. Se encontraron
correlaciones evidentes entre: tamano grande y habitates humedos en wagneri,
2) apariencia ventral mas oscura y una cantidad mayor de Iluvia anual en
melanonotus, 3) apariencia ventral palida y una mayor extension de las
glandulas ventrolaterales en wagneri, y 4) mayor extension de las glandulas
ventrolaterales y mayor cantidad anual de Iluvia en wagneri.
Se postula la hipotesis de que existe displazamiento de caracteres en la
estricta diferencia en tamano y apariencia tanto entre poblaciones de mela-
nonotus y wagneri como entre podocipinus y wagneri las cuales viven sim-
patricamente.
Leptodactylus melanonotus es la especie mas primitiva, mientras que
discodactylus y pustulatus son mas avanzadas.
Leptodactylus melanonotus, podicipinus, y wagneri estan asociadas con
regiones antiguas y son especies adaptadas a condiciones xericas. Leptodacty-
lus dantasi y discodactylus estan limitados al bosque tropical humedo del valle
al oeste del Amazonas. Leptodactylus pustulatus se encuentra en las regiones
orientales xericas de Brasil.
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Accepted for publication February 17, 1970
LOS
ANGELES
' COUNTY
MUSEUM
CONTRIBUTIONS
c:lIs in science
UMBER 192
June 26, 1970
SYSTEMATICS OF THE LIZARDS OF THE GEKKONID GENUS
PHYLLODACTYLUS OF MAINLAND SOUTH AMERICA
By James R. Dixon and Raymond B. Huey
Los Angeles County Museum of Natural History • Exposition Park
Los Angeles, California 90007
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SYSTEMATICS OF THE LIZARDS OF THE GEKKONID GENUS
PHYLLODACTYLUS OF MAINLAND SOUTH AMERICA
By James R. Dixon^ and Raymond B. Huey“
Abstract: A brief description of the habitat of species of
Phyliodactyliis of mainland South America is presented. Micro-
habitat preferences are given for each species where possible.
Little or no microhabitat information is available for P. heter-
urus, sentosus, and cUnatus. Four species, P. angiistidigitus, ger-
rhopygiis, microphyUus, and sentosiis are primarily terrestrial
and allopatric in distribution. Nine species, P. clinatus, dixoni,
inaequalis, interandinus, johnwrighti, kofordi, lepidopygus, piimi-
lus, reissi, and ventralis are scansorial and only johnwrighti,
ventralis, and dixoni are not found sympatric with one or more
of the other scansorial species. Where three or more species are
sympatric, there is little overlap in daytime retreats and foraging
areas.
One egg is laid per clutch by most species of Phyliodactyliis
of mainland South America except reissi, which lays two eggs.
Information on clutch size is not available for F. heteriirus, sen-
tosus, and clinatus. Communal egg laying appears to be common
among most species, including such sympatric species as F. inter-
andinus and reissi, kofordi and reissi.
Species of Phyllodactylus appear to have specific adapta-
tions to their microhabitat. Enlarged terminal digital lamellae,
dorsoventrally compressed head and body, and lateral nostrils
characterize species of Phyllodactylus from a boulder micro-
habitat; small terminal digital lamellae, round body and head,
dorsolateral valvular nostrils for species from a sand microhabi-
tat; small terminal digital lamellae with enlarged claw, slim body
and head, dorsolateral valvular nostrils for species from shell
beaches. Depending upon the degree of scansorial or terrestrial
behavior some species of Phyllodactylus are intermediate in the
above adaptations.
Distribution maps and photographs are given for each spe-
cies, except F. heterurus, and a gazetteer of important localities
is presented.
Of the holotypes and syntypes for species described from
the mainland of South America, ten (dixoni, inaequilis, lepido-
pygus, magister, microphyUus, nigrofasciatus, phacophorus, pu-
milus, reissi, and ventralis) are extant and eight have been exam-
ined. Seven new species (angustidigitus, clinatus, interandinus,
johnwrighti, kofordi, pumilus, sentosus) are described herein and
nine older names (abrupteseriatus, baessleri, guayaquilensis,
lobensis, magister, nigrofasciatus, phacophorus, underwoodi,
variegatus) are relegated to the synonomy of other species.
^Department of Wildlife Science, Texas A&M University, College Station, Texas.
^Department of Zoology, University of Texas, Austin, Texas.
1
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Contributions in Science
No. 192
Introduction
The leaf-toed geckos, genus Phyllodactylus, form two distinct geographic
units in South America, a northeastern group in Colombia, Venezuela and the
Caribbean Islands, and a western group in Ecuador (including the Galapagos
Islands), Peru, and Chile. A study of the northeastern representatives was
completed by Dixon ( 1962, 1964a). The present work treats the systematics
and biology of the mainland South American Phyllodactylus. Although the
Galapagos Islands species are evolutionarily close relatives to the mainland
species of western South America, they are a special problem and will be
reported on at a later date.
The earliest description of a species for the mainland of South America
was by Wiegmann (1835) for P. gerrhopygus and the most recent by Rivero-
Bianco and Lancini (1968) for P. dixoni. A total of 19 species have been
described, ranging from the coast and mountain areas of Ecuador to Chile,
including the upper Amazon Basin of northwestern Peru in western South
America; from the lowlands and foothills of eastern Colombia to central
Venezuela in eastern South America.
A common Mexican and Central American species, P. tuberculosus,
described by Wiegmann ( 1835), has also been reported in the literature from
Ecuador, Peru, the Galapagos Islands, and the Greater Antilles. The distribu-
tion of this species is Mexican and Central American (Dixon, 1960), and
specimens on the mainland and islands of South America to which this name
has been misapplied represent populations of P. reissi, P. darwini, and P.
wirshingi.
Individual species were studied and described by Wiegmann (1835),
Tschudi (1846), W. Peters (1862), Cope (1876, 1877), Werner (1901, 1907,
1910, 1912), and Noble (1924). Short notes on distribution, habitat and
taxonomy of mainland South American species were given by Parker (1935),
Taylor (1942), Codoceo ( 1950), Titschack (1951), Yanez (1951), Dixon
(1962, 1964a), J. Peters (1967), Donoso-Barros ( 1966), and Rivero-Bianco
and Lancini ( 1968).
As a group, the mainland South American species were studied by Bou-
lenger (1885) and Werner (1912), both of whom published a key to the arid
and tropical American species known at the time. Burt and Burt 0933) pub-
lished a preliminary checklist of lizards of South America. Wermuth (1965)
published a world checklist of the genera, species, and subspecies of the fam-
ily Gekkonidae, and Vanzolini ( 1968a) published a paper on the geographic
distribution of South American Gekkonidae. Vanzolini (1968b) also pub-
lished a key and list of Gekkonidae known to occur in Brazil, and included
a list of the species of Phyllodactylus known to occur in South America.
Mechler (1968) published a key, descriptions, and maps for all gekkonids in
Colombia.
Some of the latter studies contain errors in species determination, through
the uncritical acceptance of publications of earlier workers who did not have
1970
Lizards of the Genus Phyllodactylus
3
adequate knowledge of the distribution and ecology of the species. With
some of these problems in mind, a study of the genus was initiated by one of
us (Dixon) in 1957. Recent collections of Phyllodactylus by Huey in 1967
and 1968, and by Dixon and Wright in 1968, from the coast and upper Amazon
Basin of Peru and an examination of extant syntypes and holotypes of South
American species has provided the necessary material to complete this study.
A detailed study of the sympatric relations of P. kofordi, P. microphyllus,
P. reissi, and F. clinatus will be presented by one of us (Huey) at a later date.
Type Material
Of the 19 available names for mainland South American species of
Phyllodactylus, only nine have extant type material that has been examined
by us: lepidopygus, phacophorus, reissi, inaequalis, microphyllus, nigrofascia-
tus, magister, ventralis, and dixoni.
The remaining 10 typical series are either definitely destroyed or lost.
The museums in Hamburg and Dresden were seriously damaged during the
fire bombings of the second world war. Among type material destroyed in
the Dresden Museum were the nine syntypes of F. variegatus and the holotype
of F. baessleri. The Hamburg Museum lost two syntypes each of F. abrup-
seriatus and F, guayaqidlensis, and type material (if any existed) of F. lobensis
(nomen nudum). The syntypes of F. gerrhopygus, reported to be housed in
the Berlin Museum, are lost (Wermuth, pers. comm.). The holotype of F.
heterurus, housed in the School of Medicine, University of Chile, was destroyed
by a fire in the early 1950’s (Capurro, pers. comm.).
Fortunately, Donoso-Barros (1947) was able to examine the type of F.
heterurus in detail prior to its destruction by fire. However, some of the early
type descriptions by Werner (1912) were brief and failed to give a detailed
description that could be used to identify his species with certainty. There-
fore, our analyses of his original descriptions and our subsequent synonomies
based on these descriptions may be in error.
A problem related to the loss of types is knowledge of the exact type
locality for some species. Werner (1912) gives the type locality for F. abrup-
terseriatiis as “probably Brazil.” Members of this genus have never been found
in Brazil and we assume, as have earlier workers (e.g. Burt and Burt, 1931),
that the type series probably came from Ecuador or northern Peru. The type
description clearly describes individuals of Phyllodactylus reissi Peters in
these areas. Tschudi’s (1846) original descriptions of F. lepidopygus and
F. phacophorus gave only the type locality for the former species (Chorillos,
Peru). The only statement made on the type locality of the latter species was
“a wooded region.” Werner’s (1901) description of F. baessleri gave the town
of Chanchamayo, Peru, as the type locality. Several towns of that name were
located in west central and southern Peru. Most of these towns are sufficiently
low in elevation to sustain populations of Phyllodactylus, but only two species
are known to occur in the above areas, F. gerrhopygus and F. lepidopygus.
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No. 192
The only known populations fitting Werner’s description are found 1600 km
NW of the nearest town named Chanchamayo. Therefore, the type locality
of P. baessleri is presumed to be in error.
These and other problems have been responsible for the misapplication
of species names in a number of publications on Phyllodactylus over the past
100 years. Consequently, we present the following table of the chronological
synonymy of sixty-one specific names proposed for North, Central, and South
American species.
Table 1
Chronological synonymy of sixty-one specific names proposed for the
genus Phyllodactylus from North, Central and South America
Date, original name and author
present name
1830 pulcher Gray
pulcher
1835 tuberculosiis Wiegmann
tuberculosus
1835 gerrhopygus (Wiegmann)
gerrhopygus
1836 gymnopygiis Dumeril & Bibron
gerrhopygus
1846 lepidopygus (Tschudi)
lepidopygus
1846 phacophorus (Tschudi)
lepidopygus
1862 reissi Peters
reissi
1862 spatiilatus Cope
pulcher
1862 julieni Cope
julieni
1863 iinctus Cope
line t us
1863 xanti Cope
xanti
1869 galapagoensis Peters
galapagoensis
1875 ventralis O’Shaughnessy
ventralis
1876 inaeqiialis Cope
inaequalis
1876 microphyllus Cope
microphyllus
1877 nigrofasciatus Cope
lepidopygus
1887 martini Van Lidth de Jeude
martini
1889 leei Cope
leei
1892 bauri Garman
bauri
1900 variegatus Werner
lepidopygus
1900 baessleri Werner
reissi
1903 gilberti Heller
gilberti
1907 he ter unis Werner
heterurus
1910 mentalis Werner
xanti
1910 guayaquilensis Werner
reissi
1910 lobensis Werner
microphyllus
1912 abrupteseriatus Werner
reissi
1912 barringtonensis Van Denburgh
barringtonensis
1924 magister Noble
reissi
1935 mulleri Parker
ventralis
1935 homolepidurus Smith
homolepidurus
1935 lanei Smith
lanei
1936 delcampi Mosauer
delcampi
1940 magnatuberciilatiis Taylor
lanei
1970
Lizards of the Genus Phyllodactylus
5
1940 muralis Taylor
muralis
1940 rutteni Hummelinck
rutteni
1942 darwini Taylor
darwini
1942 bordai Taylor
bordai
1942 magnus Taylor
tiibercidosiis
1952 eduardofischeri Mertens
tubercidosus
1955 wirshingi Kerster & Smith
wirshingi
1960 duellmam Dixon
duellmam
1960 paucitiiberculatiis Dixon
paucitiiberculatiis
1960 insidaris Dixon
insidaris
1962 iinderwoodi Dixon
ventral is
1964 davisi Dixon
davisi
1966 angelensis Dixon
angelensis
1966 partidus Dixon
partidus
1966 tinklei Dixon
tinklei
1966 bugastrolepsis Dixon
bugastrolepis
1966 santacruzensis Dixon
santacruzensis
1 966 apricus Dixon
apricus
1968 palmeiis Dixon
pal me us
1968 dixoni Rivero-Bianco & Lancini
dixoni
1970 kofordi new species
kofordi
1970 interandinus new species
interandinus
1970 johnwrighti new species
johnwrighti
1970 angustidigitus new species
angustidigitus
1970 clinatus new species
clinatus
1970 sentosus new species
sentosus
1970 pumilus new species
pumilus
Materials and Methods
Approximately 1,800 specimens of Phyllodactylus were examined from
mainland South America. The material is housed in the American Museum
of Natural History (AMNH); Academy of Natural Sciences, Philadelphia
(ANSP); Berlin Museum (BM); British Museum (Natural History)
(BMNH); California Academy of Science (CAS); Escuela Politecnica
Nacional, Quayaquil (EPN); Field Museum of Natural History (FMNH);
Hamburg Museum (HM); Los Angeles County Museum of Natural History
(LACM); Museo de Ciencias Naturales, Caracas (MCNC); Museo de Bio-
logia de la Universidad Central de Venezuela (MBUCV); Museum of Com-
parative Zoology (MCZ); Museo “Javier Prado,” Peru (MJP); Museum of
Vertebrate Zoology (MVZ); Neuchatel Museum, Switzerland (NM); San
Diego Society of Natural History (SDSNH); Senckenberg Museum, Frank-
furt am Main (SMFM); Texas Cooperative Wildlife Collection (TCWC);
University of Illinois Museum of Natural History (UIMNH); University of
Michigan Museum of Zoology (UMMZ); United States National Museum
(USNM); private collections of Charles M. Fugler (CMF), and Raymond
B. Huey (RBH).
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No. 192
Descriptions of habitat, behavior, ecology, and color notes of living
specimens were obtained in the field by Huey during July, August, and Sep-
tember, 1967 and 1968 and by Dixon and Wright during November and
December, 1968. Climatological data were obtained from Robinson ( 1964)
and details of the vegetation were obtained from Macbride ( 1936). Measure-
ments and analysis of squamation were made by methods described by Dixon
(1964b). Sex was determined by the presence of cloacal bones and enlarged
postanal tubercles in males, or by dissection.
When we present a description of a species, it will describe all individuals
except those covered in the sections on variation.
General Description of the Habitat of
Western South America
The Andes of western South America divide the coastal countries of
Ecuador and Peru into three relatively distinct geographic and biological
units: the narrow Pacific coast, the Andes and associated valleys, and the
tropical forests of the east. Species of Phyllodactylus and other lizard genera
such as Tropidurus, Dicrodon, and Callopistes are restricted primarily to the
arid coast of western South America and to the northwestern interandean
basins of Peru.
The coastal zone, buttressed by the moist Chocoan forests in northern
Ecuador (Peters J., 1960), passes south from the thorn scrub and mangrove
forests of Ecuador to the barren deserts of Peru and northern Chile. With the
exception of a few low passes in northern Peru the Andes to the east form a
high barrier generally more than 4,250 meters in areas relevant to discussion.
The deciduous thorn scrub forests extend from the Pacific coast and
Andean foothills of Ecuador to the extreme northwestern corner of Peru in
the Department of Tumbes, east to the base and foothills of the Andes in the
Department of Piura, and then south along the foothills of the Andes into
the Department of Lambayeque. Some of the dominant plants of this region
include species of Loxopterygium, Caesalpinia, Bursura, Bombax, Cap pans,
Mimosa, and Cereus (Weberbauer, 1936) .
The junction of the Ecuadorian thorn scrub and the Peruvian deserts
in Tumbes is very narrow and has been attributed to the climatic changes
associated with the boundary between the cold waters of the Humboldt Cur-
rent and the warmer Equatorial Counter Current (Dawson, 1963). The coast
of Peru below Tumbes to northern Chile is a barren, arid desert. This desert
is not induced by excessively high temperature regimes; rather, it is apparently
the result of the aridity associated with the cold Humboldt Current and a
rainshadow produced by the Andes (Dawson, 1963; Kendrew, 1961; Murphy,
1948) . Rainfall is very low and sporadic throughout much of the area. Most
of the water available to plants comes from rivers penetrating the coast from
the western slopes of the Andes, high water tables in the Quebradas, and fault
zones of northern Peru and fog.
1970
Lizards of the Genus Phyllodactylus
7
Weberbauer (1936) recognizes two major plant zones in the Peru and
Chile deserts. The first runs from Tumbes to a latitude of about 8° south. This
area is essentially rainless, except for stormy El Nino years, and extremely
barren. Encompassed are such areas as the Sechura Desert in the Department
of Piura. Plants tend to be widely scattered and grow as shrubs rather than
trees. The substratum is sandy throughout much of the region with a few
rocky cerros and the rocky, hard soils of the Quebrada zones. Characteristic
features are shifting barchan dunes intermingled with large hummock-like
dunes stabilized by Cap parts scabrida (Koford, 1968) . Major plants inhabitat-
ing this region include species of Cryptocarpus, Prosopis, Capparis, and A cacia
(Weberbauer, 1936) .
The second zone follows the narrow coast to northern Chile and contains
the famous Lomas and Tillandsia plant formations. The Lomas are chiefly
seasonal carpets of herbs and occasional low trees in areas where the coastal
foothills of the Andes rise to heights sufficient to intercept the winter and
spring clouds. The increased elevation and lowered temperatures produce con-
siderable condensation in the form of mists making the existence of vegetation
possible in an otherwise arid climate. The average annual precipitation
recorded in the open is 201 mm for 29 years at Lomas de Lachay in the
Department of Ancash (elevation 450 m) compared with only 46 mm for
18 years at nearby Lima (Holdridge, 1964) . Major Lomas plants include
species of Aniantum, Bowlesia, Astrephia, Sicyos, and abundant lichens,
mosses and ferns (Weberbauer, 1936).
Unlike the Lomas, the Tillandsia association is not dependent on sea-
sonal changes and appears to survive upon atmospheric moisture. Tillandsia
is most often found on the leeward side and at a lower elevation of hills
covered with Lomas vegetation, in drier valleys inland from the coast, and
along the coastal plain where no other plants survive. There are large expanses
of the deserts where even Tillandsia is not found and one is confronted only
with barren sand reaching from the Pacific tides to several thousand feet up
the Andean foothills.
The deserts of northern Chile extend south to a latitude of about 27°.
Extremely barren and arid, much of these deserts are colonized only by Til-
landsia, South of 21° , the desert is bordered by brush and savanna (Mann
1960). To the east the height of the Andes has apparently prevented invasion
by species of Phyllodactylus into the Larrea deserts of western Argentina.
The northern interandean basins, lying between latitudes 4° to 6° south
and longitudes 78° to 80° west (Weberbauer, 1936), are loosely connected
with the Peruvian coast by the relatively low passes in the Department of
Piura (e.g. Porculla Pass at 2,160 m). These low passes have served as an
invasion route for species of Phyllodactylus and other species of coastal lizards
into a portion of the Amazon drainage.
Surrounded by the higher Andean passes to the north and south and by
the Amazonian forests to the east, the interandean basins are limited in extent
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Contributions in Science
No. 192
and similar in appearance to the coastal foothills of northwestern Peru. The
general aridity of these basins, less pronounced than the coast, apparently
results from a rain-shadow produced by the eastern chain of the Andes cutting
the basins off from the Amazonian moisture.
The interandean basins vary considerably in species composition and
physiognomy but are somewhat uniform in vegetative cover with cacti and
deciduous plants covering the basin floors except along water courses. The
higher slopes are generally matted with dense growths of woody plants (Weber-
bauer, 1936).
Further details of the habitat of Ecuador, Peru, and Chile can be obtained
from Weberbauer (1936), MacBride (1936), Tosi (1960), Koepcke, M.
(1954), Koepcke, H. (1961), Dawson (1963), Murphy (1948), Mann
(1960), Robinson (1964), Kendrew (1961), Donoso-Barros (1966), and
Verdoorn (1945).
Details of the arid and semiarid habitats of Colombia and Venezuela
may be obtained from Ruthven (1922) and Shelford (1926).
Key
The following key is based upon approximately 2,000 specimens of 15
species of Phyllodactylus from mainland South America.
The size, shape, and number of scales in the median row beneath the
tail is variable within some species but it is generally consistent if a careful
examination of the entire, original tail is made. Regenerated tails do not have
tubercular scales even if they were present on the original tail. In some
instances, the width of the scales of the median row beneath the regenerated
tail may be about the same size as those of the original tail.
Key to the Species of PHYLLODACTYLUS of Mainland South America
la. Preanal scale single, greatly enlarged, 10-20 times larger than belly
scales ................................................... 2
lb. Preanal scales small, equal in size to other belly scales ............ 4
2a. Tail smooth, without enlarged rows of lateral tubercles ........... 3
2b. Tail keeled, with four rows of lateral tubercles (northern Chile) heterurus
3a. Digits short, fourth toe lamellae 12-15 (13.1), terminal lamellae large
(southern Peru to northern Chile) .................. . gerrhopygus
3b. Digits long, fourth toe lamellae 15-20 (16.8), terminal lamellae very
small (coastal south central Peru) . . angustidigitus
4a. Dorsal tubercles small, rounded, sometimes slightly keeled, not arranged
in regular rows; ventral surface of tail with all scales of equal size. . . 5
4b. Dorsal tubercles large, trihedral, arranged in regular rows; ventral sur-
face of tail with or without enlarged scales in median row ......... 6
5a. Terminal lamellae of digits much enlarged, most of claw hidden when
viewed from above; nostril not swollen (extreme northwestern Peru
and probably southwestern Ecuador) . inaequalis
1970
Lizards of the Genus Phyllodactylus
9
5b. Terminal lamellae of digits very small, most of claw exposed when
viewed from above; nostril greatly swollen (central to extreme north-
western Peru) microphyllus
6a. Dorsal surface of tibia without enlarged tubercles 7
6b. Dorsal surface of tibia with enlarged tubercles 9
7a. Dorsal surface of tibia with very small granular scales; ventral surface
of tail with all scales of median row enlarged, rectangular (northwestern
Peru) clinatus
7b. Dorsal surface of tibia with round, elevated scales; ventral surface of
tail with or without some enlarged, rounded scales in median row. . . 8
8a. Tubercles in paravertebral row from head to base of tail 44-53; no
scales on ventral surface of tail enlarged (coastal foothills of central
Peru) lepidopygus
8b. Tubercles in paravertebral row from head to base of tail 65-81; some
scales in median row of ventral surface of tail enlarged, rounded (Mara-
non Basin, northwestern Peru) interandinus
9a. Tubercles present on dorsal surface of forearm 10
9b. Tubercles absent on dorsal surface of forearm 13
10a. Terminal lamellae of digits much enlarged; claw mostly hidden when
viewed from above; ventral surface of tail with all scales of median row
enlarged, rectangular 11
10b. Terminal lamellae of digits small, narrow; claw mostly exposed when
viewed from above; ventral surface of tail with a few scales of median
row enlarged, rounded (Lima, Peru) sentosus
11a. Large species, average snout-vent length of 64 mm; 19-26 paravertebral
tubercles between axilla and groin 12
1 lb. Small species, average snout-vent length of 38 mm; 16-21 paravertebral
tubercles between axilla and groin; (northwestern Peru and probably
southwestern Ecuador) kofordi
12a. Distal one-third of tail with at least two or more rows of tubercles; scales
across venter 18-24; (northern Venezuela and Colombia) . . . . ventralis
12b. Distal one-third of tail without tubercles; scales across venter 26-28;
(central Venezuela) dixoni
13a. Scales of supraocular region largest of interorbital scales; enlarged tuber-
cles usually present on dorsal surface of thigh; maximum snout-vent
length 51 mm 14
13b. Scales of supraocular region smallest of interorbital scales; enlarged
tubercles usually absent on dorsal surface of thigh; maximum snout-vent
length 75 mm (northwestern Peru and west central Ecuador) . . . . reissi
14a. Scales on proximal one-fourth of tail homogeneous; internasals usually
separated along median edges (Huancabamba Basin, northwestern
Peru) johnwrighti
14b. Scales on proximal one-fourth of tail heterogeneous; internasals usually
in contact along median edges (west central Ecuador) pumilus
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Contributions in Science
No. 192
Species Accounts
Phyllodactylus heterurus Werner
Phyllodactylus heterurus Werner 1907. Ann. Univ. Chile, 121 : 149.
Holotype. Adult male, (lost in a fire), collected by C. Reiche 1902. Type-
locality. Oasis de Pica (Desierto de Tarapaca), Department of Tarapaca,
Chile.
Definition and Diagnosis. This species is known only from the holotype
that was destroyed by a fire. Phyllodactylus heterurus is a small gecko with a
maximum snout-vent length of 38 mm; dorsal surfaces of head and body
smooth, without tubercles; tail with four rows of tubercles, two on each lateral
surface; preanal plate present; terminal lamellae of digits moderately large.
This species is distinguished from all other species of Phyllodactylus
except gerrhopygus and angustidigitus, by the presence of an enlarged preanal
plate; from gerrhopygus and angustidigitus by the presence of four rows of
tail tubercles, with anteriorly projecting tips.
Description of holotype. Since the holotype is no longer extant, we sub-
mit a translated version of Werner’s (1907) original description: Head ovoid,
very much longer than wide; snout longer than the distance from the posterior
corner of eye to ear and two times the length of eye; frontal region not con-
cave; ear opening small, horizontal and elliptical; body decidedly thin; inferior
aspect of digits similar on anterior and posterior limbs, with wide plates whose
distal ends are visibly wider than their proximal portion; dorsal aspect of body
with small scales that are ovoid, smooth, somewhat convex; rostral scales
pentagonal, somewhat concave posteriorly, with a groove beginning on pos-
terior border; nasal opening surrounded by rostral, supralabial and three
nasals; nine supralabials and seven infralabials; nasal (internasals) scales
separated on median line by an azygous scale; mental two times longer than
wide with posterior border rounded, lateral sides parallel; chin without enlarged
scales (postmentals absent) ; scales in gular region very small; ventral scales
small, smooth, circular and overlapping; tail cylindrical, with transverse series
of robust, conical tubercles whose points are directed anteriorly.
Color and pattern on dorsum lead gray with black irregular spots and
lines; lines unite on dorsal side to form transverse bands with obscure out-
lines; tubercles on tail brick red.
Total length 71 mm, head 12 mm in length by 7.5 mm in width; tail
30 mm in length.
Comments. Donoso-Barros ( 1947) redefined the holotype prior to its
destruction and further distinguished the species from gerrhopygus by its
large, round eye, ovoid head with sharply pointed snout, and gave preserva-
tion colors that differed from those given by Werner ( 1907) .
As far as we can determine, no additional specimens have been taken
since 1902, therefore data are not available on its habits, habitat, or natural
history.
Distribution. Known only from the type locality (fig. 5).
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Lizards of the Genus Phyllodactylus
11
Phyllodactylus gerrhopygus (Wiegmann)
Diplodactylus gerrhopygus Wiegmann 1835. Nova Acta Leop. -Carol.,
14:242.
Phyllodactylus gymnopygus Dumeril and Bibron 1836. Erp. Gen., 3:394.
Phyllodactylus gerrhopygus, Boulenger 1885. Cat. Liz. Brit. Mus., 1:95.
Phyllodactylus inaequalis, (part) Burt & Burt, 1931. Bull. Amer. Mus.
Nat. Hist., 61(7) :250.
Holotype. Lost, formerly in Berlin Museum, collector and date of col-
lection unknown. Type-locality. Chile.
Definition. An average size gecko with a maximum known snout-vent
length of 56 mm; large preanal plate present; dorsal tubercles absent; tibia
and femur tubercles absent; ear denticulate on anterior and posterior margins;
median row of scales beneath tail equal in size to other ventral scales; tail
about 50 percent of total length; mental much longer than wide and post-
mentals absent; terminal lamellae of digits large, longer than wide; claws of
digits large, usually extending slightly beyond tip of terminal lamellae; dorsal
markings present or absent, if present, consisting of broad bands or blotches;
tail usually banded (fig. 8a).
Diagnosis. This species may be distinguished from all other species,
except angustidigitus and heterurus, by the presence of an enlarged preanal
plate; from heterurus by the absence of enlarged lateral tubercles on the tail;
from angustidigitus by the larger toe pads and shorter digits.
Description. Rostral about as high as wide, its dorsal edge “M” shaped,
with a short median vertical groove; two internasals, somewhat triangular in
shape, their median edges not in contact but separated by three azygous scales
(snout granules) ; internasals bordered posteriorly and medially by small
granules and postnasal of each side; nostril surrounded by rostral, labial, inter-
nasal, and two postnasals; nostril area somewhat swollen by elevation of
postnasals and internasal; first supralabial in broad contact with ventral edge
of nostril; moderate depression between internasals, slight depression in frontal
region; scales in posterior loreal region about twice the size of midorbital
scales; eye large, its diameter contained in snout length about 1.5 times; eyelid
with two rows of granules and one larger outer row of scales, the last three
to six are pointed; diameter of ear contained in eye diameter about 2 times;
ear opening denticulate on anterior and posterior margins, rear of head granu-
lar; usually 8 supralabials and 5 or 6 infralabials to point below center of eye;
mental straight-edged, rounded posteriorly, about 2.5 times longer than wide;
postmentals absent, mental bordered by small chin scales.
Dorsum without enlarged tubercular rows, scales subequal in size, some-
what flattened, about half size of midventral scales; latero-ventral scales
smaller than dorsal and ventral scales; very large preanal plate (scaleless),
surrounded by about 25 ventral scales; preanal plate about 20 to 30 times
larger than adjoining ventral scales; enlarged postanal scales on each side,
somewhat flattened.
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Figure 1. Lateral, ventral and dorsal view of the distal end of the fourth toe of (A)
P. gerrhopygus and (B) P. angustidigitus.
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Lizards of the Genus Phyllodactylus
13
Dorsal surface of upper arm with flattened scales, forearm with larger,
slightly elevated scales; dorsal surface of thigh and lower leg with flattened,
but slightly elevated scales; claw moderately exposed when viewed from
below; terminal lamellae twice as long as wide, somewhat reduced in size
with claw extending beyond tip of terminal lamellae; tail with all scales of
ventral surface of equal size, scales of dorsal surface smaller than those of
ventral surface.
Color in alcohol: ground color tan, with light orange tint; dorsum with
four broad light brown bands with undulating edges bands composed of suf-
fusion of brown and tan scales intermixed (freckled) , with tan interspaces
slightly wider than dark bands; upper and lower arms and legs with faint
brownish bands; tail with nine bands similar in color to dorsal bands, with
yellowish tan interspaces; posterior part of head and nape with reticulating
brown lines on ground color; top of head spotted with brown; brownish stripe
from nostril to eye; brownish reticulating lines between eye and ear; labials
faintly flecked with dark brown; ventral surfaces of head, body, and tail
immaculate white.
Variation. Snout-vent length of young adult and adult males varies from
32 to 56 mm (43.9), females from 32 to 55 mm (42.8), juveniles from 25 to
31 mm (28.9), and males and females together average 42.7 mm; postmentals
absent in 96.4 percent of sample; three of 98 specimens have two postmentals,
one specimen had one postmental on one side, none on the other side; mental
much longer than wide, extending posteriorly beyond the tips of adjoining
labials; number of transverse chin scales immediately following and touching
mental vary from 3 to 8 (4.6) ; scales across midorbital region vary from 13 to
20 (15.9), across the snout at level of third labial from 12 to 18 (14.7); num-
ber of scales bordering internasals vary from 6 to 10 (7.7), internasals always
separated along median line by 1 to 3 granules; transverse row of scales across
venter vary from 20 to 30 (24.7), longitudinally 64 to 76 (69.1); preanal
shield large, without scales, generally 20 to 30 times larger than adjoining
scales; scales between eye and nostril vary from 8 to 12 (10.4); number of
lamellae beneath fourth toe vary from 12 to 15 (13.1); all scales of dorsum,
limbs, and tail subequal in size, relatively smooth, and slightly elevated to
some extent; supralabials vary from 7 to 9 to a point below center of eye,
8 being most common; dorsal bands (if present) number 4 to 6, tail bands
6 to 9.
Color pattern consists of body bands or blotches that tend to appear
freckled with tan and brown scales intermixed, but occasionally some speci-
mens are uniform tan or light brown; tail usually banded; belly usually immacu-
late white; parietal region and lateral area of head behind ear usually with
reticulated brown lines on ground color; limbs usually faintly banded with
brown.
Habitat and natural history. Of 98 individuals examined, only five were
collected by one of us. One of these was collected during the day from beneath
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No. 192
a piece of tin in an open sand dune area where some vegetative cover was
present. Four individuals were found active after dark along the base of sand
dunes approximately 150 m from a sand and rock rubble beach, and approxi-
mately 50 m from a ravine where some plant cover was present. The dunes
were approximately 50 to 75 m in height and without vegetation. Silverfish,
beetles, and spiders appear to be the only food items available in the area where
the five specimens were taken.
The absence of individuals along the beach might be explained by the
presence of hundreds of ghost crabs occupying the intertidal zone.
One specimen in the University of California Museum of Vertebrate
Zoology with field data was taken from beneath a stone in a sandy wash.
Carman (1875) mentions that his specimens were found beneath stones in
a sandy plain near Arequipa, Peru.
Through the courtesy of Stanley W. Taft, a Peace Corps Volunteer from
New Mexico State University, we were able to obtain some habitat data on
P. gerrhopygus from the San Juan de Marcona area of Peru. The San Juan
Marcona area is approximately 157 airline km SSE of lea, at an elevation
of 0 to 150 m. The general terrain is rolling sand hills with sandy beaches and
large outcroppings of a red limestone-like formation. Taft (personal com-
munication) indicated that large hemitite deposits were present in the area
which probably give the red color to the sandy areas. The only vegetation of
the immediate area consisted of a few scattered Capparis sp. shrubs.
A series of P. gerrhopygus collected by Taft were taken from rock walled
terraces, outside walls of houses, and from sidewalks. Taft indicated that the
geckos were occupying small holes made by the absence of plaster or mortar
between the rocks and cement blocks of terrace walls and houses. All of his
specimens were collected at night, and two were taken from a terrace wall
near a sodium vapor light. Taft stated that several were seen along the sandy
beaches at night but were not collected.
From an examination of living and preserved females, there appears to
be only one egg laid per clutch. Nests of this species have not been found
under natural conditions.
Distribution. This gecko apparently occupies a variety of habitats along
the desert coast, foothills, and mountains from central Peru to northern Chile
between the elevations of sea level and 2,750 m (fig. 2). We have examined
98 specimens from the following localities: CHILE. Tarapaca: Arica UMMZ
1 1576; 3 mi S Arica AMNH 65343-59. 50 km from Iquique MCZ 56247.
Pica FMNH 6327-29, 9984. Pintados FMNH 6326. Putre FMNH 6330,
6331(3), 6332-41. PERU. Arequipa: Arequipa AMNH 22238, 36525-26,
FMNH 3577(4), 43769. 12 mi SE Camana CAS 84769. Chucarapi FMNH
34236-37. 16 km S. La Joya MVZ 84636. Mollendo CAS 84760-68, FMNH
34248(4). Tambo FMNH 4005-06. Vitor FMNH 34179. Yura FMNH
34047(9), 34205(5), 34182-83, 34190. Ica: 1 km NNW Jaguay LACM
48465-66, TCWC 28064-65: Marcona TCWC 27874-78. 6.3 km ESE Pozo
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Lizards of the Genus Phyllodactylus
15
Santo TCWC 28063. San Juan Port (Salamanquejo) SDSNH 31091-92.
Lima: Isla San Lorenzo USNM 17542 (probably in error).
Figure 2. Distribution of P. gerrhopygus, P. microphyllus and P. angiistidigitus in
western South America.
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No. 192
Phyllodactyius angustidigitus sp. nov.
Phyllodactylus gerrhopygus, (part) Burt and Burt 1930; Proc. U. S.
Nat. Mus., 78(2849) :4.
Holotype. Adult male, TCWC 27915, collected by James R. Dixon and
John W. Wright, 1 1 November 1968. Type locality. One km S new port of
Pisco, northeast side of Paracas Peninsula, 19 km by road south and west of
Paracas, Department of Ica, Peru, sea level.
Definition. A moderate-sized gecko, maximum known snout-vent length
of 57 mm; large preanal plate present; scales of head, body, tail, and limbs
uniform in size; tubercles absent; no enlarged scales beneath tail; tail 48 to
52 percent of total length; mental much longer than wide, without enlarged
postmentals; terminal lamellae very small, claw greatly extended beyond tip
of terminal lamellae; fourth toe lamellae numerous ( 15 to 20) (fig. 8a, 8b).
Diagnosis. Phyllodactylus angustidigitus differs from all other species of
Phyllodactylus except gerrhopygus and heterurus, by having an enlarged
preanal plate; from heterurus by absence of tail tubercles and small terminal
lamellae; from gerrhopygus by small terminal lamellae and high number of
fourth toe lamellae, 15 to 20, rather than 12 to 15.
Description of holotype. Rostral about twice as wide as high, its dorsal
edge with a broad “M” shape, with dorsal vertical groove one-half depth of
rostral; two internasals, somewhat rounded, their median edges separated by
two granules, bordered posteriorly by two granules and postnasal of each
side; nostril surrounded by rostral, labial, internasal, and two postnasals; first
labial in narrow contact with ventral edge of nostril; shallow depression
between internasals, no depression in frontal region; 10 scales on line between
nostril and eye; scales in posterior loreal region about twice size of midorbital
scales; 15 scales across snout at level of third labial, 14 scales at anterior edge
of orbits, 18 midorbital scales; eye large, its diameter contained in snout length
1.7 times; eyelid (brille) with one row of granules and one larger outer row
of scales, last three to four are pointed; diameter of ear contained in diameter
of eye 2.3 times; ear opening denticulate on posterior margin; all dorsal scales
small, smooth, slightly convex; seven supralabials and six infralabials to point
below center of eye; mental with parallel sides, about 2.5 times longer than
wide, bordered posteriorly by six chin scales; enlarged postmentals absent;
dorsum with 75 granules and scales around body; venter with 29 scales across
belly, from throat to vent 79; postanal tubercles number four on one side,
three on opposite side; dorsal surfaces of limbs similar to dorsum; lamellae
formula for hand 7-11-12-13-11, foot 7-12-16-18-16; claw completely visible
when viewed from below; terminal lamellae very small, not much wider than
width of fifth lamella counted from base of digit.
Measurements in mm; snout-vent length 53.5, axilla-groin length 25.8,
length of leg 24.0, length of arm 18.0, length of tail 55.0, length of head 13.3,
depth of head 6.2, width of head 10.5, length of snout 5.0, diameter of eye
2.9, diameter of ear 1.2, distance from eye to ear 3.8.
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Lizards of the Genus Phyllodactylus
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Color in life: ground color yellowish tan; dorsum with fine network of
thin, dark brown, reticulating lines, broken at midbody by median stripe of
ground color, bordered on each side by dark brown line; paravertebral lines
begin at shoulder and terminate at level of groin; arm with four light brown
bands with interspaces of ground color; lower part of leg with three indistinct
light brown bands; upper part of thigh with faint indication of bands with
irregular edges; tail with 1 1 broad, brownish black bands that contain light
brown diffuse spots within their borders, interspaces ground color, about
one-third width of dark bands; head with fine network of light brown, reticulat-
ing lines; diffuse brownish black stripe from nostril to eye; iris of eye blue
gray with black and silver flecks; ventral surfaces dirty white, with grayish
wash across chest and throat.
Variation. Snout-vent lengths of young adult and adult males vary from
41 to 57 mm (50.3), females from 37 to 54 mm (47.8), juveniles from 30
to 36 mm (32.8), and males plus females average 48.7 mm; postmentals absent
in 99.3 per cent of sample, two of 254 individuals had three enlarged post-
mentals; number of chin scales immediately following and touching mental
vary from 3 to 8 (5.8); scales across snout at level of third labial vary from
13 to 17 (15.1), across midorbital region 16 to 20 (18.2); internasals sepa-
rated by small granules in 99.5 per cent of sample; scales bordering inter-
nasals vary from 7 to 10 (8.7) ; scales across venter vary from 23 to 28 (25.6) ,
from throat to vent 64 to 70 (67.5); enlarged preanal shield without scale
covering, 20 to 25 times larger than surrounded scales; scales between eye
and nostril vary from 10 to 12 (11.0); number of fourth toe lamellae vary
from 15 to 20 ( 16.8) .
Ventral color generally uniform dirty white with some grayish wash on
chest and throat; dorsal coloration highly variable, from almost uniform wash
of yellowish tan to broad, dark brown bands; majority of individuals with
some form of dark dorsal reticulations or bands, usually heavily suffused with
ground color; tail generally banded, but width of dark bands and light inter-
spaces highly variable; regenerated tails either near black with light areas or
ground color with dark flecks; ground color variable, from light tan to very
dark grayish brown.
Comment. Since populations of angustidigitus appear to be closely
related to gerrhopygus, the following statement of our ideas on the evolu-
tionary trends in this population emphasizes its rate of speciation. The Paracas
Peninsula was apparently an island, probably as late as the Pleistocene, and
within recent times has become connected to the mainland. The attributes of
angustidigitus populations on Islas Sangallan and Viejas are identical with
those of the Paracas Peninsula, lending some support to this idea. The original
invasion of the islands by prototypes of a angustidigitus-Yiko. population may
have occurred by rafting, or by land invasion while the land masses were con-
nected early in the formation of the continental shelf.
The islands and peninsula populations are presently restricted in distribu-
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No. 192
tion to the narrow shell and rubble beaches probably due to cover and food
availability. The adaptations for living in the shell and rubble resulted in the
reduction of the size of the terminal lamellae and increase in the exposure
of the claw as a means of walking and digging in this environment. The longer
toe length affords better traction on a shell and rubble substratum. The large
terminal lamellae adapted to sand or a scansorial habit are unsuited for life
on a shell or rubble beach.
The presence of a few angustidigitus living along a mud beach towards
the southeast part of the peninsula where it connects with the mainland and
also two individuals found in beach dunes on the mainland a few kilometers
away may indicate a recent invasion of the mainland by this species. This may
be due, in part, to the great density of the population now occupying the shell
and rubble beaches. Density dependent factors related to population pressure
may be forcing individuals farther away from their preferred habitat. Hence,
a few hardy individuals are surviving in a marginal habitat which is generally
unfit for the population as a whole.
An individual of P. gerrhopygus was found 25 kilometers from the
easternmost record of P. angustidigitus, with no indication of intergradation
or hybridization between the two species. Thus some period of genetic isola-
tion of the peninsular population has allowed angustidigitus to evolve through
natural selection.
Habitat and natural history. This species is apparently restricted to the
narrow shell and rubble beaches of the Paracas Peninsula and its associated
islands. Of 241 specimens taken from the east side of the peninsula only three
were found more than 10 m from the storm tide zone. One was found beneath a
stone at the edge of a bluff approximately 20 m from the ocean and two were
found about the same distance away on a small granite hill covered with a
thin layer of sand. There are no macroscopic plants on the peninsula and no
surface water. Away from the beach the only physical cover consists of a
few outcrops of granite and small amounts of condensed salt forming a hard
surface crust. The only food items available appear to be silverfish, amphipods,
and spiders that live in beach debris of the storm and intertidal zones. The
paucity of food and cover elsewhere on the peninsula may account for the
concentration of individuals along the storm and intertidal zones. A total of
157 individuals were taken along a 100 m stretch of shell beach in one eve-
ning between the hours of 8:00 and 9 : 30 p.m.
Individuals of this species emerge from cover soon after dark, but they
seem to be most abundant two to three hours following darkness. Several
individuals were observed emerging from beneath dried seaweed and shell
debris about 9 p.m. During the day individuals were found beneath the car-
casses of dead birds, dried seaweed, and assorted trash cover. Seven specimens
were found along a narrow mud beach where the only retreat consisted of
cracks in a vertical mudstone cliff. All specimens were found in cracks one
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Lizards of the Genus Phyllodactylus
19
to two meters above the mud beach. The only food items observed were a
few amphipods under damp seaweed on the beach and pseudoscorpions in
cracks along the mudstone cliff.
Wherever the beach was composed principally of sand, only a few
individuals of this species could be found. These were taken along the base
of cliffs bordering the beach even though there was considerable storm debris
near the beach. The near absence of geckos along these stretches of beach may
be due to the great abundance of ghost crabs occupying the storm and inter-
tidal zones of these areas. Crabs were not observed feeding on geckos but
their nocturnal activity corresponded with that of the geckos, and the crabs
may have been possible predators.
When disturbed, the majority of geckos attempted to escape by running
towards the cliffs above the beach or hiding beneath shell debris. However, on
two occasions individuals ran into the ocean. One immediately turned and
made its way back to shore while the other swam a few meters out to sea.
The latter individual, apparently tired, spread its limbs horizontal to the body
and allowed wave action to return it to shore.
A few individuals were observed climbing boulders and cliffs during their
nocturnal activities. Their actions were slow and deliberate and when dis-
turbed, frequently fell off the surface in an attempt to escape. They seem to
have difficulty in clinging to vertical surfaces and any rapid movement on
their part dislodges their hold on the surface. The absence of large terminal
lamellae normally found in scansorial species is a probable factor in their
inability to move rapidly on vertical surfaces.
Body temperatures were taken from several individuals during their
nocturnal activities and found to be closely associated with the substratum
temperature, varying approximately 0,5 °C above or below the latter tem-
perature.
This species appears to lay only one egg at a time. All gravid females
examined contained one large developing egg and those confined in captivity
laid only one egg.
Distribution. This species is apparently confined to the narrow shell and
rubble beaches of the Paracas Peninsula and associated islands of Peru
although a few individuals may be found on the periphery of the former
area (fig. 2).
Specimens (246) have been examined from: Ica: Islas Sangallan USNM
38570; Has Viejas SDSNH 31093-94; Lagunillas (Gallinazo) Bay FMNH
34337-40; Paracas Peninsula AMNH 36505-07, 3651042; 7.2 km SW Para-
cas LACM 48705, TCWC 27983; 1 km S Puerto Pisco (new) LACM 48593-
627, TCWC 27915, 27944-79; 3 km S Puerto Pisco (new) LACM 48632-704,
48706, TCWC 27984-8062; 7 km S Puerto Pisco (new) LACM 48628-31,
TCWC 27980-82.
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No. 192
Phyllodactylus inaequaiis Cope
Phyllodactylus inaequaiis Cope 1876. Jour. Acad. Nat. Sci., Phila.,
ser. 2, 8:174.
Holotype. Adult female, ANSP 11365, collected by James Orton in
1867-1868. Type-locality, from Pacasmayo, Department of Lambayeque,
Peru.
Definition. A small gecko with a maximum known snout-vent length of
42 mm; tibia and femur tubercles absent; ear denticulation absent; tubercles
on rear of head absent; median row of scales below tail equal in size to adjoin-
ing scales (occasionally widened in some specimens) ; enlarged preanal plate
absent; enlarged dorsal rows of flattened tubercles usually absent at midbody
(occasionally two rows present) ; tail about 45 per cent of total length; tail
tubercles absent; terminal lamellae of digits distinctly widened and truncate;
ground color light gray with dark gray bands usually present; tail usually
banded with dark gray on light ground color; belly brilliant white to light
yellow (fig. 12b).
Diagnosis. Phyllodactylus inaequaiis may be distinguished from ger-
rhopygus, angustidigitus, and heterurus by the absence of an enlarged preanal
plate; from clinatus, dixoni, internadinus, pumilus, johnwrighti, kofordi,
lepidopygus, sentosus, reissi, and ventralis by having less than 10 dorsal rows
of enlarged, flattened tubercles at midbody; from microphyllus by its smaller
maximum snout-vent length (42 mm rather than 56 mm), and by having the
terminal lamellae of the digits distinctly enlarged and truncate with claw
mostly hidden rather than very small terminal lamellae with claw mostly
exposed.
Description. Rostral twice as wide as high, its dorsal edge almost straight
with a short median vertical groove; internasals somewhat rounded, their
median edges in narrow contact, bordered posteriorly by small granules and
postnasal of each side; nostril surrounded by rostral, labial, internasal and
two postnasals; first supralabial in narrow contact with ventral edge of nostril;
shallow depression between internasals, slight depression in frontal region;
scales in posterior loreal region about two to three times larger than scales in
midorbital region; eye large, its diameter contained in snout length about
1.4 times; eyelid with two rows of granules and one large outer row of scales,
last three to five are pointed; diameter of ear contained in eye diameter four
times; ear opening small, scales on anterior edge rounded, subequal, those on
posterior margin rounded but with elevated scales lying within the ear open-
ing; rear of head granular, without larger intermixed tubercles; six to seven
supralabials and five to six infralabials to a point below center of eye; mental
bell-shaped, longer than wide, bordered posteriorly by two postmentals;
postmentals wider than long, their median edges in narrow contact, followed
by transverse row of granular scales.
Distal one-third of dorsum with none to 10 indistinct rows of slightly
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Lizards of the Genus Phyllodactylus
21
enlarged, flattened tubercles, paravertebral row of each side reach to shoulder
in some individuals; paravertebral rows usually separated from each other
by four rows of granules; each tubercle of enlarged dorsal series separated
from preceding tubercle by two to four granules; postanal tubercles number
four on each side, distinct and elevated.
Dorsal surface of upper arm with flattened, rounded scales, forearm
with slightly smaller scales that are rounded and slightly elevated; dorsal sur-
face of thigh and lower leg with rounded, slightly elevated scales; claw short
but visible when viewed from below; terminal pad large, longer than wide
with somewhat rounded edges; about every other median scale on ventral
surface of tail slightly widened; cloacal sacs with external plicate lips.
Color in alcohol: ground color light gray; dorsum with dark gray trans-
verse bands, somewhat broken along midline of back; width of bands equal
to ground color interspaces; dorsal bands somewhat diffuse with white granules
intermixed with few black granules; lateral area of body with suffusion of
intermixed white, gray, and black granules; arms and legs with suffuse spot-
ting of gray, brownish gray, and black granules and scales, without definite
pattern; tail with five dark gray bands, twice as wide as whitish interspaces;
top of head with suffusion of dark gray and ground color, no definite pattern;
dark gray stripe from nostril through eye, over ear, to just posterior of arm
insertion, bordered above and below by pale whitish line from internasal to
just posterior of ear; belly brilliant white.
Variation. Snout-vent length of subadult and adult males varies from
33 to 40 mm (37.0), females from 30 to 42 mm (36.8), juveniles from 19
to 28 mm (23.0), males and females together average 36.9 mm; two post-
mentals in all specimens examined; postmentals contacting first labial in only
54.6 per cent of the sample, first and second labials in 22.7 per cent, and no
labials contact in 22.7 per cent; number of transverse scales immediately fol-
lowing the postmentals vary from 7 to 10 (8.3) ; scales across the midorbital
region range from 20 to 22 (20.8), across the snout at level of third labial
20 to 24 (21.3) ; number of scales bordering internasals vary from 7 to 8
(7.3) with an azygous scale separating the internasals in 3 to 59 specimens;
transverse row of scales across belly vary from 26 to 32 (29.4), longitudinally
55 to 62 (58.4) ; scales on a line between nostril and eye vary from 1 1 to 12
(11.6); lamellae beneath fourth toe vary from 10 to 12 (10.7); enlarged rows
of dorsal tubercles scarcely evident at midbody, none present in seven speci-
mens, two rows present in four specimens; dorsal tubercular rows more evident
across rump and base of tail; rows of tubercles across base of tail vary from
four to eight, six being most common.
Color pattern fairly consistant, with shades of gray ground color and
dark gray bands somewhat variable; bands may be straight or slightly undulat-
ing; all have dark grayish brown spot in center of parietal area; dark head
stripe from nostril to arm in seven specimens, to anterior third of body in
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No. 192
three, to ear in one; body bands (if present) number 4 to 6, five being most
common; tail bands (if present) number 6 to 7 in about equal numbers; tail
bands generally fade towards distal fourth of tail.
Comments. (See comments under Phyllodactylus microphyllus.)
Habitat and natural history. Phyllodactylus inaequalis principally inhab-
its the desert foothills of northwestern Peru. Specimens have been taken in a
variety of foothill localities where the vegetation may be classified as very
sparse to dense.
The Cerro Amotape area rises out of the northern end of the Sechura
Desert and is contiguous to the northeast foothills of the western chain of
the Andes. This area is moderately barren with exfoliating granite boulders
and abundant cacti, but the vegetation is less dense than other foothill local-
ities to the east. Three specimens were obtained in a dry wash on the south
slope of the cerro where scattered small shrubs were present.
The Las Lomas area lies at the foot of the Andes and contrasts with the
Cerro Amotape region in having more dense vegetation, principally thorn
scrub with scattered larger trees. One lizard was found foraging on the ground
beneath small leafless shrubs.
The Abra de Naupe area lies at the foot of the Andes south and east of
the Las Lomas region. This region consists of low hills composed of weather-
ing granite (nonexfoliating) with scattered cacti, mesquite, acacia, and small
amounts of grass. One specimen was found during the day beneath a rotting
cactus.
The Pacasmayo area (type locality) lies near the base of the Andes
where the southern end of the Sechura Desert meets the Jequetepeque River.
The vegetation is very sparse, consisting of mesquite groves along the river and
a few cacti on the foothill slopes. A few individual plants of Cap par is sp.
occur in sandy flats at the base of the foothills. Seven individuals were taken at
dusk from a flat area of hardpan soils at the base of the foothills on the north
side of the Jequetepeque River. All seven specimens were found beneath such
trash items as cardboard, cement sacks, and tarpaper.
The majority of specimens (47) collected by us were taken in the Cerro
de la Vieja region. Although geographically a foothill locality, this cerro is
separated from the main foothills lying several kilometers to the north and
east. The cerro is approximately 150 m in height, with large exfoliating granite
boulders and moderate plant cover. The larger plants of the area are mesquite,
palo verde, cacti, and Capparis sp. (tree form). During the day P. inaequalis
were found beneath a bridge under a stone and under small exfoliating flakes
of granite on large boulders. Individuals were found actively foraging on the
ground or on small boulders at night. Most individuals were found on the
ground within one-half meter of the base of boulders. One lizard was taken
on open soil approximately five meters from the nearest cover, and one indi-
vidual was observed climbing a small shrub.
In all regions mentioned above except for the Pacasmayo area, P. inae-
1970
Lizards of the Genus Phyllodactylus
23
qualis is sympatric with reissi and kofordi. There are however, some micro-
habitat preferences for daytime retreats unique to each species. Large exfoliat-
ing granite flakes were utilized by P. reissi, small flakes by inaequalis, and
small burrows under boulders or cracks in the soil by kofordi. After dark,
P. reissi generally preferred large vertical faces of boulders on which to
forage, P. inaequalis generally on the ground at the base of boulders, and
P. kofordi on boulders and on the ground.
The nocturnal body temperatures of P. inaequalis usually ranged between
the substratum and air temperatures but generally closer to the substratum
than to the air temperature. This was found to be true for other species as well.
This species lays only one egg at a time. Captive females laid only one
egg and preserved gravid females contained a single egg, either in the right
or left oviduct.
Distribution. This species occurs in widely scattered localities in north-
western Peru. It appears to be restricted to the desert foothills bordering the
Sechura Desert (fig. 4). Specimens (59) have been examined from the fol-
lowing localities: Lambayeque: 1 km S Motupe LACM 48495, 48500-18,
TCWC 28308-27, MVZ 82174, 85329-33, 7.2 km W Naupe TCWC 28328.
Libertad: Pacasmayo ANSP 1 1365; 16.7 km ENE Pacasmayo LACM 48496-
99, TCWC 28305-07. Piura: Cerro Amotape MVZ 82170, 82172, 82283;
5 km E Las Lomas MVZ 82171.
Phyllodactylus microphyllus Cope
Phyllodactylus microphyllus Cope 1876. Jour. Acad. Nat. Sci. Phila.,
ser. 2, 8:175
Phyllodactylus inaequalis, (part) Noble 1924. Occ. Paps. Boston Soc.
Nat. Hist., 5:111.
Phyllodactylus Qerrhopycus, (part) Burt and Burt 1931; Bull. Amer.
Mus. Nat. Hist., 61(7) :250.
Holotype. Adult male, ANSP 11364, collected by James Orton in 1868-
69. Type locality. Valley of Jequetepeque, Peru (Department of Lambayeque) .
Definition. An average-sized gecko with a known maximum snout-vent
length of 58 mm; very small terminal lamellae on all digits with fingers and
toes short; dorsal tubercular rows indistinct, composed of small flat, oval
tubercles that seldom form regular rows; enlarged tubercles absent from head,
limbs, and tail; scales of ventral surface of tail uniform in size; scales sur-
rounding nostril greatly swollen; ventral color of body, limbs, and tail bril-
liant white (fig. 9b).
Diagnosis. This species is readily distinguished from P. gerrhopygus,
heterurus, and angustidigitus by the absence of an enlarged preanal plate;
from clinatus, dixoni, lepidopygus, interandinus, kofordi, sentosiis, john-
w right i, reissi, ventralis and pumilus by the absence of enlarged, regular rows
of trihedral tubercles on dorsum; from inaequalis by the presence of a distinct
swollen region about the nostril, larger size, and very small terminal lamellae.
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No. 192
Description. Rostral about twice as wide as high, its outer posterior edges
elevated along inner edge of nostril; internasals small, rounded and swollen,
their median edges separated in most specimens by three rows of granules;
median and posterior borders of internasals contact small granules and post-
nasal of each side; nostril surrounded by rostral, internasal, and two postnasals,
all greatly swollen about nostril; first supralabial not in contact with ventral
edge of nostril; shallow depression between internasals and in frontal region;
scales in posterior loreal region about twice size of midorbital scales; eye
large, its diameter contained in snout length about 1.8 times; eyelid with
2 rows of granules and one larger outer row of scales, last 8 to 12 are pointed;
diameter of ear contained in diameter of eye about 2.8 times; ear opening
small, not denticulate, scales on anterior and posterior margins rounded; rear
of head granular; usually six supralabials and five infralabials to point below
center of eye; mental bell-shaped, slightly longer than wide, its distal tip
extends beyond posterior border of first pair of labials; postmentals much
longer than wide, their median edges usually in narrow contact, nearly sep-
arated by posterior projection of mental; postmentals bordered across chin
by very small granules.
Dorsum with irregular rows of small, oval tubercles that tend to be flat,
posterior dorsal edge with faint keel; dorsal tubercles not regular enough to
count with accuracy; postanal tubercles usually number four on each side;
limbs with somewhat small, flattened scales; main portion of claw visible
when viewed from below; terminal lamellae very small.
Color pattern in life: ground color yellowish tan; dorsum with profusion
of dark brown, brown, light tan, and white granules and larger scales (tuber-
cles) that tend to give entire dorsal color a sandy appearance; arms and legs
similar to dorsum; dorsal surface of tail faintly banded with light gray bands
on yellowish tan ground color; dorsal surface of head similar to dorsum;
loreal region with larger diffuse spots of brown and tan granules; ventral sur-
faces brilliant white.
Variation. Snout-vent lengths of subadult and adult males vary from 33
to 56 mm (46.7), females from 32 to 58 mm (46.5), juveniles 23 to 31 mm
(27.1 ), males plus females average 46.6 mm; tail length varies from 42 to
53 per cent of total length; postmentals number from two to four, two occur-
ring in 90.6 per cent three in 8.0 per cent and four in 1.4 per cent of sample;
postmentals contacting adjacent labials vary from none to two on each side,
none in 4 per cent, one on one side and none on opposite side in 4 per cent,
one on each side in 89.2 per cent one on one side and two on opposite side
in 1.4 per cent and two on each side in 1.4 per cent of sample; number of
scales across chin immediately following postmentals varies from 6 to 9 (7.4);
scales across midorbital region range from 18 to 25 (20.8), across snout at
level of third labial 18 to 26 (21.8); median and posterior scales bordering
internasals vary from 7 to 12 (8.7), with median edges of internasals sep-
arated by granules in 99.2 per cent of sample; scales from nostril to eye range
1970
Lizards of the Genus Phyllodactylus
25
from 10 to 15 (12.4); scales across venter vary from 21 to 38 (27.3), main-
land populations from 26 to 38 (31.0), island populations 21 to 33 (25.8);
scales from throat to vent range from 62 to 72 (66.3) in all populations; dor-
sal tubercular rows range from 10 to 14 (11.9) when arranged in somewhat
regular rows, but majority of individuals have irregular rows; lamellae beneath
fourth toe vary from 12 to 15 (13.3); terminal lamellae generally very small,
but variable in size dependent upon soil type from which specimens are
obtained; tail generally shorter than snout-vent length and always devoid of
tubercular rows, ventral surface with all scales of equal size.
Dorsal color pattern variable, from light brown, brown, and white
freckled appearance on yellowish tan ground color to dark brown to black
reticulating lines on gray, tan or light brown ground color; broad, but indis-
tinct brown bands are occasionally present in adults, more frequently in
young and juveniles; a brownish line usually present from nostril to eye,
occasionally extends to ear; ventral surfaces brilliant white.
Comments. There has been considerable confusion in the application of
the name microphyllus to the original population Cope described in 1876.
Cope described microphyllus and inaequalis in the same paper, but failed to
make clear the type of digital expansion found in inaequalis that would sepa-
rate it from microphyllus. Noble (1924) used the name inaequalis (— micro-
phyllus) for populations inhabiting the sandy deserts of the coast, apparently
being unable to distinguish between Cope’s two species. Cope (1877) later
described another species, nigrofasciatus, and in the same paper explained the
differences in digital expansion between microphyllus and inaequalis, appar-
ently overlooked by Noble and later workers.
Burt and Burt (1930), apparently confused by Noble’s use of the name
inaequalis for the sandy desert form, listed microphyllus, inaequalis, and
gerrhopygus from Peru, but had misidentified a specimen of gerrhopygus as
being inaequalis. They later (1931) pertpetuated this mistake by listing inae-
qualis (= gerrhopygus) from Arequipa, Peru, and in part, confused speci-
mens of microphyllus with inaequalis in northwestern Peru. This mistake was
repeated by Donoso-Barros ( 1966).
Habitat and natural history. Phyllodactylus microphyllus is a widespread
species, but generally restricted to the western sandy areas of the Sechura
Desert, sandy and sand rubble beaches of the Peruvian coast and offshore
islands. The absence of vegetation does not seem to restrict its distribution in
the drier parts of Peru. We have found specimens in areas where vegetation
was absent, but these individuals were found within one kilometer of the
beach. The storm and intertidal zones of the beach are probably the principal
sources of food. The absence of food elsewhere is a possible factor limiting
the distribution of this species. In northwestern Peru where vegetation is more
abundant this species has been taken 60 kilometers (airline) from the nearest
beach.
During the day individuals have been found under rocks, cow chips.
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No. 192
boards, bones, paper, tin, and other objects strewn on sandy surfaces. In one
instance individuals were found in open Indian graves where remnants of
burial cloth were exposed. Some of these daytime retreats are often fully
exposed to sunlight and the body temperatures of individuals found under
these conditions are about 10°C higher than their nocturnal temperatures. In
the vicinity of Culebras, this species was observed emerging just prior to com-
plete darkness and actively foraging for about four hours. In the Cerro Illescas
area of northwestern Peru this species was found foraging in leaf litter and
on the branches and trunks of shrubs, mainly Cap pans avicennifolia, during
winter months. We observed this species foraging on guano mounds on Cerro
Aziil and it was reported from guano mounds on Isla Lobos de Afuera by
Burt and Myers ( 1942).
The behavior of this species is similar to that of members of the genus
Coleonyx. When disturbed, the tail is raised, back is arched, and it feigns an
attack accompanied by noise and walks diagonally away with full side view
of body exposed to a potential predator. The movements of this lizard are
deliberate, somewhat slow, and when disturbed, it does not seem able to run
rapidly. One individual was observed utilizing a small rock to scrape shedding
skin from its body.
This species apparently lays a single egg per clutch. Preserved females
contained a single egg in either the right or left oviduct.
One specimen was removed from the stomach of a snake, Liophis sp.,
but actual predation was not observed. Potential predators consist of ghost
crabs, snakes, and foxes.
As in other species of Phyllodactylus, the nocturnal body temperatures
of this species closely approximates that of the substratum.
Distribution. This species is known only from Peru. It ranges from the
northwestern coastal village of Los Organos, southward to the coastal village
of Cerro Azul, an airline distance of 1,150 km. The western part of the Sechura
Desert of northwestern Peru is the only area where its range extends inland
from the coast (fig. 2).
Specimens (277) have been examined from the following localities :
Ancash: 2 km N Culebras LACM 48589-92, TCWC 28151-55. 12 km N,
1 km W Pativilca LACM 48570-72, TCWC 28148-50. 24.7 km N Pativilca
MVZ 82145, 82231. Lambayeque: 1 1 km S Chiclayo MVZ 82181. Eten
MCZ 18138, 17692-94, AMNH 28517-18. 10 km S Mocupe TCWC 28162.
3 km SE Morrope MVZ 82146. Libertad: Jequetepeque Valley ANSP 11364.
4 km N Pacanguilla LACM 48574, TCWC 28156-57. Lima: Ancon AMNH
36513-19. Cerro Azul LACM 48567-69, TCWC 28145-47. 7 km SSE Chilca
MVZ 85430-34, 85436-38, 85440-41. Chorillos MJP 3 (no numbers). Cruz
de Hueso USNM 75390. Isla San Lorenzo USNM 17541. Lima MJP 5 (no
numbers), 3 km N San Antonio TCWC 28174. 5.5 km NE San Bartolo
MVZ 85421-29. Piura: 1 km S Bayovar, Cerro Illescas MVZ 85353-61. 5 km
SSE Bayovar MVZ 85342-52. 12 km S, 8 km E Bayovar MVZ 82157-61,
1970
Lizards of the Genus Phyllodactylus
27
82270. 20 km SE Bayovar MVZ 85362-63. Isla Lobos de Afuera SDSNH
31123-53, USNM 38571-74. Isla Lobos de Tierra FMNH 34226(7),
34234 (5), 40113-14, AMNH 38753-55, UMMZ 83099 (2). 3.7 km ENE
Los Organos LACM 48588, TCWC 28173. Negritos FMNH 5726-27, 8363,
8453-54. Paita AMNH 28461, HM 1499 (2). mouth of Parinas Valley
FMNH 41543-45. 13 mi N Piura CAS 85163. 10 km E Piura MVZ 82162.
Punta Aguja MVZ 82148-52, 82228, 82248. Reventazon MVZ 82153-56.
30 km SSE Sechura MVZ 82147. Talara FMNH 53891, 53894, 57457. 2 km
E Talara LACM 48575-78, TCWC 28158-61. 2 km N Talara TCWC 28163.
3 km S Talara LACM 48579-86, TCWC 28164-71. 5 km S Talara LACM
48587, TCWC 28172. Trie Trac MVZ 82163-68, 85334-41.
Phyllodactylus clinatus sp. nov.
Holotype. Adult female, MVZ 82227, collected by Raymond B. Huey,
19 July 1967. Type locality. Punta Aguja, 37 km SW Sechura, Department of
Piura, Peru.
Definition: A small gecko with a known snout-vent length of 46 mm;
dorsum with 10 to 12 rows of enlarged, trihedral tubercles; tubercles absent
on tail, thigh, tibia, and forearm; granules absent between internasals; ear
very large, 56.7 per cent of diameter of eye; ear strongly denticulated on
anterior and posterior margins; terminal lamellae of digits large, truncate;
all scales of median row beneath tail widened, rectangular (fig. lib).
Diagnosis. This species may be distinguished from P. heterurus, angusti-
digitus, and gerrhopygus by the absence of an enlarged preanal plate; from
inaequalis and microphyllus by much larger ear, dorsal rows of enlarged,
trihedral tubercles arranged in regular series, and by a widened row of scales
beneath tail; from dixoni, pumilus, kofordi, sentosus, johnwrighti, reissi, and
ventralis by the absence of tubercles on tibia and much larger ear; from
lepidopygus and interandinus by presence of a row of widened, rectangular
scales beneath tail, larger ear, and small, flattened granular scales on tibia.
Description of holotype. Rostral twice as wide as high with short median
groove about one-third depth of rostral; two internasals, somewhat rectangu-
lar, their median edges in broad contact, and bordered posteriorly by three
granules and postnasal of each side; nostril surrounded by rostral, labial,
internasal, and two postnasals; first labial in broad contact with ventral edge
of nostril; deep depression between internasals, moderate depression in frontal
region; scales in posterior loreal region about 2.5 times larger than midorbital
scales; 22 scales across snout at level of third labial, 14 scales across anterior
edge of orbits, 20 midorbital scales; eye large, its diameter contained in snout
length 1.5 times; eyelid with two rows of granules and one larger outer row
of scales, last 4 to 5 are pointed; ear diameter contained in eye diameter 1.8
times; ear opening large, denticulate; rear of head granular with larger inter-
mixed tubercles; six supralabials and five infralabials to point below center of
eye; mental bell-shaped, as wide as long; bordered posteriorly by two post-
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No. 192
mentals; postmental as wide as long, their median edges in broad contact,
immediately followed by transverse row of 9 scales, followed by second row
of 13 smaller scales; postmentals contact first labial only of each side.
Dorsum with 10 rows of enlarged, trihedral tubercles that are somewhat
flattened; paravertebral row with 45 tubercles from head to base of tail,
24 between axilla and groin; paravertebral rows separated from each other
by 4 to 6 rows of granules; 4 rows of dorsal tubercles reach to nape, 6 to base
of tail; each tubercle of enlarged dorsal series separated from preceding
tubercle by 1 to 3 granules; postanal tubercles number two on each side,
indistinct; 26 scales across venter; 60 from throat to vent; dorsal surfaces of
arm with granules and larger flattened scales; dorsal surface of thigh with
granules and larger flattened scales, tibia with small flat granules; lamellae
formula for hand 6-8-10-13-9, foot 7-9-11-14-10; claw hidden when viewed
from below; terminal lamellae large, longer than wide, truncate; digits slender.
Measurements in mm: snout-vent length 46.0, axilla-groin length 19.0,
length of leg 16.5, length of arm 14.4, length of tail 46.0, length of head
14.0, depth of head 4.9, width of head 8.7, length of snout 4.9, diameter of
eye 2.3, diameter of ear 1.8, distance from eye to ear 3.6.
Color in life: ground color sand brown; dorsum with six narrow, choco-
late brown bars interrupted along median line, ground color interspaces four
to five times width of dark bars; sides of body with light tan spots between
dark bars, tending to form ocellated pattern with network of fine brown lines
intermixed; arms with chocolate brown reticulating lines, tending to form
bands on upper arm; hind limb with five chocolate brown bands, one-third
as wide as ground color interspaces; tail with 1 1 dark brown bands, ground
color interspaces twice width of dark bands; head with dark brown parietal
spot, rear of head with thin, dark brown reticulating lines: dashed and partly
wavy line from rear of eye to hind limb, connecting with lower edge of dark
body bars; supralabials with upper half heavily pigmented with dark brown,
lower half and all of infralabials with suffusion of minute black dots; dark
brown stripe from nostril to eye, tending to be black from eye to side of
body; ventral surfaces with faint brownish dots scattered along sides, more
dense on tail; general ventral color pale yellow.
Variation. Snout-vent length of juvenile male 28 mm, females vary from
34 to 46 mm (40.3); tail length about 45 to 50 per cent of total length; post-
mentals number 2 in all, contacting first labial only in all; number of scales
across chin immediately following postmentals range from 6 to 9 (7.3); scales
across snout at level of third labial range from 21 to 23 (21.8), midorbital
scales from 19 to 22 (19.5); scales bordering posterior edge of internasals
number 5 in all; scales from nostril to eye range from 13 to 15 (13.8); scales
across venter vary from 26 to 32 (28.8), from throat to vent 60 to 64 (61.8);
dorsal tubercular rows range from 10 to 12 (10.5); paravertebral tubercles
from head to base of tail range from 42 to 46 (44.5), from axilla to groin
24 to 29 (27.5); lamellae beneath fourth toe range from 13 to 15 (14.0);
1970
Lizards of the Genus Phyllodactylus
29
terminal lamellae of digits large and truncate in all; ratio of ear diameter to
that of eye ranges from .500 to .653 (.567); ratio of ear diameter to distance
between eye and ear ranges from .419 to .487 (.444) .
Dorsal color pattern varies from distinct bands, broken along median
line, to series of dorsolateral spots connected longitudinally by thin dark
brown line; bands or spots (one side) vary from 6 to 8 (6.5) ; tail bands num-
ber 11 in only specimens with complete tail; limbs usually banded but bands
may be obscured by dark ground color; ventral surfaces pale yellow in all
specimens.
Habitat and natural history. Having taken only four individuals of this
species, very little natural history information of the species could be obtained.
All four specimens were taken from the vicinity of Cerro Illescas where Cap-
paris avicennifolia, C. scabrada and mesquite are the dominant plants.
Three specimens were obtained in July and August of 1967. One was
taken from a snap trap at the base of a small boulder near the edge of a cliff.
The substratum was sandy, although it was only one-third meter from a frag-
mented shale area. One was captured on the trunk of a mesquite about one m
above the ground. The third specimen was taken one afternoon under a small
rock on sandy substratum, near rock outcrops and mesquite.
In spite of intensive collecting in this general area in 1968, only one
additional specimen was obtained. It was found 2.5 m above the ground on a
mesquite branch.
Distribution. This species is known only from the Cerro Illescas area of
northwestern Peru (fig. 4).
Four specimens have been examined: Piura: 4 km SSE Bayovar MVZ
82271; 5 km SSE Bayovar TCWC 28332; 10 km SE Bayovar MVZ 82269;
Punta Aguja MVZ 82227.
Phyllodactylus lepidopygus (Tschudi)
Diplodactylus lepidopygus Tschudi 1844-46, Untersuchungen uber die
Fauna Peruana. Herp.: 38.
Discodactylus phacophorus Tschudi 1844-46, Untersuchungen uber die
Fauna Peruana. Herp.: 38-39.
Phyllodactylus nigrofasciatus Cope 1877. Proc. Amer. Philos. Soc.
Phila., 17:36.
Phyllodactylus variegatus Werner 1901. Abh. Mus. Dresden, 9(2): 2.
Holotype. NM 43, adult male, obtained by Tschudi prior to 1844. Type
locality, Chorillos, Peru.
Definition. A moderate-sized gecko with known maximum snout-vent
length of 55 mm; tibia and femur tubercles absent; ear denticulate on anterior
margin; slightly enlarged tubercles among granules on rear of head; median
row of scales on ventral surface of tail not enlarged; enlarged preanal scale
absent; 10 to 16 (13.1) rows of enlarged trihedral tubercles on dorsum; tail
length about 45 to 55 percent of total length, tail without enlarged tubercles;
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No. 192
terminal lamellae of digits moderately enlarged; body bands (may be broken
on midline) 6 to 9 (7.2), tail bands 9 to 13 ( 1 1.0) (fig. 13a).
Diagnosis. This species is easily distinguished from P. heterurus, angusti-
digitus, and gerrhopygus by the absence of an enlarged preanal scale; from
kofordi, reissi, cUnatus, dixoni, johnwrighti, pumilus, ventralis, and inter-
andinus by having all scales beneath the tail of equal size, rather than median
row distinctly widened or enlarged; from inaequalis and microphyllus by hav-
ing distinct dorsal rows of enlarged trihedral tubercles, rather than indistinct
dorsal rows of small flattened tubercles or rows occasionally absent; from
sentosus by the absence of tail, tibia, and thigh tubercles, rather than very
large tubercles present.
Description. Rostral more than twice as wide as high; two internasals
and two postnasals; nostril bordered by rostral, internasal, labial, and two
postnasals; second supralabial separated from nostril by two granules and
postnasal; posterio-dorsal loreal scales about four times larger than inter-
orbital scales; internasals slightly wider than long, bordered posteriorly by
small scales; auricular opening very small, slightly denticulate, contained in
snout length about eight times; eye large, contained in snout length about one
and one-third times; rear of head granular with few larger, rounded tubercles;
7 to 9 supralabials and 5 to 6 infralabials to point below center of eye; mental
slightly longer than wide, bordered posteriorly by two to three well defined
postmentals; postmentals followed by irregular row of granular scales; eyelid
with two rows of granules and larger palebral row of scales; last 2 to 5 scales
in larger supercilliary series bearing long spines; regular rows of longitudinal,
enlarged keeled tubercles on dorsum, 8 rows reach to rear of head and 8 to
base of tail; all rows equally spaced apart, no wide granular area separating
paravertebral rows; four well defined postnasal tubercles on either side of
anus in males; venter scales moderately differentiated from lateral body
scales; no enlarged preanal shield; arm covered with flat imbricated scales,
no enlarged tubercles intermixed with flattened scales; femur with flat, imbri-
cated scales on dorsal and ventral surfaces, posterior-ventral surface with
granular scales; lower leg similar to femur; terminal pads slightly longer than
wide; claw short, hidden between terminal pads.
Variation. Specimens of this species with a minimum snout-vent length
of 32 mm are considered young adults. In a sample of 36 specimens, 8 are
less than 32 mm snout-vent length, 13 are males with snout-vent from 32 to
50 mm (40.8), 15 are females with snout-vent from 36 to 55 mm (46.8) .
The average snout-vent length for males and females is 44.0 mm.
The number of postmentals varies from two to four, two occurring 58.4
per cent of the time, three 38.9 per cent and four 2.7 per cent; number of
scales bordering the postmentals range from 5 to 8 (6.9) ; number of scales
across the midorbital region 15 to 21 (18.2); number of scales across the
snout at the level of the third supralabial 17 to 22 (20.1); scales bordering
the internasal shields posteriorly number 5 to 1 1 (6.7), with one or two small
1970
Lizards of the Genus Phyllodactylus
31
azygous scales separating the internasals in 7 of 36 specimens; transverse
venter scales number 26 to 34 (29.4), longitudinally 65 to 74 (69.6); scales
from posterior edge of nostril to eye number 10 to 14 (11.4); fourth toe
lamellae number 10 to 14 (11.5); dorsal tubercular rows number 10 to 16
(13.1); tubercles in paravertebral row from rear of head to base of tail num-
ber 42 to 53 (47.8), axilla to groin 24 to 34 (28.9); body bands or blotches
number 6 to 9 (7.2), tail bands 9 to 13 ( 1 1.0).
The general ground color in life varies from a greenish brown to a deep
blackish brown; dorsal body markings vary from black spots to brown cross-
bands or light tan blotches; general appearance consists of light ground color
with darker spots irregularly arranged; head with blackish line from nostril
through eye to above arm, occasionally continuing along side of body to hind
limb: top of head from prefrontal area to posterior tip of parietal region con-
sistently darker than remainder of body in all specimens but one; one speci-
men has reversal pattern of dark ground color with light blotches; tail gen-
erally banded with alternating black or brown bands with light tan to whitish
interspaces; interspaces usually about half the width of dark bands but occa-
sionally as wide or wider than dark bands; regenerated tails usually reticulated
with dark lines on tan ground color; juvenile pattern similar to that of adults;
venter yellowish tan to yellowish brown with dark brown spotting along lateral
edges or occasionally to center of venter; chin and throat creamy yellow in
young and subadults, becoming progressively darker with suffusion of brownish
dots in old adults; ventral surface of tail generally darker than venter.
Comments. We have examined the holotype of P. lepidopygus, three
syntypes of P. phacophorus, holotype of P. nigrofasciatus and find that they
are identical in general shape, size of head and body. Their scale characters
are within the range of variation of a relatively large sample of P. lepidopygus
from a single locality and we therefore consider P. phacophorus and P. nigro-
fasciatus as subjective synonyms of P. lepidopygus. The nine syntypes of
P. variegatus were destroyed during the fire bombing of Dresden, Germany,
during World War II and Werner’s (1901) original description is the only
evidence we have for a comparison of P. lepidopygus and P. variegatus.
Werner’s description agrees with the characters attributed to P. lepidopygus.
His description agrees to some extent with the salient features of P. reissi, a
species found further to the north. The major exception is the presence of
tibia tubercles in P. reissi, absent in variegatus, and to a minor degree, the
short outer row of dorsal tubercles in P. reissi, not so in variegatus. Werner
gave the type locality as Lima (9 specimens) and Chanchamayo ( 1 specimen),
both areas some 1,500 km south of the nearest locality where P. reissi has
been taken. However, P. lepidopygus occurs in the vicinity of Lima and
P. variegatus is considered a synonym of P. lepidopygus.
Habitat and natural history. This species has been found associated with
the lomas vegetation of the Lima region, the drier Tillandsia association of
the desert coast, and the coastal foothills to an elevation of 1,400 m. The
32
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Figure 3. Distribution of P. pumilus, P. kofordi, P. lepidopygus, and P. interandinus
in western South America.
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Lizards of the Genus Phyllodactylus
33
microhabitat appears to be one of vertical surfaces of buildings, rock walls,
large and small granite boulders, and crevices therein.
A series of 34 specimens taken by Huey, Dixon, and Wright from Lomas
de Lachay were found exclusively on vertical surfaces of granite boulders,
rock walls, or walls of buildings. No specimens were observed on rocks with
abundant mossy growth, but lichen covered surfaces were utilized. Occasion-
ally P. lepidopygus were seen in crevices or under smaller rocks resting on
boulders during the day. This species became active as soon as it became dark,
even on cold misty nights. The air temperature on 4 July 1967 was 13.2°C.
approximately an hour and half after nightfall and seven geckos were taken
subsequent to that time. Potential food items (e.g., Porcellio, spiders, moths)
in the Lomas de Lachay region were plentiful on the rock surfaces at night.
One specimen taken 24.7 km north of Pativilca was collected from a burro
femur lying in a rolling sand hill region with few exposed surface rocks and
Tillandsia as the only plant.
The only gecko observed in the Lomas de Lachay region was P. lepido-
pygus, but the rolling sand hill region 123 km north of Lima also contained
P. microphyllus.
Of four specimens taken from the Asia area south of Lima, three were
found actively foraging at night on a rock corral wall, and one was found
on a small cerro on a rotting granite outcrop. All four were found on surfaces
relatively free of heavy lichen growth.
Two individuals were found east of Chosica at an elevation of 1,700 m.
Both specimens were taken from a wall of an abandoned adobe building that
had been plastered. The coloration of both specimens was atypical, being
almost translucent, without pattern, and inconspicuous against the light colored
plaster. The two individuals had a more normal color pattern following a
24-hour period in a cloth sack.
An examination of the oviducts of preserved females reveals that this
species lays a single egg per clutch. Natural nests were not obesrved but we
suspect that eggs are deposited in cracks of large boulders in the Lomas de
Lachey area. Several shells of Phyllodactylus lepidopygus eggs were found
40 km NNW Pativilca, 600 m, in dead cactus plants in a dry arroyo strewn
with small boulders. No geckos were observed at this locality, but the location
it within the known distribution of lepidopygus.
Distribution. The known range of this species is approximately 300 km
north and 300 km south of Lima, and appears to be restricted to the rocky
foothills from near sealevel to an elevation of 1,700 m (fig. 3).
Specimens (63) have been examined from the following localities:
PERU, no specific locality BM 25440. Ancasli: Chimbote Valley ANSP 1 1366.
24.7 km N Pativilca MVZ 82175. Lima: 2 km ESE Asia LACM 48473-74,
TCWC 28142-43. 8 mi SE Chilca MVZ 85435. Chorillos NM 43. vicinity
of Chorillos NM 41 (3). Chosica FMNH 34175. 26 km ENE Chosica LACM
48475, TCWC 28144. Lima MJP (3) unnumbered. Lomas de Lachey CAS
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85143, LACM 48467-72, MVZ 82182-91, 82222-25, 82252-65, TCWC
28135-41. 20 km E Lurin MVZ 85411; Verrugas Canyon MCZ 26674. lea:
lea FMNH 39367.
Phyllodactylus interandinus sp. nov.
Phyllodactylus phacophorus, (part) Noble 1924, Occ. Paps. Bost. Soc.
Nat. Hist., 5: 111.
Holotype. Adult female, TCWC 27914, collected by John W. Wright and
James R. Dixon, 26 November 1968. Type locality, 4.8 km S (by road) Bel-
lavista, 520 m. Department of Cajamarca, Peru.
Definition. A small gecko with a known snout-vent length of 47 mm;
numerous rows of enlarged dorsal tubercles and tubercles in paravertebral
row from head to tail and axilla to groin; terminal lamellae moderately large,
digits short; distinctly banded, black, white, and yellowish green tail; tubercles
absent on thigh, tibia, and forearm; distinct dorsal color pattern of three light
and four dark longitudinal stripes (fig. 10a).
Diagnosis. This species is distinguished from P. heterurus, angusti digitus
and gerrhopygus by the absence of an enlarged preanal shield; from inaequalis
and microphyllus by having some scales beneath tail enlarged, and by numer-
ous rows of enlarged trihedral tubercles on dorsum; from clinatus, dixoni,
ventralis, lepidopygus, kofordi, sentosus, johnwrighti, pumilus, and reissi by
presence of 65 or more tubercles in paravertebral row from head to base of
tail, and distinct dorsal color pattern of light and dark stripes.
Description of holotype. Rostral twice as wide as high, its dorsal edge
with vertical groove one-half depth of rostral; two internasals, somewhat
rounded, their median edges in broad contact, bordered posteriorly by six
granules and postnasal of each side; nostril surrounded by rostral, labial, inter-
nasal, and two postnasals; first supralabial in narrow contact with ventral edge
of nostril; shallow depression between internasals and in frontal region; 12
scales on line between nostril and eye; scales of posterior loreal region about
two times larger than midorbital scales; 22 scales across snout at level of third
labial, 14 scales across anterior edge of orbits, 16 midorbital scales; eye large,
its diameter contained in snout length 1.6 times; eyelid with two rows of
granules and one larger outer row of scales, last three are pointed; ear con-
tained in eye diameter 2.6 times; ear opening denticulate, but denticulate
scales low and somewhat rounded; rear of head with subequal scales; five
supralabials and four infralabials to point below center of eye; mental bell-
shaped, slightly longer than wide, bordered posteriorly by three postmentals;
postmentals rounded, their median edges in broad contact, immediately fol-
lowed by transverse row of 8 scales, followed by second row of 12 smaller
scales; postmentals contact first labial only on each side.
Dorsum with 20 longitudinal rows of enlarged trihedral tubercles, some-
what flattened, paravertebral row with 7 1 tubercles from head to base of tail,
44 between axilla and groin; paravertebral rows separated from each other by
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Lizards of the Genus Phyllodactylus
35
Figure 4. Distribution of P. johnwrighti, P. inaequalis and P. clinatiis in western
South America.
36
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No. 192
5 rows of granules, each tubercle of paravertebral row separated from preced-
ing tubercle by 0 to 1 granule; 14 rows of tubercles extend across nape, 8 at
base of tail; postanal tubercles number three on each side; 34 scales across
venter, 58 from throat to vent; dorsal surface of arm with small flattened
scales; dorsal surface of thigh without tubercles; tibia with subequal rounded,
somewhat elevated scales; lamellae formula for hand 6-7-8-9-7, foot 6-9-10-
11-10; claw barely visible when viewed from below; terminal lamellae moder-
ately large, slightly longer than wide, rounded.
Measurements in mm: snout-vent length 41.0, axilla-groin length 18.6,
length of leg 15.2, length of arm 12.1, length of tail 43.0, length of head
11.3, depth of head 5.2, width of head 8.1, length of snout 4.2, diameter of
eye 2.6, diameter of ear 1.0, distance from eye to ear 3.3.
Color in life: ground color dark gray; dorsum with one median and two
dorsolateral grayish white stripes, two dorsal and two lateral dark brown
stripes; median light stripe from rear of head to base of tail, anterior part of
median stripe bifurcates and extends anteriorly to above each eye; dorsal dark
stripes tend to form series of loosely connected spots because of small clusters
of grayish white spots scattered in dark fields; lateral dark stripe extends from
snout to groin; limbs heavily mottled with grayish white and grayish black
spots; tail with 1 1 black bands, posterior edge of each black band with white
tipped scales; interspaces three times width of black bands, greenish yellow;
dorsal surface of head with diffuse dark brown lines; ventral surfaces dusky,
chin generally whitish, each scale with black dots.
Variation. Snout-vent lengths of subadult and adult males range from
32 to 45 mm (39.2), females 33 to 47 mm (41.1), young and juveniles 22 to
30 mm (29.0), males plus females average 40.1 mm; tail lengths vary from
48 to 55 per cent of total length; postmentals range from two to four, two
occurring in 58.4 per cent, three in 38.9 per cent, and four in 2.7 per cent of
sample; chin scales bordering postmentals range from 5 to 8 (6.9) ; postmentals
contact first labial only of each side in 99.6 per cent, first labial on one side,
none on opposite side in 0.4 per cent of sample; scales bordering internasals
number 5 to 7 (6.2); scales from nostril to eye vary from 9 to 12 (10.8);
scales across snout at level of third labial range from 22 to 27 (24.3), mid-
orbital scales 18 to 22 (20.3); scales across venter vary from 26 to 33 (29.6),
from throat to vent 56 to 65 (60.8); fourth toe lamellae range from 10 to 12
(11.6); longitudinal rows of dorsal tubercles vary from 14 to 22 (17.4);
tubercles in paravertebral row from head to base of tail range from 65 to 85
(70.1 ), between axilla and groin 35 to 60 (40.7).
Color pattern consists of one median and two dorsolateral grayish white
to yellowish tan stripes, separated by two dorsal and two lateral grayish brown
to grayish black stripes; dark stripes usually bifurcates on nape, each arm
extending anteriorly to posterior edge of eye or to tip of snout; broad, dark
brown stripe usually present from snout to rear of head, lying between dor-
solateral light stripes; ventral surfaces usually dusky with black dots on each
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Lizards of the Genus Phyllodactylus
37
Figure 5. Distribution of P. reissi, P. heterurus and P. sentosus in western South
America.
38
Contributions in Science
No. 192
scale, chin and throat generally lighter in color than remainder of body;
dorsal surface of tail with 11 to 16 narrow black bands edged with white
posteriorly; broad interspaces bright reddish orange in juveniles, yellowish
orange in subadults, yellowish green in adults.
Comments. '^oh\Q (1924) labeled a relatively large series of specimens
from northwestern Peru as P. phacophorus, but the majority of the specimens
are referable to P. interandinus. A critical examination of the entire series
reveals four species were present in the collection, interandinus, johnwrighti,
kofordi, and reissi.
Habitat a?td natural history. Noble (1924) recorded taking phacophorus
(—interandinus) and magister (^reissi) from houses and deserted huts in Bel-
lavista. We were unable to obtain interandinus in Bellavista but found reissi
on the walls of houses. Phyllodactylus interandinus was abundant on a thorn
scrub hillside a few kilometers south of Bellavista. Standing, dead Cereus sp.
are the principal daytime retreats of this species with as many as seven indi-
viduals occupying one plant. Some individuals were taken as high as two
meters above the ground, but the majority of specimens were found within
one meter of the ground, usually beneath the bark, in the hollow trunks or
branches. Phyllodactylus reissi and interandinus were taken from the same
cactus plant, but the former species was not abundant. The distribution of
interandinus is spotty, for similar areas of vegetation on other hillsides yielded
only reissi. Hillsides and valleys were examined in the Department of Ama-
zonas, east of the Rio Marahon, but only reissi were found in abundance. Only
six individuals of interandinus were found east of the river, taken beneath
rotting agave stumps and dead cactus litter. A total of 79 reissi were taken
east of the river in a wide variety of habitats, including agave.
There seems to be potential competition between these two species for
space but not for food. Phyllodactylus reissi is primarily scansorial while
interandinus forages on the ground. Eggs of both species were found in a
variety of rotting debris, but reissi eggs were far more numerous than those
of interandinus.
An examination of preserved females of both species indicates that P.
interandinus lays one egg per clutch and reissi two eggs per clutch.
There are distinct behavioral differences between P. interandinus and
reissi when suddenly exposed to sunlight while in their daytime retreats. Phyl-
lodactylus interandinus remains motionless for a few seconds, then seeks
cover under debris within a few centimeters of where it was uncovered. Phyl-
lodactylus reissi immediately runs up or down the plant upon which it was
exposed, or jumps to the ground and runs to the nearest cover to hide.
The uneven distribution of P. interandinus in the Maranon Valley sug-
gests that its niche requisites are more restricted than those of reissi.
Distribution. This species is an inhabitant of the Chinchipe, Maranon,
and Utcubamba river valleys of the departments of Amazonas and Cajamarca,
Peru. The locality of a series of specimens in the Museum of Zoology, Uni-
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Lizards of the Genus Phyllodactylus
39
versity of Michigan, is given as Sullana, Peru. This locality is approximately
70 km west of the Andes, and lies at the north end of the Sechura Desert. The
specimens are in a similar state of color and preservation as those collected
by Noble in 1916. We believe this locality to be in error, and represent some
of the specimens taken by Noble from the Chinchipe or Maranon river valley
(fig. 3).
Specimens (149) have been examined from the following localities:
PERU: Amazonas: 17 km S Bagua Chica LACM 48538, TCWC 28087.
10 km WSW Bagua Grande MVZ 82178-80, 82251. Cajamarca: Bellavista
AMNH 39338, 39341-42, MCZ 18139, 18140(80), SMFM 8295, TCWC
24924-27. 4.8 km S Bellavista LACM 48519-37, 48539, TCWC 27914,
28066-68, 28070-86. Perico AMNH 28428-36. Pima: Sullana UMMZ
59034(4) (in error).
Phyllodactylus kofordi sp. nov.
Holotype. Adult male, TCWC 27912, collected by James R. Dixon and
John W. Wright, 19-20 Nov. 1968. Type locality. 7 km S Motupe, Cerro de la
Vieja, Department of Lambayeque, Peru, 150 m elev.
Definition. A small gecko with a maximum known snout-vent length of
46 mm; tibia and femur tubercles present; ear denticulate on anterior and
posterior margins; moderately enlarged tubercles on rear of head intermixed
among granules; median row of scales below tail distinctly widened; enlarged
preanal scale absent; 12 to 14 (12.1) rows of enlarged trihedral tubercles on
dorsum; tail 50 to 55 per cent of total length; tail with an 8-6-4-2 or 6-6-4-2
reduction of enlarged tubercles per whorl from base to distal half of tail;
terminal lamellae of digits moderately enlarged, truncate; body markings of
three types, with spots, dots, or uniform coloration; tail markings of two types,
uniform or banded; trihedral tubercles of tibia greatly enlarged (fig. 11a).
Diagnosis. This species may be distinguished from P. heterurus, angiisti-
digitus and gerrhopygus by the absence of an enlarged preanal plate; from
johnwrighti, pumilus, interandiniis, clinatus, inaequalis, lepidopygus, and
microphyllus by the presence of greatly enlarged trihedral tubercles on the
dorsum, thigh, tibia, and forearm and by whorls of tubercles on the entire
length of dorsal surface of the tail; from sentosus by having much larger
terminal lamellae, smaller dorsal tubercles, and smaller snout-vent length;
from re is si by the presence of tail tubercles, tubefcles on the thigh, smaller
body size (maximum 46 mm rather than 75 mm), fewer paravertebral tuber-
cles from rear of head to base of tail (31 to 36 rather than 46 to 60), and
fewer paravertebral tubercles between axilla and groin (16 to 21 rather than
28 to 38) ; from dixoni and ventralis by the latter two characters and smaller
maximum snout-vent length.
Description of holotype. Rostral twice as wide as high; depression
between internasals continues posteriorly through frontal and interorbital
regions; internasals bordered posteriorly by 5 granular scales; 18 scales across
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snout between third labials, 13 across snout between second labials; 14 between
middle of orbits; 1 1 scales between eye and nostril; posterodorsal loreals
slightly more than three times larger than interorbital scales; two postnasals;
auricular opening strongly denticulate, two or three auricular scales bearing
long spines, projecting posteriorly from anterior margin; eye diameter slightly
less than two times the snout length; ear diameter contained in eye diameter
three times, in snout length 5 times; second supralabial separated from nostril
by two loreal scales and postnasal; anterio-dorsal border of first supralabial
forms projecting edge bordering nostril below; 6 supralabial and 5 infralabial
scales to center of eye; mental slightly longer than wide, bordered posteriorly
by two postmentals which contact only the first labial on one side and none
on the opposite side; postmentals followed by irregular transverse row of
6 scales, followed by second row of 10 smaller scales;
Dorsum with 12 longitudinal rows of enlarged, keeled tubercles, 8 reach
to rear of head, 6 reach base of tail; 28 scales across venter; 56 from gular
region to vent; venter scales abruptly differentiated from lateral and dorsal
scales, about 5 to 6 times larger; 6 rows of enlarged tubercles on anterior six
whorls of tail, separated longitudinally by two annular rows of flat scales;
distal half of tail with rapid reduction of tubercular rows, 6-4-2-0; ventral
surface of tail with longitudinal row of 43 widened scales, about twice to three
times larger than adjoining scales; three postanal tubercles on either side of
vent; dorsal surface of femur and tibia with large trihedral tubercles inter-
mixed with granular scales; posterio-ventral surface of femur granular; upper
arm with large, flat scales; forearm with enlarged tubercles intermixed with
smaller flattened scales; terminal pads slightly longer than wide; claw not
extending beyond pad tips nor visible from below; lamellae formula for hand
7-8-9-11-8, foot 7-9-10-12-11.
Measurements in mm: Snout-vent length 44.0; head width 9.0; head
length 13.1; head depth 6.4; axilla-groin length 21.2; tail length 47.0; leg
18.6; arm 12.5; snout length 5.0; eye diameter 2.7; ear diameter 1.2; distance
from eye to ear 4. 1 .
Color in alcohol : Narrow broken, diagonal and transverse black lines
across dorsum, 13 complete bands on tail; dark brownish stripe from nostril
through eye to arm insertion; each enlarged dorsal tubercle flecked with black;
venter immaculate, pale white; labials stippled with brown; limbs faintly
spotted with brown.
Variation. Snout-vent length of subadult and adult males ranges from
30 to 45 mm (38.0), females from 30 to 46 mm (38.6), juveniles from 18
to 29 mm (25.4), males and females together average 38.3 mm; postmentals
vary from two to four, two occurring 104 times, three 5 times, and 4 one
time; postmentals contact the first labial only in 89.8 per cent of the sample,
first and second labial in 1 1.2 per cent; number of scales immediately fol-
lowing postmentals across the chin vary from 5 to 8 (6.7) ; scales across the
midorbital region range from 13 to 19 ( 15.8), across the snout at the level
1970
Lizards of the Genus Phyllodactylus
41
of the third labial 16 to 20 (17.3); number of scales bordering the internasals
vary from 4 to 8 (5.8) with an azygous scale separating the internasals in 14
of 1 10 specimens; transverse rows of scales across the venter number 22 to 30
(25.0), longitudinally 47 to 56 (50.6); dorsal tubercular rows number 12 to
14 (12.1); paravertebral tubercles from head to tail number 31 to 36 (34.0),
from axilla to groin 16 to 21 ( 18.0); scales on a line between nostril and eye
number 10 to 13 (11.5); lamellae beneath fourth toe number 11 to 13 (11.5);
body markings (if present) number 5 to 8 (7.1); tail bands (if present)
8 to 11 (9.9).
Tubercles on the dorsum, tibia, and thigh usually much enlarged, strongly
keeled and trihedral; tail tubercles large, somewhat flattened, keeled and ele-
vated posteriorly.
Color pattern consists of three distinct types; (1) presence of distinct
black spots, lines or blotches, (2) distinct but very small black dots, (3) uni-
form coloration; ground color generally pale brown in all specimens; in those
with dorsal markings, the head is usually spotted with profusion of dark brown
on ground color; dark brown to black stripe present from nostril through eye
to above arm or slightly beyond; tail generally banded with dark brown to
black on proximal three-fourths of tail, black and white on distal one-fourth;
those with uniform pale brown dorsal color occasionally have dark lateral
head stripe and faint tail bands; venter of all specimens pale white to white;
chin, throat, and chest with suffusion of light brown flecks on each scale;
ventral surface of tail usually heavily suffused with dark brown to black flecks,
much darker than venter color.
Comment. This species is named in honor of Carl Koford, who has
added much to our knowledge of the mammalian and reptilian fauna of Peru
in the past twenty years.
Habitat and natural history. This small gecko is widespread and abundant
in northwestern Peru. It has been found in every type of plant community
from sea level to 650 m, except for the main dune areas of the Sechura Desert.
This species is primarily a ground forager although it has been taken occa-
sionally on boulders and plants. The daytime retreats of this lizard are usually
holes beneath rocks, cracks in the ground, holes in sand banks, leaf litter at
the base of plants, under or in hollow limbs of cacti, under bark of shrubs
or stumps of rotting trees, in beach debris or trash dumped along the roadside.
Occasionally a few individuals were found beneath exfoliating flakes of granite.
Eggs of this species were found under rotting cacti (mainly Cereus) in
the Piura and Tumbes areas. One dead Cereus sp. plant contained 15 eggs of
P. kofordi and 4 eggs of reissi. Eggs were not found in other types of cactus
litter, nor in debris of acacia and mesquite. It appears that dead Cereus sp.
affords the necessary protection for the survival of the eggs and this plant is
sought out by females for egg deposition. This species lays one egg per clutch.
Captive females laid a single egg and preserved females contained only one egg.
Gravid females and small juveniles were found in July, August, Novem-
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No. 192
ber, and December, indicating at least summer and winter breeding periods.
Diurnal body temperatures were 8°C above the lizard’s nocturnal tempera-
tures during foraging periods.
This species is sympatric with P. reissi throughout most of its range, with
reissi and inaequalis along the desert foothills, and with reissi, clinatus, and
microphyllus in the Cerro Illescas area near the coast. Somewhere between
Los Organos and Cancas (on the northwest coastline) the northern distribu-
tion of microphyllus terminates, and the habitat (beach dunes community)
left vacant is effectively taken over by kofordi from Cancas to the Ecuadorian
border. Both were found in the same general microhabitat near Talara (beach
dunes) but microphyllus was much more abundant than kofordi.
Distribution. Phyllodactylus kofordi is a Peruvian species, found on the
periphery of the Sechura Desert, western foothills of the Andes, and along
the coast from Eten north to the Ecuadorian border. It probably occurs in the
southwestern part of Lojas and the southern part of El Oro Departments,
Ecuador (fig. 3).
Specimens (167) have been examined from the following localities:
Lambayeque: Chiclayo MVZ 82221. Eten BMNH 1900.2.26.1. 5 km SSW
Motupe MVZ 82209. 7 km S Motupe MVZ 82249-50, LACM 48540, 48559-
66, TCWC 27912, 28107-14. 7.2 km W Naupe LACM 48543-46, TCWC
28117-20. 22 km E Olmos LACM 48541-42, TCWC 28115-16. Piura: Bayo-
var FMNH 9803. 1 km S Bayovar, Cerro Illescas MVZ 85268, 85290-91,
85293-304. 4 km SSE Bayovar MVZ 82203-05, 82230. 5 km SSE Bayovar
MVZ 85287-89. 12 km S, 8 km E Bayovar MVZ 82176. Cerro Amotape
MVZ 82193-94, 82210-16, 82233-37, 82239-40, 82242, 82281, 82284.
Chongollapi AMNH 28465. 8 km S, 8.4 km SSE Chulucanas LACM 48547,
TCWC 28121. Fondo MVZ 82274. 2 km NE Las Lomas MVZ 85259, 85278-
84. 3 km ENE Las Lomas LACM 48556-58, TCWC 28131-34. 5 km E Las
Lomas MVZ 82243-47, 82296. 30 mi S Lobitos BMNH 1932.9.5.1-2. 2 mi
S Lobitos BMNH 1926.3.24.5. Negritos FMNH 8362. Paita AMNH 28460,
28462-64. Punta Aguja MVZ 82192, 82226, 82266-69. Reventazon MVZ
82229. 40 km WNW Sullana LACM 48548, TCWC 28122. Talara FMNH
57455. 2 km N Talara TCWC 28123. Trie Trac MVZ 82206, 85252-58,
85260-67, 85269-77, 85285, 85292, 85305-06. Tumbes: 1.2 km S Cancas
LACM 48549-50, TCWC 28125-27. 1.2 km S, 1 km E Puerto Pizzaro LACM
48551-55, TCWC 28128-30. Quebrada Seca FMNH 9810, TCWC 28124.
Phyllodactylus ventralis O’Shaughnessy
Phyllodactylus ventralis O’Shaughnessy 1875. Ann. & Mag. Nat. Hist.,
(4)16:263.
Phyllodactylus mulleri Parker 1935. Ann. & Mag. Nat. Hist., ( 10)
15:483.
Phyllodactylus underwoodi Dixon 1962. Southwest. Nat., 7:218.
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Lizards of the Genus Phyllodactylus
43
75 70 65
R DIXON I 3
P. V ENTR ALIS 0 kilometers
Figure 6. Distribution of P. dixoni and P. ventralis in northeastern South America.
Holotype. Adult male, BMNH 53.6.1.33, collector unknown, date of
collection unknown. Type locality. Island of Jamaica (in error).
Definition. A large gecko with a maximum known snout-vent length of
75 mm; tibia and thigh tubercles present; ear denticulation usually present;
enlarged preanal scale absent; terminal lamallae of digits distinctly widened
and usually truncate at tips; median row of scales beneath tail distinctly wid-
ened; dorsal rows of enlarged tubercles close together, trihedral, and strongly
carinate; six to eight rows of tubercles across base of tail; rear of head with
rounded tubercles dispersed among granular scales (fig. 14a).
Diagnosis. Phyllodactylus ventralis is distinguished from heterurus, ger-
rhopygus, and angustidigitus by the absence of an enlarged preanal scale;
from inaequalis and microphyllus by the presence of regular rows of large
trihedral, strongly keeled dorsal tubercles; from clinatus, lepidopygus, and
interandinus by the presence of large tibia tubercles; from johnwrighti, pumi-
lus and reissi by the presence of large tubercles on forearm; from sentosus
by having terminal digital lamellae distinctly widened and enlarged; from
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No. 192
kofordi by greater maximum snout-vent length of 75 mm, rather than 46 mm;
and 19 or more paravertebral tubercles between axilla and groin; from dixoni
by having 24 or less scales across venter, rather than 26 or more, and dorsal
tubercles four or more times the size of those in dixoni.
Description. Rostral about one and one half times higher than wide; two
internasals, bordered posteriorly by small granular scales and postnasal of
each side; nostril surrounded by rostral, labial, two postnasals and internasal;
ventral edge of nostril scale in broad contact with first labial; deep depression
between internasals and in frontal region; scales of posterior loreal region
six to seven times larger than midorbital scales; eye large, contained in snout
length about two times; eye brille with one to two rows of granules and one
larger outer row of scales, posterior half are pointed; ear diameter contained
in eye diameter about three times; ear opening slightly denticulate anteriorly,
none posteriorly; rear of head with very large tubercles dispersed among
granular scales; supralabials usually six to seven and infralabials five to six
to a point below center of eye; mental slightly longer than wide, somewhat
triangular in shape.
Dorsum with regular longitudinal rows of very large, trihedral tubercles,
10 to 14 rows reach nape, 6 to 8 rows reach base of tail; paravertebral rows
of tubercles separated along median line by four to five rows of granules;
postanal tubercles usually number three to four and very distinct in adult
males.
Forearm with large trihedral tubercles dispersed among granular scales;
upper arm with large flat scales; dorsal surface of thigh and tibia with 10 to
20 very large tubercles dispersed among granular scales; terminal digital
lamellae large, either rounded or truncate; claw short, hidden when viewed
from below.
Variation. Snout-vent lengths of adult males from 50 to 74 mm (62.4),
adult females from 52 to 75 mm (62.3), subadults 40 to 45 (42.2), juveniles
24 and 29 mm; postmentals normally two, three occurring once in 22 speci-
mens; postmentals contact first labial only in 25 per cent; first and second
labials in 75 per cent of sample; scales bordering internasals vary from 4 to 6
(5.1); scales from nostril to eye vary from 9 to 12 ( 10.3); scales across snout
at level of third labials vary from 17 to 23 (19.4); midorbital scales 12 to 17
(14.4); dorsum with 14 to 19 (16.4) longitudinal rows of enlarged trihedral
tubercles; paravertebral tubercles from head to base of tail vary from 34 to
49 (41.2), axilla to groin 19 to 25 (22.1); venter scales from gular region
to vent vary from 46 to 54 (52.0), across venter from 18 to 24 (20.6); fourth
toe lamellae vary from 12 to 14 (13.6).
Color pattern varies from seven to 10 narrow, undulating, dark brown
cross bands on a light brown to tan ground color, to a linear series of 10 to
20 dark brown spots on a yellowish brown ground color; dorsal surface of
head generally brown with or without dark brown spots or reticulating lines;
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Lizards of the Genus Phyllodactylus
45
side of head usually with a dark brown line from nostril through eye to arm
insertion, bordered above and below by whitish lines or ground color; labials
usually spotted or mottled with brown and dirty white; venter dirty white
to yellowish brown.
Comments. In 1962 one of us (Dixon) described a new species, Phyllo-
dactyliis underwoodi, from the island of Grenada, Windward Islands. The
holotype is the only available specimen and is unique in several features of
scalation. Miss Alice G. C. Grandison (personal communication) informed us
that there was an error in the labeling of this specimen, and it should have
been labeled as Colombia. This specimen was a part of a relatively small
collection made by F. A. Simons, purchased from a Mr. Janson by the British
Museum of Natural History, and bore the locality New Grenada.
Dixon (1962) noted the similarity of P. underwoodi to ventralis, but it
differed in three major scale characters. It is now apparent that P. underwoodi
is likely an aberrant ventralis, having three postmentals, 14 rows of enlarged
dorsal tubercles, and six rows of tubercles across the base of the tail. No
specimens of ventralis from Colombia or Venezuela have this combination
of scale counts. However, only 22 specimens of P. ventralis have been
examined, and additional material may reveal a wider range of variation.
Habitat and natural history. This species has not been collected by either
of us, therefore we rely upon information from the literature and from per-
sonal observations of others. Ruthven (1922) records P. ventralis from
houses, buildings, under stones and bark of trees. Two specimens were found
in a dry thorn forest and one in a thorny scrub and cacti association under a
stone. Mechler (1968) found three specimens in debris of an abandoned
house in Bonda, Colombia. Janis A. Roze (personal communication) states
that P. ventralis occurs in dry, semixerophytic areas, but occasionally pene-
trates more moist habitats where cactus and thorny vegetation is replaced by
a rich variety of shrubs and trees. Roze ( 1964) states that P. ventralis is often
found in occupied houses, in abandoned buildings, and trees near buildings.
Roze indicated that competition between Thecadactylus rapicauda and P.
ventralis for human dwellings is being won by T. rapiciiada.
Distribution. This species is found in the dry thorn forests and savannahs
of northern Venezuela and Colombia (fig. 6).
Specimens (22) have been examined from the following localities:
COLOMBIA. Magdalena: Bolivar UMMZ 54737. “Grenada” BMNH
1880.2.26.6. Santa Marta UMMZ 48177. Santa Marta Mountains MCZ 24881,
UMMZ 45462-63. VENEZUELA. Anzodtegui: Barcelona MBUCV 3144.
A ragua: Maracay MCZ 39980. Carabobo: Caizuire MBUCV 3151. Isla de
Burro MBUCV 3143. Falcon: Coro MBUCV 3148. Gudrico: San Juan de los
Morros MBUCV 3149. Sosa MBUCV 3145. Portuguesa: Acarigua MCZ
(unnumbered). Aparicion MBUCV 3146. Sucre: Isla Patos BMNH 1935.10.1.
10-11, 1940.2.15.1, 1945.11.1.41-42. Puerto de Hierro MCZ 50744-45.
46
Contributions in Science
No. 192
Phyllodactylus dixoni Rivero-Bianco and Lancini
Phyllodactylus dixoni Rivero-Bianco and Lancini 1968. Mem. Soc. Cienc.
Nat. La Salle, 27:168.
Holotype. Adult female, MBUCV III-8.450, collected by Juhani Ojasti
in a cave on 5 October 1964. Type locality, Desembocadura del Rio Parguaza,
afluente del Orinoco Medio en el Estado Bolivar, Venezuela.
Definition. A large gecko with a maximum known snout-vent length of
76 mm; tibia, thigh, and forearm tubercles present; ear with little or no den-
ticulation; enlarged preanal scale absent; terminal lamellae of digits distinctly
widened and truncate; median row of scales beneath tail distinctly widened;
dorsal rows of enlarged tubercles somewhat small, low and slightly trihedral;
each dorsal row of tubercles separated from the other by three to five rows
of granules; six to eight rows of dorsal tubercles across base of tail; rear of
head with rounded tubercles dispersed among granular scales, almost subequal
in size (fig. 14b).
Diagnosis. Phyllodactylus dixoni may be distinguished from all other
mainland species except ventralis, by having no enlarged preanal scale; regu-
lar dorsal rows of keeled tubercles; presence of tubercles on forearm, thigh
and tibia; distinctly enlarged and widened terminal digital lamellae; large
maximum snout-vent length of 76 mm; 23 or more paravertebral tubercles
between axilla and groin; from ventralis by having dorsal tubercles about one
fourth the size of those of ventralis; 26 or more scales across the venter rather
than 24 or less.
Description. Rostral twice as wide as high, its dorsal edge with median
vertical groove about one half depth of rostral; two internasals, bordered
posteriorly by small granules and postnasal of each side; nostril surrounded
by rostral, labial, internasal and two postnasals; first labial in broad contact
with ventral edge of nostril scale; shallow depression between internasals,
moderate depression in frontal region; scales in posterior area of loreal region
about three to four times larger than scales in midorbital region; eye large, its
diameter contained in snout length about 1.5 times; eye brille with two rows
of granules and one larger outer row of scales, last five to eight pointed; ear
diameter contained in eye diameter about two times; ear opening without
denticulation, scales on posterior and anterior borders rounded; rear of head
with granular scales with slightly larger rounded and elevated scales inter-
mixed; supralabials six to seven and infralabials five to six to a point below
center of eye; mental bell shaped, about as wide as long; two postmentals,
slightly longer than wide and in contact on median line; postmentals contact
only first labial.
Dorsum with regular rows of enlarged, keeled tubercles that are some-
what oval in outline; paravertebral rows separated from each other by four
to five rows of granules; 10 rows of dorsal tubercles reach nape, 6 to 8 reach
1970
Lizards of the Genus Phyllodactylus
47
base of tail; postanal tubercles number three to four on each side in males;
venter scales abruptly differentiated from lateral body granules.
Dorsal surface of upper arm with flattened scales; posterodorsal surface
of forearm with tubercles interspersed among small scales; dorsal surface of
thigh with 15 to 25 tubercles dispersed among smaller flattened scales; tibia
with large tubercles dispersed among granular scales; terminal lamellae large,
somewhat truncate, about twice as long as wide; claw long, visible when
viewed from below.
Variation. The known snout- vent lengths of adults range from 56 to
76 mm (67.8), subadults 39 and 46 mm, juvenile lengths unknown; scales
bordering postmentals vary from 5 to 8 (6.1); scales from nostril to eye vary
from 11 to 12 (11.3); scales across snout at level of third labials vary from
17 to 20 (18.6), midorbital scales 15 to 19 (17.5); scales bordering inter-
nasals vary from 5 to 6 (5.1); longitudinal rows of enlarged dorsal tubercles
vary from 13 to 16 (14.9); number of paravertebral tubercles from head to
base of tail vary from 41 to 50 (46.7), axilla to groin 23 to 26 (24.7); venter
scales from gular region to vent vary from 59 to 66 (61.7), across venter
26 to 28 (27.1); fourth toe lamellae vary from 11 to 13 (11.3).
Dorsal color pattern usually consists of five to eight undulating, dark
brown cross bands on a brownish gray ground color from occiput to base of
tail; tail banded with dark brown and light gray in juveniles, sometimes irregu-
lar bands or spots in adults; head usually brownish gray with a suffusion of
dark brown spots; labials spotted with dark brown; venter usually brownish
cream; lower sides of body brownish yellow; ventral surfaces of toes dark
brown.
Habitat and natural history. Dr. Janis Roze (personal communication)
states that the type series of P. dixoni was taken from beneath flakes of black
basaltic rocks. The flakes were associated with immense, grayish black rock
bluffs some 500 meters from the mouth of the Rio Parguaza. The immediate
vicinity was essentially void of vegetation except for a few scattered shrubs
and bromeliads. During the day, the temperature of the rocks reached 60°+C
and the lizards took refuge beneath flakes or within cracks of boulders.
The majority of lizards were taken by breaking off the rock flakes or by
beating upon the flakes until the lizards ran out. Some lizards were observed
running about in the late afternoon in the shade of scant vegetation or in the
shade of rock boulders (Roze, personal communication) .
Distribution. Phyllodactylus dixoni is known only from the type locality.
The type locality is approximately 35 km NE of Puerto Carreno, Colombia,
and 385 km (airline) south of San Juan de los Morros, Venezuela, the nearest
locality of P. ventralis (fig. 6) .
Seven specimens have been examined from the following locality:
VENEZUELA. Bolivar: mouth of Rio Parguaza, on the Rio Orinoco
MCNC 5479-80, MBUCV III 8451-54, TCWC 23827.
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Contributions in Science
No. 192
Phyllodactylus sentosus sp. nov.
Phyllodactylus phacophorus, (part) Boulenger 1885, Cat. Lizards in the
British Museum (Natural History) 2:84.
Holotype. Adult female, TCWC 27913. Collected by Fortunato Blancas,
November 1949; Type locality, Lima, Department of Lima, Peru.
Definition. An average-sized gecko with maximum snout-vent length of
56 mm; large preanal plate absent; very large trihedral tubercles present on
dorsum, forearm, thigh, tibia, and tail; ear denticulate on anterior and pos-
terior margins; median row of scales beneath tail with few scales slightly
enlarged, rounded; tail 45 to 50 per cent of total length; terminal lamellae of
digits separated along median line, small, claw visible between and beyond
tip of terminal lamellae when viewed from below (fig. 12a).
Diagnosis. This species may be distinguished from all other western
South American species except P. kofordi, by the presence of very large
tubercles on dorsum, thigh, tibia, forearm, and tail. Phyllodactylus sentosus
differs from kofordi by having very small terminal lamellae on the digits and
generally uniform sized scales beneath ventral surface of tail, rather than
terminal lamellae large and all scales of median row beneath tail enlarged,
rectangular.
Description of holotype. Rostral completely divided by vertical groove,
twice as wide as high, its dorsal and lateral edges straight; two internasals, some-
what rounded, their median edges in narrow contact, bordered posteriorly by
four granules and supranasal of each side; nostril surrounded by rostral, labial,
supranasal, and one postnasal; first supralabial in narrow contact with ventral
edge of nostril; no depression between internasals, slight depression in frontal
region; 9 scales between eye and nostril, scales in posterior loreal region about
three to four times larger than scales in midorbital region; 18 scales across
snout at level of third labials, 12 scales across head between anterior edge of
eyes, 14 midorbital scales; eye large, its diameter contained in snout length
1.7 times; eyelid with one to two rows of granules and one larger outer row
of scales, last four to five are pointed; ear diameter contained in eye diameter
2.3 times; ear opening denticulate, scales on anterior and posterior margins
pointed; rear of head granular with many larger intermixed tubercles; six
supralabials and five infralabials to a point below center of eye; mental bell-
shaped, slightly longer than wide, bordered posteriorly by three postmentals;
postmentals rounded, slightly longer than wide, their median edges in broad
contact, followed by transverse row of eight small scales, followed by second
row of 1 1 smaller scales; postmentals contact first labial.
Dorsum with 16 longitudinal rows of enlarged, trihedral tubercles,
paravertebral row with 26 tubercles from rear of head to base of tail, 13
between axilla and groin; paravertebral rows separated from each other by
three rows of granules; 10 rows of tubercles reach to rear of head, 8 to base
of tail; each tubercle of enlarged dorsal series separated from preceding
tubercle by one to two granules; postanal tubercles number three on each side;
1970
Lizards of the Genus Phyllodactylus
49
venter with 18 longitudinal and 48 transverse rows of scales; tail with 8 rows
of tubercles in proximal eight whorls, distal one-third of tail regenerated.
Dorsal surface of upper arm with flattened scales, forearm with larger
tubercles interspersed among smaller flattened scales; dorsal surface of thigh
with 8 to 9 tubercles interspersed among granular scales, tibia with 13 to 15
tubercles; lamellae formula for hand 6-8-12-11-8, foot 7-10-11-12-10; claw
largely visible between and beyond tip of terminal lamellae when viewed
from below; terminal lamellae small, about three times as long as wide.
Measurements in mm: Snout-vent length 50.0; axilla-groin length 25.0;
leg length 19.3; arm length 16.0; tail length 46.3; head length 9.6; head height
7.8; head width 9.6; snout length 5.1; distance from eye to ear 4.6; eye diam-
eter 3.0; ear diameter 1.3.
Color in alcohol: ground color fawn; dorsum with nine undulating cross-
bands of reddish brown, first two crossbands complete, remainder broken
along midline of dorsum by cream middorsal stripe that begins about level of
shoulder; each dorsal tubercle generally flecked with dark brown; limbs with
brownish spots, forming faint reticulations on ground color; tail with six
brown crossbands, about one-half width of ground color interspaces; distal
one-third of tail regenerated; head brownish with wash of light brown; dark
brown stripe from nostril to posterior edge of ear, bordered above by cream
line on snout, above and below by cream line between eye and ear; ventral
surfaces pale white, scales below tail with flecks of brown.
Variation. Snout-vent length of two males 49 and 56 mm (52.5); two
females 39 and 50 mm (44.5); postmentals number two in three specimens,
three in one; postmentals contact first labial only in all; number of scales
across chin immediately following postmentals varies from 6 to 8 (7.3);
number of scales bordering posterior edge of internasals vary from 5 to 7
(6.3), with internasals completely separated by an azygous scale in two of
four specimens; scales across snout at level of third labial varies from 16 to
18 (16.5) midorbital scales vary from 13 to 14 (13.3); scales across venter
vary from 18 to 21 ( 19.3) ; longitudinally 47 to 55 (50.3) ; number of enlarged
rows of dorsal tubercles vary from 14 to 16 (15.0); number of paravertebral
tubercles from rear of head to base of tail vary from 26 to 31 (28.8); from
axilla to groin 13 to 16 ( 15.0) ; scales along line from nostril to eye vary from
9 to 10 (9.3); number of fourth toe lamellae vary from 12 to 13 (12.5);
terminal lamellae of all digits very small, twice as long as wide; claw visible
beyond tip of pad when viewed from below; number of enlarged rows of
tubercles at base of tail number 8 in three specimens, 6 in one; 8 tubercles
occur in each of 8 to 1 1 whorls on proximal two-thirds of tail, reducing rapidly
in sequence to 6-4-2-0 on distal third of tail; median row of scales beneath
tail generally uniform in size to other scales but occasionally a larger, rounded
scale may be present.
Number of dorsal body bands varies from 8 to 10 (9.0), with majority
of bands broken along midline of body in three specimens, bands loosely con-
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No. 192
nect and form reticulated pattern in one; tail bands number 13 and 14 in two
specimens with complete tails; venter pale white to dirty cream; dark lateral
face stripe continues to shoulder region as dashed line in two specimens,
reaches to ear in two.
Comments. Phyllodactylus sentosus closely resembles kofordi (NW
Peru), darwini (Galapagos Islands), wirshingi (Puerto Rico), and martini
(Dutch Leeward Islands) in general appearance. The latter four species have
enlarged trihedral tubercles on the dorsum, tail, thigh, tibia, and forearm.
However, the four species have the median row of scales beneath the tail
much enlarged and widened, and have moderately large, well defined ter-
minal lamellae on all digits. In addition, martini and wirshingi have higher
numbers of enlarged rows of dorsal tubercles ( 18 to 20) and higher num-
bers of paravertebral tubercles from head to base of tail (32 to 40). The num-
ber of enlarged dorsal rows of tubercles in darwini are identical with those of
sentosus, but darwini has higher numbers of paravertebral tubercles from
head to base of tail (33 to 38) and differs in several other features of squa-
mation.
Natural history. The only known habitat of this species is the grounds
surrounding the University of San Marcos in Lima. Dr. Gambini ( personal
communication) indicated that she found one specimen running across a side-
walk on the campus just after dark.
Distribution. Known from the University of San Marcos campus, Lima,
Peru, and from the vicinity of the city (fig. 5). Six specimens have been
examined: Lima: Lima BMNH 63.2.3.18-19, TCWC 27913, 28329-31.
Phyllodactylus reissi Peters
Phyllodactylus reissi Peters, W. 1862. Monatsb. Konigl. Akad. Wiss.
Berlin, Nov., 1862, pp. 626-27.
Phyllodactylus baessleri Werner 1901. Abh. u. Ber. d.k. Zool. u. Anthr.-
Ethn. Mus. zu. Dresden, 9(2) : 2.
Phyllodactylus guayaquilensis Werner 1910. Mitt. Naturh. Mus. Ham-
burg, 27(2): 4.
Phyllodactylus abrupteseriatus Werner 1912. Mitt. Naturh. Mus. Ham-
burg, 30:4.
Phyllodactylus magister Noble 1924. Occ. Pap. Boston Soc. Nat. Hist.,
5:110.
Syntypes. Berlin Mus. 3734 (4) , 4567 (2) ; BM 4567, ad. female, snout-
vent 64 mm, herein selected as the Lectotype; Carl Reiss, collector; date of
collection unknown. Type locality: Guayaquil, Ecuador.
Definition. A large gecko, maximum snout-vent length of 75 mm; femur
tubercles absent, tibia tubercles present; ear strongly denticulate on anterior
and posterior margins; enlarged tubercles on rear of head present; median
row of scales beneath tail distinctly widened; enlarged preanal plate absent;
12 to 18 rows of enlarged trihedral tubercles on dorsum; tail about 48 to 58
1970
Lizards of the Genus Phyllodactylus
51
per cent of total length, without whorls of enlarged tubercles; terminal lamellae
of digits moderately enlarged, truncate; dorsal color pattern variable, uniform
in color to bold bands; venter usually yellowish (fig. 1 3b) .
Diagnosis. This species may be distinguished from P. heterurus, angusti-
digitus, and gerrhopygiis by the absence of enlarged preanal plate; from
lepidopygus, microphyllus, clinatus, and inaequalis by the presence of tibia
tubercles and larger size; from interandinus by its larger size (average s-v
of 58.3 mm rather than 40.1 mm), fewer number of head to tail tubercles
in paravertebral row, 47 to 60 rather than 64 to 8 1 ; from kofordi by its larger
size (as above) rather than average of 44.0 mm, and by having more head to
tail tubercles in paravertebral row (as above) rather than 3 1 to 36; from
johnwrighti and pumilus by larger size, a maximum snout-vent length of
75 mm rather than 5 1 mm; thigh tubercles absent, rather than present; from
sentosus by absence of tail and forearm tubercles, larger scales in median
row beneath tail, larger terminal lamellae, and color pattern.
Description. Rostral about twice as wide as high; internasals rounded,
their median edges in broad contact, bordered posteriorly by small granules
and postnasal of each side; nostril surrounded by rostral, labial, internasal,
and two postnasals; first labial in broad contact with ventral edge of nostril;
shallow depression between internasals, deep depression in frontal region;
scales in posterior loreal region about 3 to 4 times larger than midorbital
scales; eye large, its diameter contained in snout length about 1.8 times; eyelid
with 2 rows of granules and 1 larger outer row of scales, last 3 to 6 of which
are pointed; diameter of ear contained in eye diameter about 2.1 times; ear
opening strongly denticulate with pointed scales on the anterior and posterior
margins; rear of head granular with larger intermixed tubercles; 6 to 7 supra-
labials and 5 to 6 infralabials to a point below center of eye; mental bell
shaped, as wide as long, bordered posteriorly by two to four postmentals;
postmentals slightly longer than wide, their median edges in broad contact.
Dorsum with longitudinal rows of enlarged trihedral tubercles that are
somewhat elevated; paravertebral rows separated from each other by 4 to
5 rows of granules; 6 rows of tubercles reach to nape, 6 to base of tail; each
tubercle of enlarged dorsal series separated from preceding tubercle by 0 to
2 granules; postanal tubercles number 4 on each side, distinct and slightly
elevated posteriorly in males; dorsal surface of upperarm with flattened scales,
forearm with slightly larger flattened scales; dorsal surface of thigh without
tubercles intermixed among smaller flattened scales; tibia with large tubercles
scattered among granular scales; claw slightly visible when viewed from
below; terminal pad large, slightly longer than wide, somewhat truncate at tip.
Variation. Snout-vent length of subadult and adult males range from
42 to 75 mm (59.4), females from 37 to 73 mm (57.5), juveniles from 22 to
35 mm (28.9), and males plus females average 58.3 mm; 48 per cent of total
sample (753 specimens) were 60 mm or more in snout-vent length; post-
mentals vary from two to four, two occurring 97.4 per cent of the time, three
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No. 192
1.9 per cent and four 0.7 per cent; postmentals contact the first labial only
in 96.8 per cent of sample, first and second labials 3.2 per cent; number of
scales immediately following postmentals across chin vary from 5 to 8 (6.3);
scales across midorbital region vary from 16 to 24 (20.2) ; scales across snout
at level of third labial vary from 18 to 27 (23.7); scales bordering internasals
posteriorly vary from 5 to 9 (6.7) with an azygous scale separating the inter-
nasals in 23.1 per cent of sample; transverse rows of scales across venter vary
from 24 to 37 (29.7), longitudinally 54 to 77 (60.0); dorsal tubercular rows
vary from 12 to 18 (13.2); paravertebral tubercles from head to tail vary
from 47 to 60 (53.0); between axilla and groin 25 to 38 (31.3); scales along
a line between nostril and eye vary from 10 to 16 (12.5); lamellae beneath
fourth toe vary from 11 to 17 (13.5).
Dorsal color pattern highly variable, ranging from uniform yellowish
tan without darker markings to light gray ground color with bold black undu-
lating crossbands half the width of ground color interspaces; those with dark
dorsal markings on ground color may have spots, crossbands, blotches, or
lines that are either bold black, dark brown, brown, to indistinct grayish brown;
in a few specimens the lines coalesce, forming a variegated pattern; when body
markings are distinct enough to be counted, they generally vary from 5 to 7
pairs or single bands.
Coloration of dorsal surface of tail somewhat similar to that of dorsum;
when bands are present they vary from 8 to 11 with 9 and 10 being most
common; interspaces usually lighter than those of dorsum, tending to be
more in contrast with darker bands; dorsal surface of head is either uniform
in color, spotted or reticulated with dark brown to black lines or spots on
ground color; side of head with or without distinct dark line from nostril
through eye to arm; limbs similar to dorsum in color and pattern, with fore-
arm and lower leg generally banded with dark on ground color in most
specimens.
Ventral surfaces generally grayish white in alcohol (uniform pale yellow
to tan with yellowish cast in life), with dark color of sides encroaching on
venter in large individuals; ventral surface of tail usually darker than venter.
An examination of three populations of P. reissi from the coastal and
coastal foothills of Ecuador, desert foothills surrounding the Sechura Desert
of Peru, and the Maranon-Chinchipe Valley of northern Peru reveal very
little variation in the majority of characters. The Ecuadoran population has
an azygous scale separating the internasals in 43.1 per cent of the sample,
32.3 per cent in the Sechura Desert sample, and 14.3 per cent in the Maranon-
Chinchipe sample. The number of enlarged dorsal tubercular rows vary from
12 to 16 (14 the common number), 12 to 16 (12 common), and 14 to 18
(16 common) respectively, in the three samples indicated above.
Comments. The types of P. baessleri, abrupteseriatus, and quayaquilensis
were destroyed during World War II, but the characters mentioned in the
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Lizards of the Genus Phyllodactylus
53
original descriptions of the latter two species fall within the variation present
in reissi. An analysis of the characters of a large sample of magister from the
Marafion and Chinchipe Valleys of Peru indicates only minor differences
between it and populations of reissi from Ecuador and northern Peru. We,
therefore, consider abrupteseriotus, guayaquilensis, and magister as subjective
synonyms of reissi.
Werner’s (1901) original description of P. baessleri is the only evidence
we have for comparison of his species with those known to occur in Peru.
His description does not correspond in its entirety to any population of Phyl-
lodactylus we currently recognize in Peru. A translation of Werner’s (1901,
p. 2) original description follows:
“Closely related to P. tuberculosus, snout one and a half times as long
as eye, somewhat longer than distance between eye and ear; frontal region
with slight concave; ear opening diagonal, small diameter of ear one half
that of eye; snout with large flattened, rounded scales, those of the rear of the
head smaller, here they seem to differ by being mixed with small rounded
tubercles; ear opening with three pointed scales on the anterior margin; rostral
four-sided, the posterior margin with a short median notch; nostril surrounded
by first labial and three nasals, of which the upper one contacts the upper
nasal of the other side; 7-8 upper labials, 6th below eye; 7 infralabials; mental
large, bell-shaped, followed by two small polygonal postmentals in contact
with each other; followed by successfully smaller scales in size; upper side of
rump with 12 rows of strongly keeled tubercles, tubercles of each row close
together; the two outer rows near the venter do not reach the forelimb; venter
scales flat, overlapping, cycloid-hexagonal; tibia with enlarged keeled tubercles
on the upper side; finger and toe ending with strongly enlarged lamellae,
similar to P. tuberculosus; 4th toe with 16 lamellae, of which the distal one
and mostly proximal ones are divided.
Color above, light grayish brown with black spots on head, tail, base of
tail and legs; four long, black, parallel lines, from rear of eye to base of fore-
leg, two from back of head to middle of back, between the limbs; underside
whitish.
Total length 91 mm, most of tail regenerated (39 mm); The single
example on hand (D 1780 M. Dr.) came from Chanchamayo. The species is
named in honor of Prof. A. Baessler.”
From Werner’s description (1901), we may eliminate the Peruvian
species without strongly keeled tubercles on the dorsum, i.e. P. microphyllus,
inaequalis, gerrhopygus, angustidigitus, and heterurus. His description men-
tions the presence of tibial tubercles, therefore we may eliminate clinatus,
interandinus, and lepidopygus. The remaining species, reissi, johnwrighti,
sentosus, and kofordi agree in part with his original description.
The latter four species have denticulate scales on the anterior and poste-
rior margins of the ear, tibia tubercles, and strongly keeled dorsal tubercles.
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Of these four species, only P. reissi has a fourth toe lamellae count ( 12 to 17)
that falls within the range given by Werner for baessleri. Occasionally, P.
interandinus has a few small tubercles on the tibia.
The color description given by Werner ( 1901 ) does not agree with any
species of Phyllodactylus in Peru, while all other features mentioned in his
original description agree with most of the species.
Werner’s type locality, Chanchamayo, is also questionable. Chanchamayo
is a well known, tropical lowland locality at the headwaters of the Rio Perene
on the east side of the Andes. There are no known species of Phyllodactylus
inhabitating the selva of eastern Peru.
We provisionally place P. baessleri as a subjective synonym of reissi,
based upon the destruction of the holotype, a possible error in the type locality,
and discrepancies in the original description when compared with known
species in Peru.
Habitat and natural history. Phyllodactylus reissi is the most abundant
scansorial gecko of northern Peru. Daytime retreats consist of any object that
affords protection from predators and adverse temperatures. Individuals were
found beneath bark of living, dead, standing, or prone trees, fence posts,
fallen banana plants, standing and prone cacti, cracks in adobe walls, crevices
in boulders, knot holes in trees, beneath bridges, rotting tree stumps, beneath
palm leaves, construction materials and fruit crates. Their nocturnal activities
were essentially on vertical surfaces, i.e., trunks of trees, walls of buildings,
boulders, cacti, and fence posts, but occasionally were found actively foraging
on the ground. Several were found foraging on open ground two to three
meters from large boulders and in leaf litter in cacao groves.
This species is always found in association with some type of plant cover.
When exposed to daylight, P. reissi actively seeks a dark place to hide.
When exposed in its daytime retreat, it hesitates only a few seconds before
rapidly moving to another dark retreat. Its movements are usually vertical,
but on horizontal limbs or tree trunks reissi usually runs to the ventral surfaces.
Two eggs are laid and usually deposited in rotting debris at the base of
trees, beneath leaf litter in forests, in rotting agave stumps, hollow branches
of dead cacti filled with debris, or in any rotting debris associated with its
general habitat.
Phyllodactylus reissi has been found occupying the same cactus plant
with interandinus in the upper Amazon Basin, and with kofordi in north-
western Peru. It has been found on the same tree with Pseudogonatodes bar-
bouri and Gonatodes atricucullaris, and in the same general habitat with
inaequalis, clinatus, and microphyllus.
As in all of the species of Phyllodactylus found in Peru, the nocturnal
body temperatures of reissi closely approximated that of the substratum upon
which it was active.
Distribution. This species is found from sea level to about 2,000 m in
southern Ecuador and northwestern Peru. Its principal habitat seems to be
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Lizards of the Genus Phyllodactylus
55
arid tropical scrub west of the Andes, and cacti dominated areas of the north-
ern interandian basins of Peru (fig. 5) .
Specimens (772) have been examined from the following localities:
ECUADOR, 4 km E San Francisco USNM 167183 (23). El Oro:
Machacha USNM 167182. Santa Rosa AMNH 22068. Guayas: Ancon Camp
BMNH 1931.10.21.1-2. 6 km SW Colonche USNM 167189. Daule River
MCZ 4740 (2). Guayaquil AMNH 21845. ANSP 7526. BM 3734 (4), 4567
(2). CMF 16504, 16540-46, 16658-61, 16676, 16695, 16700, 16850-51,
16889-900. EPN 4618 (18). FMNH 61210. HM 3386. Playas MVZ 77217-
20, 77268. USNM 167181. Poza Huancauilca CMF 16512, 16515. Puente
Chimbo AMNH 24343. Puna CAS 84770. 1 km NW Punta Cornero USNM
167190 (6). 0.5 km N Punta Cornero USNM 167191 (10). Salinas CMF
16531. Loja: BMNH 1930.10.12.13. Rio Casanga AMNH 18304-05. Manahi:
Manta CMF 16522; FMNH 53896-98. 1 km S Manta USNM 167186 (5).
6 km E Manta, 5 km W Montecristi USNM 167188 (8) . No specific locality
in Ecuador AMNH 64569 (4); ANSP 7530; USNM 14052-53.
PERU. A mazonas: 14 km S Bagua Chica TCWC 28259. 17 km S Bagua
Chica LACM 48766-72, TCWC 28260-64. 19 km S Bagua Chica LACM
48739-58, 48786; TCWC 28228-49. 10 km WSW Bagua Grande MVZ 82301.
8 km WSW Bagua Grande MVZ 82208, 82302-06. 15.5 km WSW Bagua
Grande LACM 48759-64. TCWC 28250-55. 3.5 km SE Bagua Grande LACM
48765. TCWC 28256-58. Cajamarca: Bellavista AMNH 28525-27, 39339-40.
FMNH 73379. LACM 48734-38. MCZ 42233 (3), 18126-29, 18141-42 (+ 208
unnumbered). TCWC 24923, 28210-21. UIMNH 41515, 40747. UMMZ
59029 (4). 4.8 km S Bellavista LACM 48734-38. TCWC 28222-27. Perico
FMNH 100182-83. MCZ 17974, 18145-50. Lamhayeque: 1 1 km N Jayanca
MVZ 82197-201. 5 km SSW Motupe MVZ 82299. 7 km S Motupe LACM
48707-21, 48811-29. MVZ 82300; TCWC 28175-208. 15 km S Naupe MVZ
82202. 7.2 km W Naupe LACM 48773-74. TCWC 28265. 22 km E Olmos
TCWC 28209. Liber tad: Chiclin FMNH 34266. Pacasmayo HM (3 unnum-
bered); SMFM 8285, 8289. USNM 167180. Piura: Bayovar FMNH 9798-
802, 9804. 1 km S Bayovar (Cerro Illescas) MVZ 85308-09, 85316-28.
1.1 km E Bocana LACM 48810. TCWC 28278-79. 1 km N, 28 km E Cabo
Blanco MVZ 82177, 82232, 82280. Capullana Point FMNH 41551. Catalinas
FMNH 9816-17. Cerro Amotape MVZ 82169, 82173, 82195, 82217-18,
82238, 82241, 82282, 82285-95. 8 km S, 8.4 km ESE Chulucanas LACM
48775-81. TCWC 28266-71. 2.4 km SW Chulucanas LACM 48782, TCWC
28272-73. El Alto FMNH 41548, 41550. Fondo MVZ 82272-73. La Breita
BMNH 1926.3.24.6. 5 km E Las Lomas MVZ 82219-20, 82297-98. 2 km NE
Las Lomas MVZ 85310-15. 3 km ENE Las Lomas LACM 48802-09. TCWC
28297-304. 5 km E, 4.2 km NNE Lobitos LACM 48798-801. TCWC 28291-
96. Monte Grande MVZ 82279. Negritos FMNH 8364-65, 8384. Parinas
MVZ 82207, 82276-77. Punta Sal SMFM 41913. Sechura Desert CAS 92346.
40 km WNW Sullana LACM 48783-84; TCWC 28274-75. 4 mi W Suyo
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TCWC 24072-91. Talara AMNH 66600-01; FMNH 41547, 41549, 53895,
53892-93, 57456. 1 km SW Talara MVZ 82275. 2 km E Talara LACM 48785;
TCWC 28276-77. Tamboa AMNH 28426-27. Trie Trac MVZ 85307. between
Verdun Alto and Negritos FMNH 41546. Tumbes: Las Vocas USNM 38569.
1.2 km S, 1 km E Puerto Pizzaro LACM 48792-97; TCWC 28285-90. Que-
brada Seca FMNH 9809, 9811-12; LACM 48787-91; TCWC 28280-84. no
specific locality in Peru ANSP 13121; FMNH 41552-53; SMFM 8290.
LEP1D0PY6US KOFORDl SENTOSUS MICROPHYLLUS J0HNWRI6HTI INTERANDINUS
ANGUSTIDIGITUS
GERRHOPYGUS
REISSI
CLINATUS
INAEQUALIS
Figure 7. Ventral view of the fourth toe of all species of Phyllodactyliis (except
hetemms and pumilis) found on the mainland of western South America.
Phyllodactylus johnwrighti sp. nov.
Phyllodactyliis phacophorus, (part) Noble 1924, Occ. Paps. Boston Soc.
Nat. Hist., 5:111.
Holotype. Adult male, TCWC 27916, collected by James R. Dixon and
John W. Wright, 28 November 1968. Type locality. 13 km SSE (by road)
Hacienda Molino Viejo, 1190 m, Department of Cajamarca, Peru.
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Lizards of the Genus Phyllodactylus
57
Definition. A small gecko with a known maximum snout-vent length of
50 mm; enlarged preanal shield absent; dorsum and tibia with tubercles, absent
from thigh, forearm, and tail; some scales of median row beneath tail widened
and rounded; tail length about 49 per cent of total length; terminal lamellae
moderately enlarged, rectangular; dark stripe from nostril to arm insertion
always present (fig. 10b).
Diagnosis. This species differs from P. angustidigitus, gerrhopygus and
heterurus by the absence of enlarged preanal shield; from interandinus by
absence of dorsal stripes, presence of tubercles on tibia, and maximum num-
ber of 49 tubercles in paravertebral row from head to tail, rather than mini-
mum of 65 in interandinus; from kofordi and sentosus by absence of forearm,
tail, and thigh tubercles; from reissi by smaller maximum size, 50 mm rather
than 75 mm, and absence of wide, rectangular row of scales beneath tail; from
microphyllus and inaequalis by presence of regular longitudinal rows of
enlarged trihedral tubercles on dorsum, from lepidopygus and clinatus by
presence of tibia tubercles; from pumilus by absence of tail tubercles and
smaller supraorbital scales.
Description of holotype. Rostral rectangular, twice as wide as high, its
dorsal edge with vertical groove one-half depth of rostral; two internasals,
subrectangular, their median edges separated by two granules; internasals
bordered by rostral, nostril, six granules and postnasal of each side; nostril
surrounded by rostral, labial, internasal, and two postnasals; first labial in
broad contact with ventral edge of nostril; slight depression between inter-
nasals and in frontal region; 10 scales between eye and nostril; scales in
posterior loreal region about 2.5 times larger than midorbital scales; 22 scales
across snout at level of third labial, 14 scales across anterior edge of orbits,
20 midorbital scales; eye large, its diameter contained in snout length 1.6
times; eyelid with two rows of granules and one larger outer row of scales,
last three to four are pointed; ear diameter contained in eye diameter 2.4
times; ear opening slightly denticulate, scales on anterior and posterior mar-
gins small, rounded and subequal; rear of head granular with intermixed
tubercles; six supralabials and five infralabials to point below center of eye;
mental bell-shaped, as wide as long, bordered posteriorly by two postmentals;
postmentals rounded, slightly larger than other chin scales, their median edges
in broad contact; postmentals immediately followed by transverse row of
7 scales, followed by second row of 10 smaller scales; postmentals contact
first labial only of each side.
Dorsum with 14 longitudinal rows of enlarged trihedral tubercles; 10
rows reach to nape, 6 to base of tail; paravertebral row of 45 tubercles from
head to tail, 26 between axilla and groin; paravertebral rows of tubercles
separated from each other by five rows of granules, each tubercle separated
from preceding tubercle by one to two granules; postanal tubercles three on
each side; 26 scales across venter, 55 from throat to vent; tail without tuber-
cles, median row of scales beneath tail with every other scale distinctly
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a
Figure 8. (a) dorsolateral view of P. angustidigitus; (b) ventral view of preanal
plate of P. angustidigitus.
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Lizards of the Genus Phyllodactylus
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widened; arms and thigh with small flattened scales; thigh with five to six
tubercles intermixed among scales; tibia with 17 to 19 tubercles intermixed
among granules; lamellae formula for hand 6-8-9-10-8, foot 7-9-1 1-12-11;
claw slightly visible when viewed from below; terminal lamellae large, rectan-
gular, longer than wide.
Measurements in mm: Snout-vent length 39.0, axilla-groin length 17.4,
leg length 14.8, arm length 11.8, tail length 34.0, head length 11.1, head
depth 4.5, head width 7.6, snout length 4.2, eye diameter 2.7, ear diameter
1.1, distance from eye to ear 3.3.
Color in life: Ground color grayish tan; dorsum without distinct bands;
grayish brown spots on anterior half of body, fading posteriorly; grayish
yellow median stripe from nape to base of tail; posterior edge of scales on
limbs stippled with dark brown, rest of scale ground color; head with diffuse
spotting of grayish brown; grayish brown line from nostril to above arm inser-
tion, bordered above by dirty cream line from nostril to ear; tail with 2
brownish black bands with yellowish brown interspaces; ventral surfaces dirty
gray, with three to five black dots on each scale, more dense on tail.
Variation. Snout-vent lengths of subadult and adult males range from
32 to 40 mm (37.9), females from 33 to 44 mm (39.0), juveniles 21 to 31 mm
(26.4) , males plus females average 38.4 mm; tail length ranges from 46 to
51 per cent (48.8% ) of total length; postmentals vary from two to four, three
occurring twice and four once in 40 specimens; postmentals always contact
first labial; number of scales immediately following postmentals vary from
5 to 8 (6.2) ; midorbital scales vary from 17 to 21 ( 18.8), scales across snout
at level of third labials 20 to 26 (21.7) ; scales bordering internasals vary
from 6 to 9 (7.5), internasals separated along median line by granules in
97.7 per cent of sample; scales from nostril to eye vary from 10 to 12 (11.1);
scales across venter range from 26 to 32 (27.4), from throat to vent 47 to
57 (51.8); dorsal tubercular rows vary from 12 to 15 (13.9); paravertebral
tubercles from head to base of tail range from 40 to 49 (45.0), between axilla
and groin 22 to 29 (26.4) ; lamellae beneath fourth toe range from 10 to 12
( 1 1.4) ; tubercles absent from tail.
Dorsal pattern generally uniform tan to pinkish tan, few individuals
with double row of 9 to 12 small, grayish brown spots or transverse lines; tail
usually banded with 1 1 to 16 black bands, with ground color interspaces of
equal width; tail bands of juveniles usually bright orange and black, subadults
yellowish orange and black, and adults yellowish brown and black; iris of
eye pinkish gold, ventral surfaces generally dirty white in all specimens.
Comments. A distinct population of this species occurs on the east slope
of the Andes at an elevation of 2100 m. One specimen of this population was
taken by Noble from near the headwaters of the Rio Huancabamba in 1916,
and three individuals were taken by us from near the summit of the Andes on
the Olmos to Bagua road in 1968. These four specimens are similar in squama-
tion to our series from near Molino Viejo, but differ in color pattern and
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have a more decidedly depressed head and body. One male measured 50 mm
and a female 46 mm in snout-vent length. The dorsal pattern consists of a
fine reticulation of dark lines on ground color with a light median stripe. The
lateral dark stripe extends from the nostril to midbody. Until additional speci-
mens of this population are available, we are not in a position to analyze this
population further.
This species is named in honor of Dr. John W. Wright, who discovered
the original population along the Rio Huancabamba near Molino Viejo, and
who provided us with an immeasurable amount of help in the field.
Habitat and natural history. This species occupies two distinct vegeta-
tion communities, the high, barren eastern slopes of the Andes where only
a few cacti and shrubs are present, and a more extensive community of cacti,
shrubs, mesquite, and broad leafed trees along the Rio Huancabamba and
associated hillsides. The specimens from 2100 m were taken beneath dead
Cereus sp. and a cholla-like cactus. Those along the Rio Huancabamba, taken
during the day, were found beneath the bark of mesquite or under fallen trees.
After dark, individuals were found emerging from beneath fallen Cereus sp.
and from their branches as high as two meters above the ground. Later in the
evening about 20 individuals were found foraging around the bases of small
rocks, Opunta sp., weeds, and on open gravel soil. When disturbed, most
individuals sought cover in small patches of weeds or among small rocks.
Their nocturnal body temperatures averaged 23 °C and closely approximated
that of the substratum. Eggs of this species were found in rotting stumps of
dead trees. An examination of the oviducts of preserved females indicates that
this species lays one egg per clutch.
Distribution. This species has been found only on the arid slopes of the
Rio Huancabamba Valley and higher barren slopes of the east side of the
Andes in northwestern Peru (fig. 4).
Specimens (41 ) have been examined from the following localities.
PERU. Cajamarca: El Arenal MVZ 82196. 4.5 km W El Tambo TCWC
28088. 23.5 km WNW El Tambo TCWC 28104-05. 25.2 km WSW El Tambo
TCWC 28106. 13 km SSE Molino Viejo LACM 48476, 48479-94, TCWC
27916, 28089-90, 28092-103. 11 km WNW Las Juntas LACM 48477-78,
TCWC 28091. Piura: Huancabamba AMNH 28466.
Phyllodactylus pumilus sp. nov.
Holotype. Adult male, USNM 167227, collected by James A. Peters,
October 1965. Type locality. 1 km S Manta, Department of Manabi, sea level,
Ecuador.
Definition. A small gecko with a known maximum snout-vent length
of 51 mm; enlarged preanal shield absent; dorsum, tibia, and part of tail with
tubercles; scales of median row beneath tail widened and rounded posteriorly;
tail about 50 per cent of total length; scales in supraocular region largest of
interorbital series; dark brown to black stripe from nostril to arm insertion
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Lizards of the Genus Phyllodactylus
61
always present; terminal digital lamellae moderately enlarged and somewhat
rectangular in shape (fig. 14c).
Diagnosis. This species differs from P. angiistidigitus, gerrhopygus, and
heterurus by absence of enlarged preanal shield; from interandinus by absence
of dorsal stripes and presence of tubercles on tail and tibia; from kofordi and
sentosus by absence of forearm tubercles and tubercles on distal two-thirds
of tail; from microphyllus and inaequalis by presence of regular rows of
enlarged trihedral tubercles on dorsum; from lepidopygus and clinatus by
presence of tibia turbercles; from reissi by smaller maximum snout-vent length
of 51 mm rather than 75 mm, scales in supraocular region largest of inter-
orbital series, and at least two rows of enlarged, flattened scales on proximal
one-fourth of tail; from johnwrighti by presence of at least two rows of
enlarged, flattened scales on proximal one-fourth of tail; more transverse
Figure 9. (a) dorsolateral view of P. gerrhopygus', (b) dorsal view of P. micro-
phyllus.
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ventral scales from throat to vent, 53 to 62 (57.6) rather than 47 to 57 (46.4) ;
fewer paravertebral tubercles from head to base of tail, 37 to 43 (39.4) rather
than 40 to 49 (45.0).
Description of holotype. Rostral twice as wide as high, its dorsal edge
with short median, vertical groove; two internasals, somewhat rectangular
and in contact along median edges; nostril surrounded by rostral, labial, inter-
nasal, and three postnasals; first supralabial in broad contact with ventral edge
of nostril; shallow depression between internasals and in frontal region; 1 1
scales between nostril and eye; scales in posterior loreal region about three
times larger than interorbital scales; 19 scales across snout at level of third
labials, 17 interorbital scales, 18 at anterior edge of orbits; eye large, its diam-
eter contained in snout length 1.7 times; eyelid (brille) with one to two rows
of granules and one larger outer row of scales; diameter of ear contained
in eye diameter about three times; ear opening strongly denticulate on anterior
and posterior margins; rear of head granular with a few larger tubercles inter-
mixed; six supralabials and five infralabials to point below center of eye;
mental bell-shaped, as wide as long, bordered posteriorly by two postmentals;
postmental about as long as wide, their median edges in broad contact; post-
mentals immediately followed by transverse row of six scales, followed by
second row of 10 smaller scales; postmentals contact first labial only.
Dorsum with 14 longitudinal rows of enlarged trihedral tubercles that are
somewhat flat; paravertebral row with 42 tubercles from rear of head to base
of tail, 25 between axilla and groin; paravertebral rows separated from each
other by four rows of granules; five rows of tubercles reach nape, six to base
of tail; each tubercle of dorsal series separated from preceding tubercle by
none to two granules; postanal tubercles number four to each side; venter with
27 longitudinal and 61 transverse rows of scales.
Dorsal surface of arm with large, flat scales; dorsal surface of thigh with
three to five tubercles scattered among granular scales; lamellae formula for
hand 7-S-9-9-7, foot 7-8-12-12-10; claw slightly visible when viewed from
below; terminal lamellae slightly longer than wide, moderately enlarged;
distal three-fourths of tail regenerated, proximal one-fourth with a 6-4-2
reduction in number of tubercles per whorl, decreasing in number towards
distal portion of tail; median row of scales beneath tail widened, rounded
posteriorly.
Measurements in mm: snout-vent length 49.0, axilla-groin length 22.0,
leg length 17.5, arm length 12.5, tail length 44.5, head length 13.0, head width
9.3, head depth 5.7, snout length 5.2, eye diameter 3.0, ear diameter 1.1, eye
to ear distance 3.7.
Color in alcohol : ground color grayish tan; dorsal surface of body with
faint indication of six light brown bands, broken along median line of body
by longitudinal stripe of ground color; limbs without pattern, ground color;
top of head from occiput to snout uniform light brown; brownish stripe from
nostril to arm insertion, passing over ear; supralabials spotted with dark brown.
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Lizards of the Genus Phyllodactylus
63
less so on infralabials; venter grayish white with a few minute black dots on
each scale; ventral surfaces of hands and feet slightly darker than venter; tail
with faint indication of banding.
Variation. Snout-vent length of young adult and adult males range from
37 to 51 mm (44.0), females from 41 to 44 mm (42.0), juvenile 25 mm;
males plus females average 43.2 mm; tail partly regenerated in all specimens
except one and averages about 50 per cent of total length; two postmentals
in all, contacting first labial only in 100 per cent of sample; number of scales
immediately following postmentals vary from 4 to 8 (6.1 ) ; midorbital scales
vary from 15 to 18 ( 16.9) , scales across snout at level of third labial 19 to 22
(20.1 ) ; scales bordering internasals vary from 5 to 8 (6.0) ; internasals in
contact along their median edges in 90.1 per cent of sample; scales between
nostril and eye vary from 10 to 13 ( 1 1.6) ; scales across venter range from
27 to 30 (28.5), from throat to vent 53 to 62 (57.5) ; dorsal tubercular rows
vary from 12 to 14 ( 13.3) ; paravertebral tubercles from head to base of tail
range from 37 to 43 (39.4), between axilla and groin 23 to 27 (24.7) ; lamel-
lae beneath fourth toe vary from 1 1 to 13 ( 12.0) ; at least two rows on enlarged,
flat scales (may be elevated and tubercular-like) on proximal one-fourth of tail.
Dorsal color pattern highly variable, from an almost uniform light brown
to definite paravertebral rows of dark brown spots or bands; head usually
uniform light brown, but faintly mottled with light gray in some specimens;
dark brown stripe from nostril to arm insertion in all but one specimen which
has stripe continuing to groin region; median grayish stripe on dorsum in all
specimens, but faintly interrupted by black reticulating lines in one; tail banded
brown and gray in only specimen with complete, original tail; venter grayish
white in all.
Habitat and natural history. Dr. James A. Peters (personal communica-
tion) states that the type locality is a vertical rocky cliff on the beach, just
above the high tide zone, 1 km south of Manta. The cliff surface is somewhat
rotten, crumbles easily, and contains numerous holes and crevices. Four speci-
mens of P. pumilus were taken from beneath rocks or in crevices along the
cliff face. Peters indicated that he had not seen other specimens of Phyllo-
dactylus this close to the ocean at any other point on the Ecuadorian coast.
Peters indicated that all specimens of this species were taken during day-
light hours. Other specimens were found beneath loose bark on trees and
stumps in a semi-desert area with low scrubby vegetation, or in a transition,
dry to subdry forest.
Phyllodactylus reissi were taken with pumilus from the stump of an old
ceiba tree 6 km E of Manta, and at the type locality.
Distribution. This species has been taken west of the Cordillera de Balzar
in the department of Manabi of Ecuador, from sea level to approximately
200 m (fig. 3).
Specimens (11) have been examined from the following localities:
ECUADOR. Manabi: 8 km W Jipijapa USNM 167184. 1 km S Manta USNM
Figure 10. (a) dorsal view of P. interandinus; (b) dorsolateral view of P. john-
wrighti.
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Lizards of the Genus Phyllodactylus
65
Figure 11. (a) dorsal view of P. kofordi (holotype); (b) dorsal view of P. clinatus
(holotype).
167227-30. 6 km E Manta, 5 km W Montecristi USNM 167233. 11 km W
Portoviejo USNM 167185, 167226. 9 km N San Vicente USNM 167187,
167231-32.
Adaptation to Habitat
Species of Phyllodactylus are, in the more restricted sense, geckos of arid
and semiarid regions. Only on the mainland of British Honduras and some
islands of the Caribbean are they found in a more tropical environment.
Adaptations of species of Phyllodactylus for living in arid deserts and
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No. 192
tropical deciduous forests are generally of two types, scansorial and terrestrial.
Most of these lizards are found in areas where sufficient physical objects are
available for scansorial habits, i.e. trees, rocks and cliffs. All species of Mexico,
Central America, and eastern South America are scansorial. Most prefer a
rock substratum, but are frequently found on trees and shrubs. Only on the
mainland of western South America does one find a shift to the terrestrial
habit, and even then the terrestrial species are occasionally scansorial.
Underwood (1954) indicates that the most primitive condition in geckos
is one of terrestrial habits with simple, clawed digits. Adaptations to a scan-
sorial habit involved the development of pilose, friction lamellae beneath the
digits, and in some cases, beneath the tip of the tail. He also states that there
has been a number of independent reversions to terrestrial habits with a loss
or reduction in the size of the lamellae.
Of the 15 mainland species of western South America, four are ter-
restrial : P. microphyllus, angustidigitus, gerrhopygus, and sentosus. Of these
four species, microphyllus has become well adapted to an open, deep sand
habitat, through reduction in size of the terminal lamellae, shorter and more
muscular limbs, and swollen scales about the nostril with the nasal opening
a very small slit and lying in an anterior, dorsolateral position.
Phyllodactylus angustidigitus has become adapted to a shell and rubble
beach habitat through reduction in the terminal lamellae, enlargement of the
claw, elevated and swollen scales about the nostril, but unlike microphyllus,
the limbs are very slender, long, and without much muscular development.
Little is known about the habits of sentosus, but all known specimens were
taken from the ground and they have relatively small terminal lamellae and
well developed claws, but the region about the nostril is unswollen. We find
it difficult to explain the presence of enlarged terminal lamellae in a terrestrial
species, such as gerrhopygus. This species has very large, well developed ter-
minal lamellae like those of scansorial species (fig. 7), but has the area about
the nostril greatly swollen, and the limbs are intermediate in size and shape
between angustidigitus and microphyllus.
There appears to be differential development for a terrestrial habit for
each of the four species, with evolutionary trends variable within each popu-
lation. For example, P. microphyllus taken from hard surface sands or con-
glomerate soils have slightly larger terminal lamellae than those from soft
sands. However, in gerrhopygus, the size of the terminal lamellae varies slightly
within each population and does not seem dependent upon soil type. The
reduction in the size of the terminal lamellae coupled with an enlargement
of the claw in angustidigitus, seems to be a special modification for living on
shell and rubble beaches. Whether the reduction in the terminal lamellae and
enlargement of claws are for walking on shell and rubble, or for turning over
these objects in search of food is not known, but the claw is approximately
four times the size of the claw of gerrhopygus, a species that is frequently
found on sandy beaches.
1970
Lizards of the Genus Phyllodactylus
67
The terminal lamellae of the scansorial species (P. reissi, lepidopygus,
and probably dinatus) are generally greatly enlarged. Phyllodactylus reissi has
exceptionally large toe pads, possibly influenced by its large body size. Several
geckos are intermediate in their microhabitat preferences, P. kofordi, inae-
qualis, phnwrighti, and interandinus, usually spending the day under bark or
exfoliations, and at night foraging about on the ground. As one might expect,
the toe pads are intermediate in size, reflecting an adaptation to both terrestrial
and scansorial habits. In addition, a toe pad of intermediate size may facilitate
terrestrial locomotion over a broken substratum and surface debris typical of
the habitats where these geckos are found. Seemingly, the more scansorial a
species becomes, the more natural selection favors an enlargement of the ter-
minal lamellae. However, this does not explain the enlarged terminal lamellae
of gerrhopygus. This species has been taken from terrace walls and walls of
houses indicating it is not completely terrestrial. There may have been a recent
shift from a scansorial to a terrestrial habit, and natural selection for small
terminal lamellae is presently acting upon the population. However, it is pos-
sible that gerrhopygus is adapted for running upon soft surface sand and for-
aging for surface food items. This would, in part, account for the small claws
and enlarged terminal lamellae. Larger and more lengthened claws may be
used for turning over surface litter or for digging under it in search of food
(microphyllus, sentosus, angustidigitus). Large terminal lamellae would likely
hinder the ability to dig or clasp objects in the search for food, wheras a
surface feeder might retain the large lamellae, enabling it to move faster,
farther, and with more ease, in search of surface food items.
The swollen area about the nostril seems to be an adaptation for living
in regions where deep sand is constantly being moved about by wind action.
The scales about the nostril are elevated, rounded (swollen), and the position
of the nostril has been shifted to a more anterior, dorsolateral position. The
nasal opening is narrow and partially protected by an anterior extension of the
upper postnasal scale into the nasal aperture. In effect, the elevated and rounded
scales surrounding the nostril form a tubular opening. This type of nostril
provides adequate air passage and prevents sand from clogging the aperture
wLile the individual rests in daytime retreats or forages about on the sand at
night.
Phyllodactylus sentosus is the only representative of a terrestrial species
that does not have a tubular condition about the nostril. The substratum where
it lives is more of a conglomerate soil, and selection for a tubular nostril is not
one of the critical adaptive features for survival. None of the scansorial species
have tubular nostrils.
Sympatric Relations of PHYLLODACTYLUS in
Western South America
An awareness of sympatry is essential in overcoming the subjectivity of
morphological taxonomy. This section will briefly document known cases of
68
Contributions in Science
No. 192
Figure 12. (a) dorsal view of P. sentosus (holotype) ; (b) dorsal view of P. inae-
qualis.
1970
Lizards of the Genus Phyllodactylus
69
sympatry in western South America as well as describe certain aspects of
ecological and evolutionary differences among the sympatric species.
The geographic distribution of species of Phyllodactylus is summarized
in figs. 2-5. Known cases of sympatry are noted in Table 2.
When geckos occur sympatrically one consistently observes distinct dif-
ferences in the degree of arboreality. For example, although both P. reissi
and interandinus utilize arboreal perches for diurnal retreats, only interan-
dinus moves to the ground for nocturnal foraging.
Phyllodactylus reissi and kofordi are commonly sympatric but have very
little overlap in perch height. Phyllodactylus reissi is usually a scansorial for-
ager, while kofordi is primarily a ground forager.
In some localities P. inaequalis is found with kofordi and reissi in sym-
patry. Phyllodactylus inaequalis, like kofordi, is a terrestrial forager. The
relative abundance of these species seems to vary from locality to locality
based on collecting records. It is not clear whether this is due to competition
and/or to subtle environmental differences favoring one species at a given
locality. There is some evidence of microhabitat separation between these
two species at Cerro de la Vieja where inaequalis forages near the bases of
boulders and kofordi forages more on open ground.
Another difference commonly found among sympatric geckos is in body
size, presumably reflecting differences in prey size. In sympatric situations the
only known exceptions to extreme body size differences are with P. kofordi
and inaequalis as well as kofordi and cUnatus. The fact that all of these geckos
are small and appear to forage in different microhabitats probably accounts
for their coexistence.
Some cases of sympatry are not clearly understood. The replacement of
P. microphyllus, a sand gecko, by kofordi, normally a foothill gecko, along
the sandy coast in extreme northwestern Peru is puzzling. Climatic, competi-
tive, or physical environmental factors may be involved.
Selander and Giller ( 1963) observed that broad sympatry in wood-
peckers of the genus Centurus occurs between distantly related species. Con-
versely, closely related species are allopatric or only narrowly sympatric. These
workers concluded . . it is suggested that the evolution of reproductive
isolating mechanisms tends to proceed more rapidly than does the evolution
of ecologic isolation, with the result that competition and other interspecific
interactions prevent extensive sympatry of closely related species . . .”
Pending completion of our osteological and karyological studies, the
intrageneric relationships of the western South America Phyllodactylus are
tentative. Strictly on the basis of external morphology, we suggest the follow-
ing species are closely related and group them: gerrhopygus, heterurus, an-
gustidigitus; microphyllus, inaequalis; lepidopygus, clinatus; johnwrighti,
pumilus, interandinus; sentosus, kofordi; and reissi. Comparison of these
groups with Table 2 shows only one known case of sympatry within a group,
that of gerrhopygus and heterurus. However, the known extent of sympatry
70
Contributions in Science
No. 192
Table 2
Sympatric relationships of 15 species of Phyllodactylus
from the mainland of South America.
5
3
3
5c
Cc
•5
3
5c
5c
s;
>3
clinatus
dixoni
CX
o
Oc
heterurii
*3
3
3^
3
3
•2?
3
-3
O
kofordi
3.
0
s.
-3
3.
2
3
’i
pumilus
reissi
sentosus
ventralis
angiistidigitus
cl i fiat us
dixoni
gerrhopygiis
heteriirus
inaequalis
interandinus
johnwrighti
kofordi
lepidopygus
microphyllus
pumilus
reissi
sentosus
V entrails
X XXX
X
X
X X X
X
X X X X
X
X XX XX
X
X XX X XX
X
(X in sympatry)
of these species is based on one locality, and the status of heterurus is in
doubt. Where broad sympatry does occur (e.g., among inaequalis, kofordi,
and reissi), it is between or among very distinct species. Hence, Selander and
Ciller’s (1963) conclusions might well be applicable to species of Phyllodac-
tylus in South America.
1970
Lizards of the Genus Phyllodactylus
71
%
Figure 13. (a) dorsolateral view of P. lepidopygus; (b) dorsal view of an adult and
young of P. reissi.
72
Contributions in Science
No. 192
Figure 14. (a) dorsal view of P. ventralis; (b) dorsal view of P. dixoni; (c) dorsal
view of P. pumilus (holotype).
1970
Lizards of the Genus Phyllodactylus
73
Gazetteer
There are a large number of collecting localities in Ecuador, Peru, and Chile
not shown in gazetteers, on maps, or generally unavailable to scientists working
with material from western South America. We present the following list of locali-
ties with coordinates, arranged by country and political division.
CHILE
Tarapaca
Arica
18°
29'S
o
O
r-
20'W
Iquique
20
13S
70
low
Pica
20
30S
69
20W
Pintados
20
38S
69
38W
Pisagua
19
36S
70
13W
Putre
18
12S
69
35W
ECUADOR
Azuay
San Erancisco
3
37S
79
30W
El Oro
Machacha
3
16S
79
57W
Santa Rosa
3
27S
79
58W
G nay as
Ancon Camp
2
20S
80
52W
Colonche
2
OIS
80
40W
Daule River
1
53S
79
58W
Guayaquil
2
lOS
79
52W
Playas
2
38S
80
23W
Puente de
Chimbo
2
IIS
79
07W
Puna
2
42S
79
55W
Punta Cornero
2
18S
80
55W
Salinas
2
13S
80
50W
Loja
Loja
4
OOS
79
13W
Rio Casanga
4
08S
79
49W
Manabi
Jipijapa
1
20S
80
35W
Manta
0
57S
80
45W
Portoviejo
1
03S
80
27W
San Vicente
0
36S
80
24W
PERU
Amazonas
Bagua Chica
5
38S
78
35W
Bagua Grande
5
47S
78
26W
Marahon
Valley
4
30S
78
27W
Ancash
Chimbote
Valley
9
07S
78
38W
Culebras
9
56S
78
14W
Pativilca
10
40S
77
48W
Areqiiipa
Arequipa
16
24S
71
32W
Camana
16
37S
72
42W
Chucarapi
17
04S
71
44W
La Joy a
16
45S
71
52W
Mollendo
17
02S
72
OIW
Tambo
17
06S
71
48W
Vitor
16
26S
71
49W
Yura
16
12S
71
42W
Cajamarca
Bellavista
5
37S
78
39W
Chinchipe
Valley
5
28S
78
32W
El Arenal
5
56S
79
17W
El Tambo
5
46S
79
24W
Las Juntas
5
57S
79
14W
Molino Viejo
5
49S
79
23W
Perico
5
15S
78
45W
Ica
Ica
14
05S
75
44W
Isla Sangallan
13
50S
76
28W
Isla Viejas
14
17S
76
12W
Jaguay
13
23S
76
12W
Lagunillas
Bay
13
54S
76
18W
Marcona
15
20S
75
low
Paraca Bay
13
50S
76
16W
Paracas
13
48S
76
15W
Paracas
Peninsula
13
48S
76
24W
Pozo Santo
13
54S
76
03W
Puerto Pisco
(new)
13
47S
76
17W
San Juan Port
15
22S
75
12W
Junin
Chanchamayo
10
42S
75
08W
74
Contributions in Science
No. 192
Lambayeque
Chiclayo
6
46S
79
SIW
Eten
6
54S
79
S3W
Jayanca
6
24S
79
SIW
Mocupe
7
oos
79
37W
Morrope
6
32S
80
02W
Motupe
6
09S
79
43W
Naupe
5
37S
79
S4W
Olmos
Libertad
6
OOS
79
4SW
Chiclin
Jequetepeque
7
SOS
79
low
Valley
7
21S
79
36W
Pacanguilla
7
lOS
79
28W
Pacasmayo
Lima
7
24S
79
34W
Ancon
11
46S
77
low
Asia
12
47S
76
3SW
Cerro Azul
13
03S
76
30W
Chorillos
10
46S
77
46W
Chosica
Cruz de
11
54S
76
42W
Hueso
Isla San
12
23S
76
46W
Lorenzo
12
OSS
77
ISW
Lima
Lomas de
12
03S
77
03W
Lechay
11
27S
77
18W
San Antonio
Verrugas
12
37S
76
37W
Canyon
Pi lira
11
S3S
76
32W
Bayovar
5
SOS
81
03W
Bocana
4
41S
80
18W
Cabo Blanco
4
ISS
81
14W
Capullana
Point
4
29S
81
17W
Catalinas
4
ns
80
S8W
Cerro Amotape
4
40S
80
S8W
Chongollapi
S
26S
79
37W
Chulucanas
S
06S
80
low
El Alto
4
17S
81
12W
Fondo
4
28S
81
09W
Huancabamba
S
14S
79
27W
Isla Lobos de
Afuera
6
S7S
80
42W
Isla Lobos de
Tierra
6
27S
80
S2W
Las Lomas
4
40S
80
ISW
Lobitos
4
26S
81
16W
Los Organos
4
09S
81
07W
Monte Grande
4
27S
81
02W
Negritos
4
38S
81
19W
Paita
S
06S
81
07W
Parinas Valley
4
30S
81
06W
Piura
S
ns
80
38W
Punta Aguja
s
47S
81
04W
Punta Sal
s
SIS
80
S6W
Reventazon
6
lOS
81
oow
Sechura
s
34S
80
sow
Sechura Desert
s
SOS
80
40W
Sullana
4
S3S
80
41W
Talara
4
34S
81
17W
Trie Trac
S
SIS
81
OOW
Tiimbes
Cancas
4
OOS
81
oow
Las Vocas
3
2SS
80
19W
Puerto Pizzaro
3
39S
80
24W
Quebrada Seca
4
23S
81
14W
Acknowledgments
We are endebted to V. Aellen, C. Rivero-Bianco, J. A. Bohlke, C. M.
Fugler, E. Gambini, A. G. C. Grandison, R. Inger, K. Klemmer, W, Ladiges,
A. E. Leviton, G. Orces, J, A. Peters, J. A. Roze, J. A. Sloan, H. M. Smith,
R. C. Stebbins, D. W. Tinkle, H. Wermuth, E. E. Williams, J. W. Wright,
and R. G. Zweifel for the loan of material in their care.
We are especially grateful to Drs. Carl Koford, John W. Wright, F.
McCollum, and R. B. Hamilton for their able assistance and companionship
while pursuing our studies of geckos in Peru; to Drs. H. W. and M. Koepcke,
Mr. and Mrs. W. Grindley, Ing. H. Ramirez, G. and Sr. S. Bertarelli, Drs. F.
Blancas and E. Gambini for the many courtesies extended to us during our
1970
Lizards of the Genus Phyllodactylus
75
trips to Peru; to Dr. Janis A. Roze for field notes and a photograph of habitat
in Venezuela; and to Dr. James A. Peters for field notes on Ecuadorian geckos.
This study was supported through a grant to Dixon from the Texas A&M
University Organized Research Fund (1968), a National Science Foundation
Traineeship to Huey through the University of Texas (1968), and a expedi-
tion to Peru ( 1967), sponsored by the Museum of Vertebrate Zoology, Uni-
versity of California.
Resumen
Se hace una breve descripcion del habitat natural de las especies de Phyllo-
dactylus encontradas en el continente de America del Sur. El micro-habitat na-
tural de cada especie se describe hasta donde ha sido posible. Poca a ninguna
informacion se aporta sobre el micro-habitat de P. heterurus, setosus y clinatus.
Cuatro especies, P. angustidigitus , gerrhopygus, microphyllus y setosus son
primariamente terrestres y alopatricas en distribucion. Diez especies, P. clina-
tus, dixoni, inaequalis, interandinus , johnwrighti, kofordi, lepidopygus, pumi-
lus, reissi y vent rails son aboricolas y unicamente johnwrighti, v entrails y
dixoni no viven simpatricamente con una o mas de las otras especies arbori-
colas. En donde tres o mas especies viven simpatricamente, las areas utilizadas
para reposar durante el dia y para la obtencion de alimento se sobreponen
escasamente.
Con la excepcion de P. reissi, la cual produce dos huevos, todas las especies
de Phyllodactylus encontradas en America del Sur continental depositan un
solo huevo por nidada. La deposicion de huevos en forma comunal parece
ocurrir con frecuencia entre el mayor numero de especies, incluyendo aquellas
que viven simpatricamente : P. interandinus y reissi, y kofordi y reissi.
Las especies del genero Phyllodactylus parecen tener adaptaciones especifi-
cas a sus micro-habitates naturales. Aquellas especies que habitan areas roco-
sas se caracterizan por la presencia de laminillas digitales terminales ensan-
chadas, cabeza y cuerpo comprimidos dorsoventralmente y ventanas de la
nariz localizadas lateralmente; en el caso de especies que habitan un micro-
habitat arenoso las laminillas digitales terminales son pequenas, la cabeza y el
cuerpo redondeados y las ventanas de la nariz son valvulares y estan localiza-
das dorsolateralmente ; finalmente, las especies que viven en playas de concha
tienen laminillas digitales con unas alargadas, cabeza y cuerpo delgado y ven-
tanas de la nariz con valvulas situadas dorsolateralmente. Dependiendo del
grado en que el comportamiento es arboricola o terrestre algunas especies de
Phyllodactylus poseen las adaptaciones descritas en forma intermedia.
Para cada especie, excepto P. heterurus, se incluye un mapa de distribucion
junto con fotografias; ademas, se presenta una lista de localidades importantes.
De los holotipos y sintipos de las especies descritas de America del Sur con-
tinental, existen diez (dixoni, inaequalis, lepidopygus, magister, microphyllus,
nigrofasciatus, phacophorus, pumilus, reissi, y v entrails), ocho de los cuales
han sido examinados.
76
Contributions in Science
No. 192
Se describen siete nuevas especies (angustidigitus, clinatus, interandinus,
johnwrighti, kofordi, pumilus y setosus); nueve nombres antiguos se relegan
al sinonimio de otras especies ( abrupteseriatus, baessleri, guayaquilensis, lo-
bensis, rnagister, nigrofasciatus, phacophorus, underwoodi, variegatus).
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Occ. Pap., 5: 107-113.
Parker, H. W., 1935. Some lizards from Venezuela and the Dutch Leeward Islands.
Ann. Mag. Nat. Hist., 15: 480-484,
Peters, J. A., 1960. The snakes of Ecuador, a check list and key. Harvard, Mus.
Comp. Zool., Bull., 122: 491-541.
Peters, W. C. H., 1862. Mittheilung uber einen neuen Phyllodactylus aus Guayaquil.
Monatsb. Akad. Berlin, pp. 626-627.
Rivero-Blanco, C., and A. R. Lancini, 1968. Phyllodactylus dixoni; Una nueva
especie de lagarto (Sauria: Gekkonidae) de Venezuela. Soc. Cienc. Nat. La
Salle, Memoria, 27: 168-175.
Robinson, D. A., 1964. Peru in four dimensions. American Studies Press, S.A.,
Lima, Peru, pp. iii-424.
Roze, J, A., 1964. La herpetologia de la isla de Margarita, Venezuela. Soc. Cienc.
Natl. La Salle, Mem., 24(69) : 209-241.
Ruthven, a. G., 1922. The amphibians and reptiles of the Sierra Nevada de Santa
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Selander, R. K., and D. R. Gillier, 1963. Species limits in the woodpecker genus
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Shelford, V. E., 1926. Naturalist’s guide to the Americas. Williams & Wilkins Co.,
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78
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— , 1968b. Lagartos Brasileiros de familia Gekkonidae (Sauria). Arq. Zool.
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Marina, 3(1-2): 1-18.
Accepted for publication February 10, 1970
I LOS
1^' ANGELES
' COUNTY
MUSEUM
CONTRIBUTIONS
IN SCIENCE
'
ikuMBER 193
July 1, 1970
HADROSAURIAN DINOSAUR BILLS— MORPHOLOGY
AND FUNCTION
By William J. Morris
*!!
1
Hi
ii
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4
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
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HADROSAURIAN DINOSAUR BILLS— MORPHOLOGY
AND FUNCTION
By William J. Morris^
Abstract: Reconsideration of the morphology and function
of the bill of hadrosaurian dinosaurs is based upon an excellently
preserved bill mold of a recently discovered specimen (LACM
23502) assigned to the genus Anatosaunis. Morphology suggests
a filtering function for the bill and adaptation for discharge of
fluid and small particles following intake and mastication of plant
or invertebrate food.
Acknowledgments
The excellent specimen of Anatosaurus cf. annectens (LACM 23502)
upon which this study is based was discovered and collected by H. Garbani
and skillfully prepared by M. Odano, both of the Los Angeles County Museum
of Natural History. The skeleton was found in the Hell Creek Formation south
of Ft. Peck Reservoir, NE Va, Sec. 36, T21N, R35E, Montana.
The project, leading to this discovery as well as other important dinosaur
finds, was generously supported by Mr. and Mrs. William T. Sesnon, Jr.,
patrons of the Los Angeles County Museum of Natural History.
Constructive criticism of the manuscript by Dale Russell, Canadian
National Museum, and Theodore Downs of the Los Angeles County Museum
of Natural History is appreciated. Others on the staff of the Los Angeles County
Museum whose help is acknowledged are: David Forstch and Alan Tabrum
for their interest and discussion, and Joseph Cocke for his drawing of Figure 2.
The remainder of the illustrations and photographs were done by the author.
Introduction
Dinosaurs of the family Hadrosauridae are among the most familiar Late
Cretaceous fossils. Indeed, the first dinosaurs described from North America,
belonged to these so called duckbilled forms. Prior to a brief description by
Cope (1883) the presence of a horny bill in hadrosaurs was conjectural. Cope,
in preparing a specimen of Anatosaurus annectens, discovered portions of a
mold of the bill. Later two other specimens in which most of the bill was
preserved were described for A. annectens by Versluys (1923) and for
Corythosaurus excavatus by Sternberg ( 1935) . In all three cases we must rely
upon the description of the bill as the original molds were either destroyed
during preparation or subsequently lost. As a result, a recently discovered
specimen of Anatosaurs cf. annectens (LACM 23502) from the Hell Creek
beds of Montana containing a well preserved mold is most important in
^Research Associate in Vertebrate Paleontology, Los Angeles County Museum of
Natural History; and Professor of Geology, Occidental College, Los Angeles,
California 90041.
1
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Contributions in Science
No. 193
evaluating the morphology and function of the hadrosaurian bill. The letter
designations are used to designate specimens in various museums: LACM,
Los Angeles County Museum of Natural History; ROM, Royal Ontario
Museum; NMC, National Museum of Natural Sciences, Ottawa.
Bill Morphology of Anatosaurus cf. A. ANNECTENS (LACM 23502)
The specimen is a mature individual approximately forty feet long. The
skull is only slightly distorted mainly along a parasagittal section approximately
midway between the orbits and narial openings. The mandibles, though offset
to the right, are not distorted relative to the midline so that the bill mold is
undistorted, (Fig. 1). Morphologically, the specimen is very similar to
A. annectens but there are some differences between it and the holotype. Until
the time when careful study has been made regarding individual variability of
hadrosaurian dinosaurs, it is hazardous to make positive, specific references for
specimens other than the type. This is especially true regarding the LACM
specimen for it was found further north than the Lance Formation, Wyoming,
localities from which the types of A. annectens were obtained.
The bill mold is composed of gray, lithic siltstone, the surface is accentu-
ated by a thin veneer of iron oxide. In the figures, the mold appears dark due to
the application of preservative although it has the same composition as the
matrix. The entire anterior surface (attached dorsally to the premaxillary
region) is preserved and only a small portion of the ventral edge is lacking.
Only the left side is preserved.
Before discussing the form and function of the bill it is important to
describe the geometry of the mold. Errors in interpretation by other authors
seems largely to have resulted from misinterpretation of the geometry of the
mold.
When the specimen was discovered, anterior and posterior sides of the
mold were preserved with a very thin surface of weakness separating them. The
surface, shown in Figures 1, A, B, C and D, is the posterior side of the mold,
the one closest to the premaxillary region of the skull. It is important to note
that matrix on both sides of the preserved surface was carefully prepared and
other structures were not found within the matrix except the surface of the
mold. Hence the entire bill, as preserved, is represented by two surfaces
separated by less than 0.01 mm, one of which is shown in the figures.
The surface of the mold is vertically fluted, containing nine or possibly
ten V-shaped channels. These are separated by dorsal-ventral pillars, rounded
Figure 1. Partially prepared skull of Anatosaurus cf. annectens, LACM 23502.
(A) Complete skull, left side. The bill mold is not preserved on this side but the
edge of the mold can be seen extending ventrally from the premaxillary below the
predentary of the lower mandible. Note that the free margin of the mold extends
below the predentary and is separated from the predentary by siltstone matrix.
(B) Anterior undulating surface of the bill mold showing the fluted channels
preserved as an iron oxide veneer on the siltstone matrix. (C) Dorsal oblique
aspect of the skull illustrating the very terminal position of the bill mold along
the ventral margin of the premaxillary. (D) Ventral oblique view showing the
bill mold and the matrix layer separating it from the predentary.
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Hadrosaurian Dinosaur Bills
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Contributions in Science
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Hadrosaurian Dinosaur Bills
5
in cross section. Laterally, towards the maxillaries, the fluted surface is re-
placed by a smooth triangular plate.
The mold is approximately 110 mm long at the junction of the pre-
maxillaries and it extends ventrally about 50 mm below the anterior margin
of the predentary, decidedly overlapping the front of the lower jaw. Although
the extreme ventral margin of the mold appears to be missing, the taper of the
pillars and channels suggests that the entire bill was not much longer than
shown in this specimen.
The premaxillary of the specimen, dorsal to the mold on the left side,
is not well preserved and the surface separating bill and upper jaw cannot be
seen. There is, however, little doubt that the bill was united with the upper
mandible. A complimentary structure that would have been attached to the
lower mandible was not found even though conditions for its preservation must
have been similar to those for the upper bill mold.
Interpretive Morphology
The surface of the mold forms an immediate extension of the pre-
maxillary without break or offset. The actual horny bill must have been
affixed anterior to the mold and, in order to be secure, must have extended up
and onto the surface of the premaxillaries. Although there is no direct evidence
indicating the posterior limit of the bill, it probably did not extend farther than
the excavation surrounding the nares in the premaxillaries.
The geometry suggested by the anterior position of the mold suggests that
the mold reflects the interior surface of the bill. Probably some compression
of the bill occurred during compaction of the silt now forming the matrix, and
Figure 2. Restoration of the bill on the skull of Anatosaunis cf. annectens, LACM
23502. The undulating, fluted inner surface of the bill is shown. When the mandibles
were closed and the predentary opposed to the bill, channels opening from the
mouth were formed.
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Contributions in Science
No. 193
it is impossible to determine the original thickness of the specimen. The fluted
surface was evidently confined to the interior of the bill as no comparable
surface was found against the more dorsal surfaces of the premaxillary. Con-
ceivably the original exterior fluting could have been destroyed during com-
paction but then why was it not also destroyed on the more ventral portion?
It seems best to reconstruct a bill with the fluted surface only on the internal
side.
The predentary is poorly preserved and furnishes no evidence for a bill on
the lower jaw. Nor was evidence for the presence of a lower bill found in the
matrix anterior to the predentary.
A restoration of the probable appearance of the upper bill is shown in
Figure 5.
Discussion of Previously Described Bill Molds
The specimen described by Cope was destroyed during preparation of
the skull. Cope did not give a reason for destroying the mold, but one cannot
help conjecturing that he thought his description vivid enough that future
reference to the specimen would be unnecessary. The mold of the Cory-
thosaurus excavatus specimen, except for a very small and unilluminating
fragment, could not be located. In the third example Versluys apparently used
the mold to cast what he took to be a plaster replica of the bill and, after
obtaining the cast, the natural mold was not retained.
Cope appears to have misinterpreted a structure very similar to that of
LACM 23502. The full text follows (1883:106) : “Dermal or corneous struc-
tures have left distinct traces in the soft matrix about the end of the beak-like
muzzle. Laminae of brown remnants of organic structures were exposed in
removing the matrix. One of these extends as a broad vertical band round the
sides, indicating a vertical rim to the lower jaw, like that which surrounds some
tea trays, and which probably represents the tomia of the horny sheath of a
bird’s beak. At the front of the muzzle its face is sharply undulate, presenting
the appearance of vertical columns with tooth-like apices. Corresponding
tooth-like processes, of much smaller size, alternate with them from the upper
jaw. These probably are the remains of a serration of the extremital part of the
horny tomia, such as exists on the lateral portions in the lamellirostral birds.”
Cope described an undulatory surface with tooth like terminations; how-
ever, he did not make it clear whether he considered the object examined a
mold or the actual bill. He implies that the structure was attached to the lower
jaw and that only small protuberances were present on the upper. Cope prob-
ably misinterpreted the area for attachment of the bill, judging from the LACM
specimen. Versluys (1923) also believed that Cope’s analysis was in error, but
Lull and Wright ( 1942) and Ostrom ( 1961 ) both accepted Cope’s (1883)
description of a horny beak on the lower jaw. Probably the smaller, tooth like
processes described by Cope were rugosities normally present distally on most
hadrosaurian premaxillaries.
The morphology of the hadrosaurian bill was further confused by Lull
and Wright who state (1942:43), “Tn the vSenckenberg specimen [the one
1970
Hadrosaurian Dinosaur Bills
7
described by Versluys] the impression shows a beak on the upper jaw which
projected with a free lower border for about 8 cm over the ventral edge of the
premaxillae. The beak stood vertically, with a regular, undulating surface
which became smoother towards the sides. Cope described the same wavy
appearance corresponding and alternating with the tooth-like processes in the
jaw itself. ... He [Cope] also described a similar horny beak on the lower jaw.”
Cope of course had not described a wavy appearing structure occurring
on both jaws, the upper being described as small tooth like processes. Ostrom
(1961:152) apparently agreed with Lull and Wright stating, “Cope (1883),
however, described beak impressions at the anterior extremities of both the
upper and lower jaws in a specimen of A. copei {Diclonius mirabilis) Yet
neither LACM 23502 nor the equally well preserved Senckenberg specimen
give any indication of a lower bill.
Sternberg figured a bill mold in describing a specimen of C. excavatus
(NMC 8676) , but in neither the plate explanation nor in the body of the paper
did he state that the bill was attached to the lower mandible (Fig. 3). Ostrom,
however, (1961:152) states, “Further evidence of a lower beak has been found
in a third specimen {Corythosaums excavatus) , (NMC 8676) , described by
Charles M. Sternberg (1935), in the form of an impression of an incomplete
horny beak in front of and along the left side of the predentary.” As the plate
from Sternberg’s paper shows (Fig. 3) the mold is a fluted surface associated
with the lower jaw but it is separated from the predentary by a considerable
thickness of rock matrix. The presence of matrix between the surface and
predentary is evidence that the mold is not the opposing surface of a predentary
sheath. Upon examination of the plate and the small fragment in the collection
of the Canadian National Museum, it appears as though the mold is that
portion of the upper bill which originally projected over and in front of the
predentary.
Although the fluted surface has been observed on three reported species,
there is a problem regarding it. Does the fluted surface belong on the external
surface or the internal surface of the bill?
Cope, (1883) having interpreted the mold as being attached to the lower
jaw, apparently assigned the undulatory surface to the exterior of the lower
bill. On the other hand, Versluys (1923) assigned the undulatory surface to
the outer surface of the upper bill. Versluys presents excellent drawings show-
ing that the bill was indeed attached to the upper mandible and that the feature
is similar to the mold in LACM 23502. However, in reconstructing the bill he
placed the undulatory surface on the exterior. Versluys removed the posterior
portion of the bill mold and then poured a substance, probably plaster, against
the anterior part of the mold, judging from the wording as translated from
German. The resulting cast was retained but the rest of the mold was not.
Versluys interpreted this cast as being the bill. He had apparently reproduced
the posterior part of the original mold formerly occupied by matrix. Such a
restoration would erroneously cause the undulatory surface to appear on the
exterior rather than on the interior of the bill.
Sternberg (1935) did not state whether he believed the undulatory surface
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No. 193
of the beak to be on the exterior or the interior. Ostrom, however, in discussing
the specimen of Corythosaurus excavatus described by Sternberg, states,
(1961:152), “This specimen indicates the presence of small, tooth-like pro-
jections on the inner surface of the beak, which probably contributed to a firm
union of this structure with the lower jaw.” Sternberg’s plate clearly shows a
layer of matrix between the mold and predentary which is evidence against a
firm union. In addition, the surface of the predentary, although rugose, is not
appressed against the mold of the bill and the rugosities do not match the
channels in the bill mold.
The various interpretations of the relationship of the undulatory surface
to the mandibles is illustrated in Figure 4.
Function of the Hadrosaurian Bill
Almost every conceivable mode of food gathering has been proposed at
one time or another for the hadrosaurian dinosaurs. There are no existing
Figure 3. Lower jaw of Corythosaurus excavatus, NMC 8676, showing the bill mold
(H.B.), predentary (P.D.), and dentary (D). This figure is reproduced from Plate
II, Sternberg (3).
1970
Hadrosaurian Dinosaur Bills
9
MORRIS
(1969)
COPE
(1883)
B-l
NEGATIVE MOLD
INTERIOR surface
UPPER BEAK
B-2
NEGATIVE MOLD
EXTERIOR SURFACE
LOWER BEAK
B-2
NEGATIVE MOLD
EXTERIOR SURFACE
LOWER BEAK
C- POSITIVE REPLICA
UPPER BEAK
b-2
NEGATIVE MOLD
OUTER SURFACE
UPPER BEAK
Figure 4. Diagram illustrating the interpretations of the four reported hadrosaurian
bill molds. All interpretations are based upon molds preserved with specimens of
Anatosaiinis annectens except Sternberg (1935) which is based upon Corythosaurus
excavatus. Blocks labeled represent the mold as discovered. Block 1 is the posterior
part of the mold and 2 is the anterior. Blocks labeled B represent the parts of the
mold preserved after preparation. Block C in the case of the interpretation by
Versluys represents a plaster cast that was poured against B-2- Each group of blocks
is followed by the interpretation placed upon them by the authors listed. Lastly,
outline restorations are shown illustrating the interpretation by each other.
10
Contributions in Science
No. 193
reptiles that have comparable feeding or masticating features. Cope suggested
a diet of soft vegetable matter but this was based on the erroneous assumption
that the teeth, in particular those of the dentary, were only loosely attached.
Lull and Wright (1942:36) give the following picture, “The analogy now
seems to be with the moose Alces whose diet is that of a browsing animal, that
is the twigs and leaves of deciduous trees . . . and also certain conifers. In
summer it turns to the tall, lush grass in swamps or near the margins of streams
and lakes and, actually invading the water, feeds upon the leaves of the water
lily or, with head entirely submerged, on the roots and stems of aquatic
plants.”
Versluys strongly rejected the idea of a ducklike feeding habit as reported
by Abel (1912). Instead, he suggested that the structure was more beak than
bill adapted for scraping bark and leaves from small scrubs and trees or even
for the uprooting of plants.
There is only one report of stomach contents associated with a hadro-
saurian skeleton. Unidentified plant debris and recognizable conifer needles
were reported (Krausel, 1922) from the stomach cavity of A. annectens. As
pointed out by Ostrom ( 1961 ), this might indicate merely that these creatures
could feed on land plants, and it does not eliminate the possibility that they
fed on aquatic vegetation as well. Indeed, the harsher fibers of land plants
might be more readily preserved than the softer materials of aquatic plants.
In any analysis of their feeding habits, several morphological features
must be taken into account and the assumption made that all of the features
functioned together in an integrated well adapted system. The hadrosaurians
were remarkably successful. Not only are their remains relatively common but
genera have been recognized in North and South America and Eurasia. Despite
considerable disparity, particularly in the shape of the skull, there are features
related to feeding habits common to all genera. One is bulk. The probable
weight of Corythosaurus and Anatosaurus was approximately 4 tons and yet
the 30 to 40 foot hadrosaurians are not the largest recognized, being surpassed
by Edmojitosauriis and Hypacrosaurus. Small hadrosaurian remains have been
collected but it is not known if these represent a truly small taxon or are
immature individuals of the larger species. The few apparently undistorted
skulls indicate that the hadrosaurians had a constricted buccal passage formed
by the dentaries and maxillaries, and leading to the throat. In large anatosaurs,
for example, it is doubtful that particles having a cross-sectional diameter
larger than three or four inches could have passed along this narrow channel.
Hadrosaurian dental batteries are very similar, consisting of dozens of teeth
arranged in closely spaced rows. The occlusional surface is a pavement formed
of the diamond-shaped crowns of the opposing dental batteries. In addition, in
all genera three to six teeth are arranged in vertical rows and in each, only the
most dorsal are in active use. The others serve for a highly effective replace-
ment mechanism.
The bill as preserved in LACM 23502, seems to impose certain limiting
parameters on hadrosaurian feeding habits. The structure seems best adapted
for filtering. Although several workers have suggested that the bill closed
1970
Hadrosaurian Dinosaur Bills
11
against the predentary of the lower jaw forming a cropping device, three
characteristics of the bill seem to negate this function: 1 ) The bill was attached
to the anterior and dorsal segment of the premaxillaries and its free edge
extended well below the anterior dorsal margin of the predentary; 2) The very
thin space between the anterior and posterior mold suggests that the bill itself
was a relatively thin structure certainly not well adapted for foraging leaves
and branches of trees and scrubs; 3) The last feature is the fluted, undulating
inner surface of the bill which would have little or no function for foraging
or browsing, and yet seems ideally suited as a filtering device.
Hadrosaurian dinosaurs, particularly such forms as Corythosauriis and
Hypacrosaurus, are envisioned as mainly aquatic animals. A filtering device
would be very important in assuring that these large animals could ingest large
amounts of concentrated food relatively free of water in a manner similar to
that of the dabbler ducks, such as Dendrocynga, Anas and Aix. These eury-
phagus forms have laminations on the inner surface of the dorsal bill which,
when closed against the ventral bill, form an efficient filtering device. Such an
interpretation certainly makes the name “duckbill” even more fitting for the
hadrosaurian dinosaurs than previously supposed.
With their very efficient food gathering system, the hadrosaurs could take
in all manner of food including mollusks and small crustaceans as well as
plant material. In this sense they would, like many ducks, be relatively high
protein as well as carbohydrate feeders. Plant material as well as more resistant
invertebrates could be crushed between the broad occlusional surfaces of the
impressive dental battery. By lowering the anterior part of the head, excess
water as well as food particles smaller than approximately one cm would drain
out of the mouth along the channels formed by the fluted bill and the opposing
predentary. The resulting masticated material, highly concentrated, could then
be made to flow down the very small passage between the dentaries to the
stomach simply by raising the head.
The presence of the fluted bill, multiple grinding surfaces, mechanism
for continuous replacement of teeth, and a constricted passage formed from
the bones of the posterior mouth region makes such a feeding habit feasible.
Ostrom (1964) presents a very thorough compilation of prior hypotheses
and information regarding the feeding habits of hadrosaurian dinosaurs. In his
analysis of the paleoecology the conclusions reached vary only in detail from
those expressed earlier (1961 ). In summary Ostrom states (1964:995), “Past
interpretations of hadrosaurian ecology have pictured these animals as pre-
dominantly aquatic in their habits, living and feeding in swamps, lakes, or
rivers and making only occasional and limited excursions out on surrounding
lands. Anatomic evidence, however, contradicts this interpretation, indicating
instead that the hadrosaurs as highly adept bipeds were primarily terrestrial
animals. Other anatomic and botanical evidence shows them to have been
active terrestrial foragers adapted for browsing on the harsher, fibrous, or even
woody tissues of trees and shrubs.”
The bill, as reconstructed from the mold of LACM 23502, is very thin
relative to its lateral and vertical extent, and suggests a structurally weak
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Contributions in Science
No. 193
feature, too weak to be of use in browsing on fibrous or woody plants. Ostrom
is in agreement with the conclusions presented here that hadrosaurian dental
batteries together with a high degree of tooth wear must have been used to
masticate food of substantial resistance; however, he suggests that this is
evidence for foraging upon more resistant terrestrial vegetation. Such a dental
battery could be used just as effectively for mastication of invertebrates. Ostrom
(1964:987-989) suggests correctly that in the hadrosaurian localities of the
Edmonton, Belly River, Judith River, Kirkland-Fruitland, Ojo Alamo and
Lance marginal and aquatic vegetation are relatively rare in the paleoeco-
systems. This might, of course, be the result of fortuitous preservation, but even
if truly representative it could be interpreted as evidence that aquatic and
marginal vegetation was rare in the area where hadrosaurians were abundant,
a position taken by Ostrom. On the other hand it could also be evidence for
the needed addition of invertebrates in the hadrosaurian diet. One must keep
in mind that the rarity of marginal and aquatic vegetation as computed by
Ostrom is an absolute occurrence affected by all vagaries of preservation, but
in the dynamics of the ecosystem it is the relative availability of the plants
compared to the needs of the hadrosaurian that is significant.
Hadrosaurians were efficient bipeds and as Ostrom indicates (1964:990-
993) the articular surfaces of the pes and hind limbs, the large fourth tro-
chanter, ossified sacral tendons, and reduction of the carpus point strongly to
this conclusion. These characteristics when treated by themselves may well
suggest a terrestrial habitat but when the specialized nature of the bill is taken
into consideration an alternative hypothesis seems more fitting. Therefore the
conclusion that hadrosaurians were either aquatic or shallow water feeders
seems appropriate. Under these circumstances the bipedalism as well as the
specialized bill would be adaptively advantageous. In addition such a feeding
habit would give credence to the more obvious adaptive significance of the
laterally compressed tail and the probable presence of webbed hind feet, rather
than to suggest that these characteristics were for protection allowing the
otherwise defenseless hadrosaurian to retreat to the water when threatened
(Ostrom, 1964:993-995) .
Restoration of Hadrosaurian Head
Figures 5, A, B, and C are sketches illustrating the appearance of billed
hadrosaurian dinosaurs. Evidence of bills have been found in Anatosaurus
annectens and Corythosaurus excavatus but not in C. intennedius as restored
in Figure 5C. In the skull of C. intermedins, however, the distal part of the
dentary is greatly deflected while the maxillaries are not. The paratype skull of
C. intermedins (ROM 4671 ) shows this deflection while, in the same skull,
the tooth rows are occluded in normal position. If the specimen did not have a
bill on the upper mandible, then there would have been a wide, opened gape
at the anterior end of the mouth. The restoration of C. intermedins was made
as this form has the most exaggerated gape of any hadrosaurian, a condition
which suggests the presence of a bill, although the magnitude of the gape may
be partly due to accidents of preservation.
1970
Hadrosaurian Dinosaur Bills
13
Figure 5. (A) Restoration of the skull of Anatosaiirus cumectens.
Figure 5. ( B) Restoration of the skull of Corythosaurus excavatiis.
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Contributions in Science
No. 193
Figure 5. (C) Restoration of the skull of Corythosauriis intermedins.
Literature Cited
Abel, O. 1912. Grunziibe der Palaeobiologie der Wirbectiere. Stuttgard, E.
Schweizerbart’s Verlagsbuchhandlung, xvi-708.
Cope, E. C. 1883. On the characters of the skull of the Hadrosauridae. Acad. Nat.
Sci. Phila. Pr. 35:97-107.
Krausel, R. 1922. Die Kaubewegung und Lebenweise von Edmontosaiirus spec, auf
Grund der mechanischkonstruktiven Analyse, Palaeobiologica 5:409-422.
Lull, R. S. and Wright, N. E. 1942. Hadrosaurian dinosaurs of North America.
Special Paper Geol. Soc. Amer. 40:xi-242.
OsTROM, J. H. 1961. Cranial morphology of the hadrosaurian dinosaurs of North
America, Amer. Mus. Nat. Hist. Bull. 122(2) : 37-186.
. 1964. A reconsideration of the paleoecology of hadrosaurian dinosaurs,
Amer. Jour. Sci. 262:975-997.
Sternberg, C. M. 1935. Hooded hadr’osaurs of the Belly River series of the Upper
Cretaceous: A comparison with descriptions of new species, Canada, Dept, of
Mines, Nat. Mus. Bull. 77(52) : 1-37-
Versluys, J. 1923. Der Schadel des Skelettes von Trachodon annectens im Senken-
berg — Museum, Abh. Senckenb. Naturf. Ges. 38: 1-19.
Accepted for publication March 17, 1970
LOS
,1
ANGELES
' COUNTY
MUSEUM
CONTRIBUTIONS
IN SCIENCE
Number 194
July 1, 1970
ANOTHER NEW NIGHT LIZARD {XANTUSIA) FROM
DURANGO, MEXICO
By Robert G. Webb
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
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Robert J. Lavenberg
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Virginia D. Miller
Editor
ANOTHER NEW NIGHT LIZARD (XANTUSIA) FROM
DURANGO, MEXICO
By Robert G. Webb^
Abstract: Ten specimens of Xantiisia from a desert habitat
in eastern Durango, Mexico are described as a new subspecies
of the previously monotypic X. henshawi. The Durango popula-
tion of X. henshawi is known only from the type locality; indi-
viduals occur in cracks and crevices of igneous (andesite)
outcrops.
Xantiisia exlonis Webb is regarded as a subspecies of X.
vigilis.
In May 1969, Dr. Charles C. Carpenter sent me one xantusiid lizard from
the area in eastern Durango from which I had previously described Xantusia
extorris (Webb, 1965). The specimen sent by Carpenter was not X. extorris,
but resembled X. henshawi. Field work in August, 1969, yielded nine addi-
tional specimens from igneous outcrops and thereby revealed the presence of
another disjunct population of Xantusia in eastern Durango. Its morphology
suggests a subspecific relationship with X. henshawi.
I am grateful to Dr. Carpenter for sending me the specimen obtained by
his field party in April, 1969; to Robert M. Kinniburgh for field assistance,
and to the University of Texas at El Paso University Research Institute for
defraying expenses in the field in August, 1969; to Dr. John W. Wright, Los
Angeles County Museum of Natural History, and Drs. Alan E. Leviton and
Steven C. Anderson, California Academy of Sciences, for the loan of com-
parative material; and to Dr. Jerry M. Hoffer, Department of Geology, Uni-
versity of Texas at El Paso, for analysis of rock samples.
Preserved specimens are in the collections of the following institutions, to
which the abbreviations in parentheses refer in the text: The California
Academy of Sciences (CAS), the Los Angeles County Museum of Natural
History (LACM), and the University of Oklahoma (OU). The terminology
for scales follows Savage (1963).
Xantusia henshawi bolsonae, new subspecies
Type material-. Holotype: LACM 55956 (Fig. 1), obtained 6-6.5 road
miles NE Pedricena on Mexican Highway 40, at an elevation of 4400 feet,
Durango, Mexico, by Robert G. Webb and Robert M. Kinniburgh (original
field number, RGW 5365) on August 24-25, 1969. Paratypes: Eight specimens
(LACM 55957-64) with same data as holotype. One specimen (OU 32848)
^ Department of Biology, University of Texas at El Paso, El Paso, Texas 79999.
1
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Contributions in Science
No. 194
from same locality as holotype, obtained by Jan Sassaman (member of field
party headed by Charles C. Carpenter) on April 1, 1969.
Diagnosis: A subspecies of X. henshawi differing from X. h. henshawi in
having fewer longitudinal rows of dorsal scales (averaging 50 rather than 63),
seven (instead of six) supralabials, fewer femoral pores (averaging 7 rather
than 11), in mostly lacking microscopic dark peppering laterally on the belly,
in having a narrower head in relation to body length (head width/ body length
averaging 0.16 rather than 0.18), and probably in attaining a smaller (body
length 57 rather than 68 mm) maximal size. For further detail, see compari-
sons and relationships.
Description of Holotype
Female (on basis of relatively small size of femoral pores) with normal
xantusiid body proportions; snout pointed, rounded at tip; nostrils partly visible
in dorsal view; eye relatively large, slightly closer to nostril than to ear opening;
ear opening ovoid and slightly diagonal; anterior pregular fold indicated only
by smaller scales that form two transverse rows medially; posterior pregular
fold, four scale rows anterior to gular fold, indicated only by short intercalary
row of scales on either side; distinct gular fold with enlarged scales on edge
abruptly differentiated from scales of fold immediately posterior; limbs rela-
tively short, fingers and toes overlapping when limbs adpressed to body.
Measurements (in mm from dial calipers) : Snout-vent or body length,
56.8; tail (seemingly regenerated), 61.9; head length, from tip of snout to
upper anterior margin of ear opening, 12.0; maximal width of head, 8.7; depth
of head, 4.4; length of large-scaled part of head, 11.5; diameter of eye, 2.4;
distance from eye to tip of snout, 4.2; distance from tip of snout to gular fold,
9.1; length of leg, about 22.0; length of fourth toe, about 6.4; axilla-groin
length, 28.3.
Squamation: Rostral broader than high, followed in order by two nasals,
a frontonasal, two prefrontals, a median, two frontals, a hexagonal inter-
parietal that separates two parietals, and two large postparietals; interparietal
with obscure parietal eyespot; postparietals separated posteriorly by small,
wedge-shaped interpostparietal that touches nuchals; nasals, prefrontals,
frontals, and postparietals in contact; labials, loreal region and dorsal head
scales minutely pustulose (pustules confined to edges of posterior head scales) ;
nostril pierced in suture between rostral, nasal, postnasal, and first supralabial;
nostril followed on side of head by postnasal, anterior loreal, large posterior
loreal, two loreolabials, uppermost smallest, and small preocular scales of
ocular ring; postnasal touching nasal and frontonasal above and first supra-
labial below; anterior loreal touching frontonasal and prefrontal above and
first and second supralabials below; posterior loreal touching prefrontal and
first supraocular above and second (barely) and third supralabials below;
small, upper loreolabial touching first supraocular and posterior loreal; large,
1970
Another New Night Lizard
3
triangular, lower loreolabial touching posterior loreal and fourth supralabial;
scales of ocular ring small, two postoculars largest; uppermost postocular
touching fifth supraocular; five supraoculars, first touching upper loreolabial,
posterior loreal, prefrontal, median, and frontal (first supraocular divided into
two scales on right side); fifth supraocular touching uppermost postocular,
pretemporal and first temporal; pretemporal, touching fourth and fifth supra-
oculars, frontal, and parietal, followed by four (left) and five (right side)
temporals that border parietal and postparietal; seven supralabials; four (left)
and five (right side) infralabials; pretympanic scales variable in size, largest
bordering temporal and auricular scales; five to seven small auricular scales;
mental about as broad as long; three large pairs of postmentals, first pair
touching medially; anteriormost pregular scale largest, between second pair of
postmentals; 35 transverse rows (some intercalary from either side) of
pregulars; 15 rows of pregulars between first pair of postmentals and anterior-
most pregular fold; 17 rows between pregular folds, and four rows (curved
from either side) of pregulars between posteriormost pregular fold and gular
fold, excluding 13 enlarged scales on edge of gular fold; five rows of small
scales in gular fold, anteriormost scales smallest.
Small, roundish, granular scales on back, sides, and limbs; some radials
and femorals enlarged; 14 longitudinal rows of squarish ventrals at middle of
body, lateralmost scales with curved outer edges and slightly smaller than
adjacent ventrals; transverse rows of ventrals alternating along midline in some
places, so that number is 32 on right and 34 on left side; two pairs of enlarged
preanal scales, posteriormost pair largest; about 49 longitudinal dorsal scale
rows at midbody; about 1 18 transverse rows of middorsal scales (from post-
parietals to level above posterior surface of thighs); 25 lamellae under fourth
toe of left foot; seven femoral pores on each side; caudal scales rectangular,
squarish proximally underneath tail.
Coloration and pattern (recently preserved, in alcohol); Dorsal ground
color of head, body, limbs, and tail pale yellow-buff; sides of body whitish;
ventral surfaces pale yellow-white; ground colors showing uneven distribution
of minute black peppering under magnification; black peppering absent on
postorbital head stripes, and mostly lacking proximally underneath tail and on
chest and belly.
Dorsal surface having pattern of dark markings; dark brown spotlike
markings on head, largest posteriorly; brown markings (paler than elsewhere)
on limbs; dark brown-black markings on back, some markings lineate or of
irregular shape; ground color of back less extensive than markings; dark
brown-black markings on tail, tending to form ringed pattern; distinct, pale
postorbital stripes extending posteriorly from upper margin of eye along
juncture of temporals, and parietal and postparietal to small granular nuchal
scales of neck; postorbital stripe with wide (as wide as pale stripe), continuous,
brown-black lower border, but having border of two dark spots above; two.
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Contributions in Science
No. 194
pretympanic dark spots (right) or distinct, barlike mark (left side) between
eye and ear opening; dark markings on side of neck and body paler and smaller
than those dorsally; dark markings on side of head; labial region heavily suf-
fused with dark punctations.
Variation
The general morphology of the nine paratypes closely approximates
that of the holotype, except that the anterior pregular fold is indicated by a
fold of skin in OU 32848. The limbs seem to overlap more than in the
holotype with the fourth toe reaching the region of the elbow when the limbs
are adpressed to the body.
The measurements (in mm, means with ranges in parentheses) are
based on the holotype and nine paratypes. The body length of the smallest
specimen is 34.4. The body length of the largest female (holotype) is 56.8, and
of the largest male is 50.8. Severed tails that have undergone complete regen-
eration are often difficult to distinguish from tails that have never been broken.
The body length/ tail length ratio of three specimens that appear to have
original tails is 0.69 in one female with a body length of 51, and 0.74 and 0.75
in two males with body lengths of 50 and 51, respectively. The head length/
body length ratio is 0.22 (0.21-0.24); the head width/body length ratio
is 0. 1 6 (0.1 5-0. 17); the head width/ head length ratio is 0.72 (0.68-0.79) ; and
the head depth/head length ratio is 0.38 (0.35-0.42).
The squamation of the paratypes resembles that of the holotype. The size
of the small interpostparietal is often variable; the scale is either lacking or very
small, or in LACM 55957, it is especially elongate. Four specimens have
either one or two small intercalary scales at the rear of the interparietal; in
LACM 55961 the intercalary scale touches the small interpostparietal, and in
LACM 55964 the two scales nearly touch. In LACM 55963 the postparietals
are completely separated by a large interpostparietal. The nasals, prefrontals,
frontals, and postparietals touch medially in all specimens. An incomplete
longitudinal suture partly divides the posterior loreal (right side) in LACM
55961. All specimens have minutely pustulose head scales. The supraoculars
are five, in one case four (second and third supraoculars fused). There is often
one (sometimes two) intercalary scale between the first and second supra-
oculars and the adjacent median head scales; the intercalary scale seems to
represent a fragment of the first supraocular. The temporals, variable in size,
are usually five, in some cases four or six. The supralabials are usually seven,
in one case six, and the infralabials are usually five, in one case four, and in
another six. The largest postmentals are usually three, in one case four.
The transverse rows of ventrals average 32.8 (31-34), excluding the two
(usually) or three rows (each of two scales) of enlarged preanals, and small
intercalary rows, usually in the chest region. The longitudinal rows of dorsal
granular scales across the middle of the back average 49.6 (48-52), and the
1970
Another New Night Lizard
5
Figure 1. Comparison of dorsal pattern of Xantiisia h. holsonae and X. Ii. henslunvi\
four individuals of holsonae at left and two individuals of henslunvi at right; museum
numbers of specimens from left to right: LACM 55956 (holotype), LACM 55960,
LACM 55962, LACM 55964, CAS 57852, CAS 58089.
femoral pores average 6.9 (6-8). The femoral pores in females are often not
well developed being marked by shallow depressions. The average number of
transverse rows of pregulars between the anterior pregiilar fold and gular fold
is 20.8 ( 1 9-23 ) , and of the enlarged scales on the gular fold is 11.8 (10-13).
The nine paratypes resemble the holotype in general features of pattern.
All specimens have distinct pale postorbital stripes, which are accentuated by
lacking microscopic black peppering and by having blackish borders that may
be narrowly interrupted, especially the upper border. The black peppering is
usually lacking proximally on the underside of the tail, is lacking or diffusely
scattered on the underside of the head, and is mostly lacking on the chest and
belly. The pattern on the neck and back of the largest specimens consists of
small, irregularly shaped, dark blotches (generally intermeshed in jigsaw
fashion) with the total area of blotching about equal to or slightly exceeding
that of the ground color. However, ontogenetic variation is suggested since the
corresponding pattern of the smallest specimen (LACM 55964, 34 mm body
length) is obscure, consisting of small, mostly punctate, dark markings. See
Figure 1 for variation in dorsal pattern.
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Contributions in Science
No. 194
Living individuals had pale yellow to buff ground color on the back,
with the color somewhat more intense on the head. The tails were pale yellow,
whereas the limbs and sides of the body were white. The dorsal markings on
head, body, limbs and tail were dark brown-black. The ventral surfaces were
white with the belly and tail tinged with yellow, especially ventrolaterally. The
irises were orange with black reticulations concentrated anteriorly and
posteriorly.
Etymology. The name bolsonae refers to the geographic position of this
race in a southern outlier of the Bolson de Mapimi.
Comparisons and Relationships
Based on gross morphological similarities there are two distinct species of
Xantusia — vigilis and henshawi. Hitherto, X. vigilis contained six subspecies,
vigilis, gilberti, wigginsi, utahensis (Savage, 1963:35), sierrae (Bezy, 1967a),
and arizonae (Bezy, 1967b). Xantusia extorhs closely resembles X. vigilis, but
because of its geographic isolation and because of the status of the mor-
phologically similar arizonae as a distinct species at that time, extorris was
considered a distinct species (Webb, 1965). The degree of morphological dif-
ference between the subspecies of X. vigilis and between X. henshawi and
X. vigilis indicates that relationships within the genus are best expressed by
considering X. extorris as a subspecies of X. vigilis, and the isolated population
described herein as bolsonae as a subspecies of the previously monotypic
X. henshawi.
The distinct species, X. henshawi, is most readily distinguished from the
other species in the genus, X. vigilis, by 14 longitudinal rows of ventral scales
(instead of 12) , and a dorsal pattern of relatively large, black blotches (instead
of small dark dots) .
The 108 specimens examined from throughout the range of X. h. hen-
shawi do not seem to show any geographic variation, except in the number of
femoral pores, and possibly in the number of temporals. In 54 specimens from
the northernmost part of the range in the vicinity of the San Jacinto Mountains,
Riverside County, California, the femoral pores average 1 1 .9 (8-16), whereas
the femoral pores in 54 specimens from the other localities to the south average
9.7 (7-12); the combined number of femoral pores averages 10.7. The num-
ber of temporals shows a slight increase from north to south. In populations to
the north the number averages 5.2 (4-8), whereas the temporals in other
populations to the south average 5.9 (4-7); the combined average number of
temporals is 5.6. There seems to be no significant geographic difference in the
number of dorsal granules across the back (62.8, 56-71), transverse rows of
ventral scales (32.7, 30-36), enlarged scales on edge of gular fold (10.3, 7-14),
transverse rows of pregulars between the anterior pregular and gular folds
(22.6, 19-29), infralabials (5.1, 4-7), supralabials (6.2, 5-8), in the ratio,
head width/ body length (0.18, 0.16-0.28), and in dorsal pattern. The average
1970
Another New Night Lizard
7
body length of the 10 largest males is 60.4 (59-62), and of the 10 largest
females is 65.3 (63-68).
Xantusia h. bolsonae has fewer longitudinal rows of dorsal granules (49.6,
48-52) than X. h. henshawi (62.8, 56-71). The average number of femoral
pores is fewer in X. h. bolsonae (6.9, 6-8) than in X. h. henshawi (10.7, 7-16) .
The number of supralabials is usually seven in X. h. bolsonae (95%) rather
than six as in X. h. henshawi (72%; the frequency of seven supralabials is
25% ). In some specimens of X. h. henshawi it is difficult to determine which
scale is the last supralabial. Xantusia h. bolsonae seems to have a slightly
narrower head in relation to body length (head width/body length averaging
0.16, 0.15-0.17) than X. h. henshawi (0.18, 0.16-0.28). The two lateralmost
longitudinal scale rows of the belly have a few scattered dots but are mostly
devoid of microscopic black peppering in X. h. bolsonae. The ventral black
peppering is extensive in X. h. henshawi but is diffuse or absent medially and
most concentrated laterally; the two, and often the four, lateralmost scale rows
are usually liberally black-peppered in X. h. henshawi. The postorbital stripes
are distinct in all specimens of X. h. bolsonae\ when present in X. h. henshawi
the stripes are often indistinct. A comparison of the dorsal patterns on the back
is difficult to evaluate, but generally the pattern is of relatively smaller, more
often irregular-shaped, dark markings in X. h. bolsonae than in X. h. henshawi
(see comparison in Fig. 1 ). Xantusia h. bolsonae may be a smaller subspecies
(maximum size, 57) than X. h. henshawi (maximum size, 68).
Comparative material examined of X. h. henshawi (108 specimens):
Baja California: 14 specimens — CAS 1 1932, 57294-96, 57455-58; LACM
36542-44, 36555-56, 36564. California. Imperial County: 20 specimens —
CAS 22574-75, 57846-59; LACM 52668-71. Riverside County: 54 specimens
—LACM 3769-72, 3774-76, 14513, 14519, 14524-26, 14528-29, 14531-49,
22370-73, 22376, 22381-84, 22389, 22391-92, 22395, 22397, 26800, 52672-
77. San Diego County: 20 specimens — CAS 22017-18, 22566, 22581-85,
58083, 58086, 58089, 58093-94, 64298; LACM 52678-83.
Geographic Range and Habitat
The ten known specimens of X. henshawi bolsonae are from within about
a one-half mile radius, 6-6.5 road miles (Mexican Highway 40) NE Pedricena,
Durango, Mexico. The nearest localities for X. henshawi, some 800 miles to
the northwest, are in southern California and adjacent northern Baja California
(see map 72 in Stebbins, 1966). The extent of geographic range of X. h. bol-
sonae is not known. Apparent habitat was intensively examined only at the type
locality. Perhaps X. h. bolsonae is restricted, at least in Durango, to the general
desert habitat south of the Rio Nazas where X. vigilis extorris, Sceloporus
maculosus, and Cnemidophorus inornatus paululus also have restricted dis-
tributions in Durango.
The general desert habitat in this part of the Chihuahuan Desert has been
8
Contributions in Science
No. 194
Figure 2. Type locality of Xantusia lienshawi bolsonae, 6-6.5 road miles NE
Pedricena, elevation 4400 ft., Durango. Photographs taken August 25, 1969. Top:
General view of hills and low mountains showing isolated, small, andesitic out-
crops. Bottom: Outcrop of large weathered boulders of andesite.
1970
Another New Night Lizard
9
described elsewhere (Webb, 1965). The general area is of relatively low
elevation and physiographically is part of the extensive Bolson de Mapimi to
the north; the topography is mostly irregular consisting of hills and low moun-
tains. At the type locality there is a series of low hills with rock outcrops
(Fig. 2). The principal vegetation covering the low hills includes lechuguilla
(Agave lecheguilla) , a maguey (Agave sp.), a treelike yucca (Yucca sp.),
ocotillo (Fouqideria splendens) , cholla (Opuntia sp.), small prickly pear
(Opimtia sp.), and leatherplant (Jaimpha cuneata) . A shrubby legume (re-
sembling Mimosa) and a small-leaved shrub (resembling Rhus) are often
associated with the rock outcrops.
Rock outcrops in the area are not extensive and generally form either a
cluster of large boulders (Fig. 2) or a rim-rock escarpment of limited extent up
to 10 feet in height. Xantiisia h. holsonae is associated with these igneous
outcrops; the rock is best described as a porphyritic andesite (with phenocrysts
of biotite and feldspar) and may have either a gray or pink hue. Most of the
rock exposures in this desert are gray dolomitic limestone; these outcrops
frequently are extensive and have rough corrugated surfaces. Although
collecting activities were limited, X. henshawi was not found to be associated
with the limestone. The andesite weathers mostly into large chunks. The
lizards were found in both vertical and horizontal crevices. They occurred
in some horizontal crevices that had small amounts of soil. All individuals
occurred singly, and were observed only after moving large chunks of andesite
by using a large crowbar. Temperatures at 2 pm on August 24 were: rock
surface in sun, 48° C; air in sun, 37° C; and air in shade of rock crevice, 34° C.
Three species of rock-dwelling Sceloporus — S. poinsetti, S. jarrovi, and S.
maculosiis — are associated with both the andesite and limestone outcrops. One
specimen of S. poinsetti and one of X. henshawi were found in the same rock
crevice.
Xantusia h. holsonae is sympatric with X. vigilis ext orris in eastern Dur-
ango; one specimen of X. vigilis was found under a fallen yucca a few feet
below an andesitic outcrop. Both populations are disjunct from the species in
the southwestern United States and northwestern Mexico. The two Durangan
populations occur east of the continental divide, and their ranges appear to be
limited to the west by grasslands at higher elevations. The morphological
differences between the two subspecies of X. henshawi seem to be greater than
those between X. vigilis extorris and X. vigilis gilberti (the subspecies that
extorris most closely resembles).
Resumen
Dies especimenes del genero Xantusia de la habitacion del desierto en el
este de Durango, Mexico, estan descritos como una nueva subspecie del
previo monotipico X. henshawi. La poblacion de Durango de X. henshawi es
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Contributions in Science
No. 194
conocida solamente por el tipo local; individuos ocurren en gfietas y hendiduras
en las pretuverancias de andesita ignea. j ^
Xantiisia extorris es ahora considerada como una subespecie de X. vigilis.
Literature Cited
Bezy, Robert L. 1967a. A new night lizard {Xantiisia vigilis sierrae) from the
southern Sierra Nevada in California. Jour. Ariz. Acad. Sci. 4(3) : 163-167.
. 1967b. Variation, distribution, and taxonomic status of the Arizona night
lizard {Xantiisia ari zonae) . 1967(3) :653-661.
Savage, Jay M. 1963. Studies on the lizard family Xantusiidae IV. The genera. Los
Angeles Co. Mus., Contrib. Sci. 71-1-38.
Stebbins, Robert C. 1966. A field guide to western reptiles and amphibians. Hough-
ton Mifflin Co., Boston, xiv-f-279 p.
Webb, Robert G. 1965. A new night lizard (genus Xantiisia) from Durango,
Mexico. Amer. Mus. Novitates 223 1 : 1-16.
Accepted for publication January 28, 1970
LOS
ANGELES
COUNTY
MUSEUM
CONTRIBUTIONS
cX,, IN SCIENCE
Number 195
July 1, 1970
THE BIRDS OF THE KALINZU FOREST,
SOUTHWESTERN ANKOLE, UGANDA
By Herbert Friedmann and John G. Williams
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
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THE BIRDS OF THE KALINZU FOREST,
SOUTHWESTERN ANKOLE, UGANDA
By Herbert Friedmann^ and John G. Williams-
Abstract: The Kalinzu Forest, southwestern Ankole,
Uganda, had never been explored zoologically before our field
team worked there late in 1969. The forest is being cut down
rapidly, and the present report on its avifauna may well be the
final as well as the first account of its bird life. In all some 123
species are here recorded, and indicate that the Kalinzu Forest is
(or was) a meeting place for some montane forest birds with the
lowland forest species of western Uganda.
The Kalinzu Forest, a medium altitude (5000 feet) moist, evergreen
forest, in southwestern Ankole Province, western Uganda, lies a short distance
to the south of Lake George and to the east of Lake Edward, at 0° 25' S,
30° 05' E. It is close to, but 1400 feet higher than, the larger Maramagambo
Forest, which extends to the border between Ankole and Kigezi, the extreme
southwestern province of Uganda.
The Kalinzu Forest had never been explored zoologically before, and,
indeed there seem to be no Kalinzu bird records in print. The survey on which
the present report is based was carried out under sponsorship of National
Science Foundation Grant GB-7787, by a team led by Robert M. Glen and
Andrew Williams. Their stay in the Kalinzu covered just a month; their first
specimens were collected on October 27, the last on November 26, 1969. The
collection consists of 433 specimens of 121 species of birds, and also sight
records of two more species. This comprises the first, and quite possibly, the
last, collection to be made in that forest.
In the original plan submitted to National Science Foundation in 1966 for
the survey of the terrestrial vertebrates of the isolated forests of western
Uganda it was pointed out that these areas have, or may be expected to have,
faunas that are west African rather than east African in their main affiliations,
and, as such, they form the easternmost outliers of the ranges of many Congo-
lese species. It was also stressed that there is an unfortunate urgency in the
survey as most of these forests are being cut down at an alarming rate to make
room for agriculture, and that if the survey were postponed by even a few
years, it might be too late, and our potential knowledge of the original, natural
distributional limits of many of these sylvan species would be thereby re-
stricted forever.
1 Director, Los Angeles County Museum of Natural History. (Currently, Research
Associate in Ornithology, Los Angeles County Museum of Natural History.)
-Research Associate in Zoology, Los Angeles County Museum of Natural History.
1
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Contributions in Science
No. 195
The Kalinzu Forest may be taken as an extreme instance of the urgency
for collecting data before it is too late; it is an area in which there is reason to
think the survey was accomplished literally at the last opportunity. In a letter
from the field, dated November 17, 1969, Robert M. Glen wrote as follows:
“. . . The Kalinzu is probably the poorest forest we have yet collected. Vast
areas, almost the whole forest, have been worked for timber with the majority
of the remaining non-timber trees poisoned by the forestry department . . .” He
was informed by the area forest officer, “. . . who spent two days in our camp,
that we are seeing the last of the forest in its present state, since it is his inten-
tion, and that of the deparment, to utilize all standing timber as charcoal to
supply a nearby tea factory which is at present being erected. Within a rela-
tively short space of time ... all vegetation will be of a secondary nature only.”
It must be emphasized that it is not our intention to deplore what is
happening to original, natural environments, as these changes are probably
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Birds of the Kalinzu Forest
3
inevitable, and no criticism of the trend is here implied. We merely wish to
stress the need for gathering information about such areas while it is still
possible to do so.
The following catalog of Kalinzu birds may well be smaller than it would
have been if made five or ten years earlier, but we must be grateful that it was
still possible to report so many kinds of birds. The fact that the forest lies at an
altitude of 5000 feet makes it a meeting place of many lowland forest species
with a smaller number ordinarily considered highland forest birds. Among the
latter may be mentioned Pycnonotus masukensis kakamegae, Coracina caesia
pura and Erithacus aequatorialis, all of which were collected, and Tauraco
johnstoni and Nectarinia purpureiventris, which were seen, but not obtained.
The name Kalinzu Forest has previously been used for the lowest level of
the Impenetrable Forest, in southwestern Kigezi Province, as was pointed out
recently by Keith et al. ( 1969: 9-10). While it is unfortunate thate this ambigu-
ous application of the name exists, it must be clear that the Kalinzu Forest with
which this paper deals is quite a different area, in Ankole Province, some 40
miles north-north-east of the nearest edge of the Impenetrable Forest, and at an
altitude of 1500 feet higher than the lowest levels of the latter.
In the following annotated list we have kept to a minimum discussions of
different systematic treatment by different authors, as our real purpose is
merely to record what birds are known to occur (possibly by the time this paper
appears, what are known to have occurred) in the Kalinzu Forest. By and large,
we follow White (1960-1965) in nomenclature, and in the recognition or
rejection of subspecies, except where more recent revisions have been issued,
principally in the volumes completing the Check-list of Birds of the World
begun by the late J. L. Peters. The order of the passerine families follows that
of Chapin (1953, 1954).
Accipitridae
Polyboroides typus Smith
On November 26 an African harrier hawk flew into a tree in an area in
the Kalinzu Forest where several mist nets had been placed. It landed among a
troop of monkeys of several species, all of which made a tremendous racket,
but which dispersed as the hawk was shot. It proved to be an adult male, testes
not enlarged; bill black, the base and soft parts whitish; iris dark brown; feet
yellow; weight 720 grams; stomach empty.
Accipiter tachiro sparsimfasciatus (Reichenow)
One specimen, male, testes not enlarged, November 19; bill black, cere
and gape greenish yellow; basal ring gray; iris yellow; feet yellow; weight 227
grams; stomach empty. This goshawk was caught in a net in open under-
growth in an area of remaining natural forest.
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No. 195
Hieraaetus africanus (Cassin)
Cassin’s hawk-eagle is still generally considered a rare bird everywhere in
its range, chiefly because it is rare in museum collections, yet three examples
were obtained in eleven days, November 13 to 23, suggesting that in the
Kalinzu Forest it is by no means uncommon. The three examples, two males
and one female, were in non-breeding condition; bill black, the base gray, the
soft parts, cere, and gape pale yellow; ring at base pale gray; iris medium
brown to brownish cream; feet pale yellow; weight, female 1153 grams; males
938 and 1049 grams; all stomachs were empty. One of these birds was shot out
of a tall tree, and while the collector’s assistants were searching for it in the
undergrowth, another one, probably its mate, flew into the same tree and was
also collected. The other example was caught in a trap baited with a small
chicken. The Kalinzu Forest is the second area in Uganda from which this
hawk is now known, the other being the Impenetrable Forest in the extreme
southwestern part of the country. It may be worth noting that in the Im-
penetrable Forest this hawk must also be common as our collectors in 1969
noted (but did not collect) the species on 21 occasions during two months in
the field.
Keith and Twomey (1968: 537-538) considered it significant that their
single male specimen from the Impenetrable Forest had an unusually long tail,
248 mm, as compared with 225 and 234 mm in two topotypical male birds from
Gabon. Our two Kalinzu males have tails 228 and 231 mm long, the female
266 mm. Our males therefore agree in this dimension with Gabon examples
and negate any suspicion that the eastern population of the species may prove
to be a race characterized by its tail length.
Rallidae
Sarothrura elegans (Smith)
One buff-spotted crake, a female with the ovary much enlarged, was
collected on October 30, when it flew into the camp at night, apparently
attracted by one of the pressure lamps hanging in the mess tent. It had the bill
blackish gray, the lower base whitish; iris dark brown; feet dark brown; eye
skin gray; weight 44.5 grams; stomach contents grit and small beetle fragments.
Sarothrura pulchra centralis Neumann
The white-spotted crake was not obtained in the Kalinzu Forest, but on
November 8 an adult male was taken in the nearby Maramagambo Forest,
3600 feet (as compared with 5000 feet at Kalinzu). It showed only slight
gonadal swelling; bill black; iris dark brown; feet dark gray; weight 46 grams;
stomach contents insect fragments.
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Birds of the Kalinzu Forest
5
COLUMBIDAE
Cohimba unicincta Cassin
The afep pigeon is represented by an unsexed adult taken on November
21; bill gray, paling to whitish gray at the tip; iris orange-red; feet pale gray;
weight 379 grams; stomach contents seeds of small fruits. The bird was seen
feeding in a fruiting tree in an open area of the forest.
Streptopelia semitorquata (Riippell)
Two adults with enlarged gonads, one male, one female, were collected
on November 14, from a flock of several red-eyed doves feeding on the small
berry-like fruits of a tall tree. Bill dull black, the gape pinkish in the male, dull
maroon in the female; iris dark red; bare skin dull crimson to dull maroon;
feet dull pale maroon; weight 162.5 grams (female), 192.5 grams (male);
stomach contents seeds of small fruits.
Tiirtiir tympanistria (Temminck)
One male with enlarged testes, November 13; bill black; soft parts dark
dull maroon; iris very dark brown; bare skin dark gray; feet dull maroon;
weight 75 grams; stomach contents seeds and grit.
Treron aiitralis gihherifrons (Madarasz)
This green pigeon was met with twice, on Novemeber 12 and 16. On each
day an adult male was taken, the second of which had large testes; bill pale
gray; soft parts and cere dull orange; iris bright metallic sky-blue; feet orange-
red; weight 225 and 226.5 grams; stomach contents small fruits.
Musophagidae
Taiiraco schutti emini (Reichenow)
On October 29 one male of this black-billed lourie was collected in a
patch of secondary forest which had recently been felled; testes not enlarged;
bill black, the lower base red; iris dark brown; feet black; bare eye skin red;
weight 267 grams; stomach contents fruit seeds.
Tauraco johnstoni (Sharpe)
Robert Glen saw, but was unable to collect, what “. . . was certainly a
Ruwenzori turaco . . .”, which would establish a new minimal altitude for this
montane bird. A specimen would not only have been welcome to corroborate
the identification, but also to determine which of the two races occurs in the
Kalinzu Forest.
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No. 195
CUCULIDAE
Ciiculus clamosus gabonensis Lafresnaye
An almost adult female in the pale-bellied ''mabirae'" type of plumage of
this forest cuckoo was taken on November 25; maxilla black; mandible slightly
grayish; iris dark brown; feet white with pale gray scales; weight 87 grams;
stomach contents hairy caterpillars. This specimen has the chin, throat and
upper breast pale rufescent barred fairly broadly with blackish; the lower
breast and abdomen whitish cream with only a small number of incomplete
fuscous cross bars, especially toward the sides of the breast and abdomen, the
under tail coverts unmarked cream.
Chrysococcyx cupreus cupreus (Shaw)
The emerald cuckoo undoubtedly occurs in every forest in Uganda.
In the Kalinzu an adult female with an enlarged ovary was taken on November
2; maxilla blackish gray; mandible below greenish; iris dark brown; eye ring
apple-green; feet blue-gray; weight 38 grams; stomach contents caterpillars. A
few of the abdominal feathers have a pronounced yellowish wash in their pale
areas — a slight variation in the direction of the richly yellow abdomen of the
male.
Ceuthf nochares aereus aereus (Viellot)
This cuckoo is represented in the collection by two males, one with large,
the other with small, gonads, November 14 and 22; bill lemon-yellow; iris
chestnut-brown to dark red; feet black; bare skin behind eye blue, in front of
eye apple-green; weight 56 and 66 grams; stomach contents caterpillars, grass-
hoppers and beetles. These birds were found feeding in creeper-covered bush
in mixed, exploited forest.
Strigidae
Glaucidium tephronotum medje Chapin
Three examples, two males and one female, of this owl were taken
November 1, 22, and 26, respectively. It follows that the species must be fairly
common in the Kalinzu Forest, and in the forests of western Uganda generally,
as six more specimens from the Bwamba, Kibale, and Impenetrable Forests
are now in the Los Angeles County Museum of Natural History. One of the
present individuals was kept alive for some days and was heard to give a rather
mournful sounding, whistled poop several times in rapid succession. This owl
was caught in mist nets in the undergrowth in both primary forest and in areas
where the trees had been heavily exploited. All three examples were in non-
breeding state; bill pale greenish yellow to dull apple-green; cere yellow to
greenish yellow; iris bright yellow (with gray flecks in one of the males); feet
1970
Birds of the Kalinzu Forest
7
yellow; eyelids olive; weight 83.5, 94, and 103 grams (the female the heaviest) ;
stomach contents insect fragments and one green beetle. There is much
individual variation among these owls, especially in the amount of rufescence
on the underparts, and in the degree of difference between the dark grayish of
the crown and occiput and the brown of the back, the distinction being more
marked in some individuals than in others. As we pointed out (Friedmann and
Williams, 1968: 15) some of the dorsal variability is probably due to wear as
the brown feathers of the back have dark grayish tips when fresh, and as these
are abraded, the brown color becomes correspondingly more prominent.
Because of the scarcity of specimens of this owl in most museum collec-
tions, the measurements of the nine examples now in Los Angeles form a useful
addition to those discussed by Keith and Twomey (1968 : 540-541 ) . Our eight
males have wing lengths of from 1 12 to 119 mm; tail lengths 80 to 88 mm; our
one female, wing, 115 mm, tail, 85 mm.
Apodidae
A pus affinis affinis (Gray)
This swift is fairly common in the Kalinzu area, where three males were
taken November 1, 4, and 6. Two of them had enlarged gonads, the third
showed only slight testicular swelling; bill black; iris dark brown; feet black;
weight 25 to 28 grams; stomach contents flying insects. The collectors noted
that this species was nesting under the eaves of the forestry department rest
house at the edge of the forest.
Chaetura sabini Gray
Sabine’s spine-tailed swift appears to be commoner and more widely
distributed in western Uganda than was formerly thought. Five females were
taken in the Kalinzu area October 31 to November 10, and others have been
reported or collected in the Bwamba, Budongo, and Kibale Forests as well. Of
our present examples only one showed much gonadal enlargement, the others
had small ovaries; bill black; iris dark brown; feet maroon to maroon-gray;
toes blackish; weight 16.5 to 18 grams; stomach contents small flying insects.
These specimens were shot from groups flying over the swampy forest together
with other swifts and with black rough-winged swallows {Psalidoprocne) .
In an earlier paper we (Friedmann and Williams, 1968 : 16) noted that
the wing length of our sole Kibale specimen was only 1 10.9 mm, which seemed
to uphold the validity of the supposedly short winged race ogowensis. Our
present five examples have wing lengths varying from 1 14 to 126 mm, and they,
in turn, make it more probable that ogowensis is not a recognizable subspecies.
The use of a binomial for this swift thus appears to correctly reflect our
knowledge of its variability.
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Contributions in Science
No. 195
Trogonidae
Apaloderma narina brachyurum Chapin
Two male trogons, taken on the same day, November 25, differ in that
in one the feathers of the rump and the upper tail coverts are somewhat bluer,
less greenish than in the other; one had enlarged testes, the other showed only
slight gonadal swelling; bill greenish gray becoming yellow at the base; iris
chestnut-brown; feet pale flesh-brown; bare skin around eye apple-green;
weight 68 and 70 grams, respectively; stomach contents caterpillars, one butter-
fly and other insect fragments. The birds were flying after insects about 20 feet
from the ground, but the presence of caterpillars in their stomachs is evidence
that not all of their food is caught on the wing.
Alcedinidae
Halcyon malimhica malimhica (Shaw)
Two specimens of this kingfisher were taken on November 18 and 19, one
of each sex; the female with large ovary, the male with slightly swollen testes;
bill crimson-red, the mandible and the lower base blackish; iris dark brown;
feet orange, scales brownish; weight 89 and 92.5 grams, respectively; stomach
of the female held a blue-green tree frog while that of the male was empty.
Meropidae
M crops gularis australis (Reichenow)
One male and two females of this bee-eater were taken October 28 to
November 4, all in non-breeding state; bill black; iris red; feet black; weight 30
to 34 grams; stomach contents beetles and one wasp. On November 8 another
male was taken in the nearby Maramagambo Forest, 3600 feet.
CORACIIDAE
Eurystonius gularis neglectus Neumann
Two male rollers with small gonads and one female with enlarged ovary
collected November 10 to 16. The males are in a changing state of plumage
with a mixture of blue and maroon-brown on the abdomen; bill yellow; iris
dark brown; feet yellowish brown, the toes darker; weight 99 (female) to
1 17.5 grams (males) ; stomach contents termites and flying black ants. These
birds were seen catching small insects on the wing in the open, exploited forest.
Phoeniculidae
Phoeniciilus hollei jacksoni (Sharpe)
Two wood hoopoes were taken October 30 and November 4, the former
Linsexed, the latter a male; bill orange-red to deep pinkish crimson; iris dark
brown; feet red; weight 48 and 55 grams; stomach contents beetle larvae and
other insect remains.
1970
Birds of the Kalinzu Forest
9
Bucerotidae
Bycanistes siihcylindriciis suhquadratiis Cabanis
One male hornbill in breeding condition, taken November 14; bill dull
scaly brown, with base and half of casque pale gray; iris rich reddish brown;
bare skin dull black; feet scaly black; weight “over 1600 grams”; stomach
contents fruits. The collectors noted many more single birds (males?) than
pairs, which seemed to suggest that the females were already holed up in
their nests.
Capitonidae
Lyhius hirsutus ansorgei (Shelley)
The hairy-chested barbet is fairly common in the Kalinzu forest. Three
examples were collected October 31 to November 22; all with slight gonadal
enlargement; bill black; iris dark brown; feet black; weight 45 to 49 grams;
stomach contents berries and fruit seeds. These specimens were collected in the
undergrowth in an unspoiled area of natural forest.
Gyninobucco honapartei cinereiceps Sharpe
The brown barbet was numerous and four specimens were taken Novem-
ber 13 to 19, three of them with large gonads, one with small ones; bill blackish
horn; iris cream to yellowish white; feet black; weight 62 to 67 grams; stomach
contents green fruits. A female, November 19, was shot at a nesting (?) hole
in a dead tree.
Buccanodon duchaillui duchaillui (Cassin)
On November 10 three adults, two females, 1 unsexed, were collected;
this was the only time the field team met with this barbet. The birds were in
non-breeding state; bill black; iris dark brown; feet dark gray; weight 33 to
36.5 grams; stomach contents small fruit seeds. All were netted in the under-
growth in a heavily exploited part of the forest.
Pogoniidus scolopaceus aloysii (Salvadori)
This little tinker bird was common in the Kalinzu. Specimens were
obtained October 28, November 14, and November 20, the first and the last
were in breeding condition; bill black; iris cream to grayish yellow; feet dark
gray; weight 15.5 to 17 grams; stomach contents insect fragments and small
fruit seeds. Found in the undergrowth in exploited forest areas.
Pogoniulus hdineatus mfiiinbiri (Ogilvie-Grant)
The commonest tinker bird in Kalinzu; seven examples, October 29 to
November 19; non-breeding condition; bill black; iris dark brown; feet black-
ish; weight 10.5 to 13.5 grams; stomach contents fruit seeds and insect
fragments.
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Contributions in Science
No. 195
Trachyphonus purpuratus elgonensis Sharpe
This brightly colored species was one of the commonest barbets in the
Kalinzu. Large numbers were caught in the mist nets (and released) where
they were catching flying termites as they escaped from the ground. Five
specimens were taken October 30 to November 12; four of them with small,
one with large, gonads; bill lemon-yellow; iris dark red; feet blackish gray;
weight 72 to 85 grams; stomach contents fruit and fruit seeds.
Indicatoridae
Indicator exilis pachyrhynchus (Heuglin)
The least honey guide is represented in the collection by one adult male,
with enlarged testes, taken October 30; bill black, the lower base whitish; iris
dark brown; eye skin dark gray; feet olive-gray; weight 21.5 grams; stomach
entirely filled with beeswax. It was collected when it came to a spot where a
number of sunbirds were excitedly mobbing a snake. Although this was the
only example noted, there is every reason to assume that this honey guide is no
less numerous in the Kalinzu than it is in the other western Uganda forests.
The local abundance of small barbets, of at least two species of Pogoniulus,
would assure an ample supply of hosts for this brood parasite as well as for
the next species.
Indicator willcocksi willcocksi Alexander
That this forest honey guide occurs in the Kalinzu is of interest in round-
ing out our concept of its range in western Uganda, where it is already known
from the Impenetrable and the Kibale Forests. It does not occur in the Bwamba
woodlands, which are at too low an altitude, as its range seems to be above
4200 feet. Three specimens, one male and two females, were taken in the
Kalinzu on October 27, November 13, and November 23, respectively; all
were in non-breeding state; bill black, the lower base pale pinkish gray; iris
dark brown; feet olive-gray; weight 16 and 17 grams in the females, 20.5 grams
in the case of the male; stomach contents entirely beeswax in two, beeswax and
small insect fragments in the third.
In an earlier paper we (Friedmann and Williams, 1968: 17-18) described
seven specimens from the Impenetrable and Kibale Forests as being slightly
paler and with more of a yellowish tone below and less heavily streaked on the
crown than were the available Cameroon and Congo specimens, and we
raised the possibility of an eastern subspecies of /. willcocksi. The Kalinzu
birds merely add to the variability, or the inconstancy of coloration as noted
in the western Uganda population, and thereby tend to negate the idea of an
eastern race of this honey guide. Furthermore, as we pointed out in the earlier
paper, it would be necessary to see specimens from localities all across the
Congo before it would be possible to ascertain where, if anywhere, such a
division might be made.
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Birds of the Kalinzu Forest
11
PiCIDAE
Campethera nivosa herberti (Alexander)
Inasmuch as the buff-spotted woodpecker is known from many forested
areas throughout Uganda, it is no surprise to find it in the Kalinzu as well. On
November 22, two males were collected as they were watched searching for
insects on a dead branch sticking up out of the undergrowth in an undisturbed
area of primary forest. Both birds showed only slight gonadal enlargement;
bill horn black becoming brown on the basal part of the mandible; iris dark
brown; bare eye skin olive-gray; feet olive; weight 35 and 39 grams; stomach
contents insect fragments.
Campethera tullbergi taeniolaema Reichenow and Neumann
The fine-banded woodpecker is a very common bird in the Kalinzu Forest.
Our collectors obtained ten specimens from October 30 to November 8, all in
non-breeding condition; bill horn-black to dark gray, the basal part of the
maxilla pale gray; iris bright crimson to deep chestnut; eye skin gray; feet olive;
weight 48 to 55 grams; stomach contents ants in all cases. These woodpeckers
were found feeding along long stems in the undergrowth in areas where the
forest had been heavily exploited.
Dendropicos fuscescens lepidus (Cabanis and Heine)
The cardinal woodpecker was met with on November 23 when one adult
male was collected as it was pecking on one of the upper branches of a dead
forest tree that had been poisoned by the forestry department as part of its
scheme to remove the remnants of the Kalinzu to make way for agriculture. It
is not known if the methods used to poison non-timber trees affects the edibility
of the insects still found on the latter, and, so, in turn, the birds that feed on
them; but the question is a real one. The specimen obtained had the bill dark
horn-gray above, paler toward the base on the mandible; iris red-brown; eye
skin dark gray; feet olive; weight 25 grams; stomach contents beetles and beetle
larvae.
Mesopicos xantholophus (Hargitt)
The yellow-crested woodpecker occurs in many Ugandan forests and its
presence in the Kalinzu is therefore not surprising. One male and two unsexed
specimens were collected, October 31 to November 25; bill dark horn-gray,
paler on the mandible; iris red-brown to deep crimson; eye skin dark gray; feet
olive-gray; weight 57.5 to 66 grams; stomach contents beetle larvae. The birds
were noted feeding extensively on spurge stems (Neobutonia) , which appar-
ently house a great many beetles. Robert Glen wrote that he had observed
many species of beetles, including even the very large goliath beetle, frequent-
ing these spurge stems.
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Mesopicos elliotii elliotii (Cassin)
Elliot’s woodpecker is generally considered to be fairly uncommon in the
forests of Uganda, but it must be common in Kalinzu, as four specimens were
obtained in less than a week, October 30 to November 4; all in non-breeding
state; bill blackish horn, paling to gray on the mandible; iris dark brown to
deep crimson; eye skin brown to dark gray; feet olive; weight 34.5 to 38 grams;
stomach contents ants, caterpillars and beetle larvae.
Eurylaemidae
Smithornis capensis meinertzhageni van Someren
The African broadbill is common in Kalinzu; five specimens, four males,
one female, all in non-breeding state, were collected, November 8 to 22; maxilla
black; mandible whitish gray to pale pink; iris dark brown; feet pale olive-
yellow to olive; weight 23 to 27.5 grams; stomach contents insects, mainly
small beetles. These birds were all netted in the dense undergrowth in natural
forest areas.
Hirundinidae
Hirundo rustica rustica Linnaeus
One male example of the European barn swallow, a winter visitor to
Uganda, was collected on November 16; weight 20 grams; stomach contents
small flies.
Psalidoprocne pristoptera ruwenzori Chapin
One male, with large testes, collected November 14, has a short tail (77.5
mm) well within the variational range of ruwenzori; bill black; iris dark brown;
feet dark brown; weight 1 1 .5 grams; stomach contents insect fragments.
Motacillidae
Motacilla drier ea drier ea Tunstall
One female gray wagtail was taken November 16; ovary small; bill
blackish horn; iris dark brown; feet pale brown; weight 16.5 grams; stomach
contents insect fragments.
Motacilla capensis wellsi Ogilvie-Grant
Common in the open areas about the Kalinzu Forest. From October 29
to November 1 3 four examples were secured, two males with large gonads,
one female with a small ovary, and one of unrecorded sex; bill black; iris dark
brown; feet blackish gray; weight 19 to 25.5 grams; stomach contents insect
remains.
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Birds of the Kalinzu Forest
13
Timaliidae
Trichastoma fulvescens ugandae (van Someren)
This brown babbler would seem to be relatively uncommon in Kalinzu,
judged by the fact that only a single example was obtained, as compared with
four or five of T. rufipennis and T. alhipectiis. One male in non-breeding state
was obtained November 4; maxilla black; mandible gray; iris brown; feet dark
gray; stomach contents beetles and small snails. The bird was caught in a mist
net in the dense undergrowth in an area of heavily exploited forest.
Trichastoma pyrrhoptera pyrrhoptera (Reichenow and Neumann)
One specimen, a non-breeding male, November 8; maxilla black; man-
dible pale gray; iris chestnut; feet gray; weight 25 grams; stomach contents
beetles and small snails.
Trichastoma rufipennis rufipennis Sharpe
Common in the Kalinzu Forest, where four specimens were caught in mist
nets placed in the undergrowth of natural, undisturbed forest, November 1 3
and 14; all were in non-breeding state; maxilla black; mandible blue-gray;
gape yellow; iris dark brown; feet dark gray, the toes paler; weight 21 to 26
grams; stomach contents insect fragments and small snails.
Trichastoma albipectus (Reichenow)
Like the previous species this one is also numerous in the Kalinzu Forest,
where it occurs with the very similar T. rufipennis. Five specimens, four males,
one of which had enlarged gonads, and one female, were taken November 1
to 23; maxilla black; mandible pale gray; gape yellow; iris brown to reddish
brown; feet opaque pale gray in some cases with a pinkish or a pale violet
tinge; weight 28 to 32 grams; stomach contents small beetle fragments and
parts of one millipede. This species was caught in nets in both natural and
exploited forest.
Pycnonotidae
Pycnonotus barbatiis tricolor (Hartlaub)
This ubiquitous bulbul is represented in the collection by an adult male
in non-breeding state, taken on November 19 outside the forest; bill black; iris
dark brown; feet black; weight 45 grams; stomach contents fruits.
Pycnonotus masukensis kakemegae (Sharpe)
The Kalinzu Forest would seem to be at the lower level of the range of
this bulbul, which is chiefly a bird of the highland forests. Between October
28 and November 21 four males and one female were collected; gonads ranging
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No. 195
from small to much enlarged; bill blackish above, dark gray below; iris red-
brown; feet blue-gray; weight 25 to 27 grams; stomach contents insect frag-
ments, and, in one case, small fruits. These birds were collected in natural or
little exploited forest areas. The Kalinzu is the only area from western Uganda
other than the Impenetrable Forest from which we have received examples of
this species; it might have been expected in the Kibale Forest, but our collectors
did not meet it there.
Pycnonotus v ire ns holochlorus (van Someren)
One female in non-breeding condition, November 4; bill brownish black;
iris dark gray; feet yellow-brown; weight 21.5 grams; stomach empty; netted
in undergrowth in heavily exploited forest.
Pycnonotus gracilis ugandae (van Someren)
Common in the Kalinzu; six specimens, four males and two females, all
with small gonads, were taken October 30 to November 22; bill black; iris
dark brown; feet olive-gray; weight 19 to 22 grams; stomach contents fruit
seeds, insects and insect larvae.
Pycnonotus curvirostris curvirostris (Cassin)
Another common Kalinzu bird; seven examples October 29 to November
24; gonads varying from small to large; bill dark horn gray to blackish, more
yellowish brown at tip; iris chestnut-brown to red-brown; feet olive; weight
23.5 to 26 grams; stomach contents fruit seeds and insect fragments.
Pycnonotus latirostris eugenius (Reichenow)
As might have been expected this bulbul proved to be numerous in the
Kalinzu Forest. Seven specimens were obtained October 28 to November 14,
three of them with enlarged gonads, four with small ones; bill black, tomium
and gape yellow; iris dark gray-brown; feet brownish yellow; weight 25.5 to
29 grams; stomach contents fruit seeds and insect remains. The yellow-
whiskered bulbul was found (netted) chiefly in undergrowth in heavily
exploited areas of the forest.
Pycnonotus gracilirostris congensis (Reichenow)
Very common. Seven examples were taken in the Kalinzu Forest and two
more in the nearby Maramagambo Forest, October 28 to November 19. A
long series of this bulbul from many of the western Uganda forests suggests the
use of the name congensis rather than the recognition of chagwensis. In this
allocation we follow White (1962; 78) rather than Rand (1960: 256) . The
birds were mostly in non-breeding state, but one female, November 2, had an
enlarged ovary; bill black; iris brick-red to crimson; feet black; weight 29 to 37
1970
Birds of the Kalinzu Forest
15
grams; stomach contents insect fragments entirely. These slender-billed bulbuls
were noted feeding in the tree tops in the ironwood climax forest.
Baeopogon indicator indicator ( Verreaux)
One male in non-breeding state, October 28; bill black; iris white; feet
dark gray; weight 47 grams; stomach contents ants; collected in a heavily
exploited area of the forest.
Phyllastrephus hypochlorus (Jackson)
Apparently fairly common in the Kalinzu Forest; three specimens, No-
vember 8 to 18; all with small gonads; bill black above, grayish below with
whitish sides at the tip; iris brown; feet gray; weight 21 to 28 grams; stomach
contents insect fragments.
Phyllastrephus fischeri sucosus Reichenow
Very common in the Kalinzu; seven examples were collected, October 28
to November 22; gonads ranging from small to much enlarged; bill blackish
horn to black, the tomium and mandible paler; iris gray to yellow-gray to pale
brown; feet gray; weight 20 to 27 grams; stomach contents ants and other
insect fragments. These birds were netted in the undergrowth in heavily
exploited areas of the forest.
Phyllastrephus xavieri xavieri (Oustalet)
Xavier’s bulbul was found to be common in the Kalinzu, where four
specimens were obtained October 31 to November 24; all with small gonads;
bill black, tomium and gape yellow, mandible gray at base; iris gray to grayish
cream; feet dark gray, the toes paler; weight 25.5 to 27 grams; stomach con-
tents insect remains. These birds were caught in nets in the undergrowth in the
heavily exploited areas of the forest.
Bleda syndactyla woosnami Ogilvie-Grant
Very common in the Kalinzu; seven specimens taken November 4 to 2 1 ;
all but one with small gonads; bill black above, pale gray below; iris dark red to
reddish brown; feet brownish gray to pinkish gray; weight 40 to 49 grams;
stomach contents beetles and other insect fragments.
Criniger calurus emini Chapin
I'his bulbul was found to be very common and seven specimens were
obtained November 4 to 22; gonads ranging from small to much enlarged; bill
black above, blue-gray below and with the tomium whitish in some but not in
all individuals; iris chestnut to crimson; feet gray to dark blue-gray; weight 23
to 31 grams; stomach contents beetles and other insect fragments. All these
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No. 195
specimens are definitely C. calurus and not C. ndussumensis, as all have broad
bills and lack any grayish on the lores.
Campephagidae
Coracina caesia pura (Sharpe)
The Kalinzu Forest (5000 feet) must be close to the lower level of the
range of this cuckoo-shrike, which does not occur in the lower Bugoma,
Bwamba or Kibale Forests. Two females, both in non-breeding condition, were
taken November 20 and 22. Both were in late stages of molt and both were in
nearly complete adult plumage; bill black; iris dark brown; feet black; weight
49 and 50 grams; stomach contents caterpillars and other insect remains.
Campephaga phoenicea petiti Oustalet
One adult male, testes much enlarged, taken on November 25; bill black;
gape rich yellow; iris dark brown; feet black; weight 34 grams; stomach con-
tents caterpillars. The yellow gape of the male is conspicuous in the field.
Muscicapidae
Muscicapa acliista pwnila (Reichenow)
One male of this little flycatcher, testes not enlarged, was taken Novem-
ber 7; bill black above, the basal half of mandible cream; iris dark brown; feet
black; weight 7.5 grams; stomach contents beetles.
Muscicapa griseigularis (Jackson)
This flycatcher is fairly common in the Kalinzu Forest; three males and
one female, all with small gonads, were collected November 4 to 18; maxilla
black; mandible gray; iris dark brown; feet gray; wegiht 10.5 to 13 grams;
stomach contents small beetle and other insect fragments. These birds were
caught in mist nets in the undergrowth of both natural and exploited areas of
the forest.
Muscicapa comitata (Cassin)
Another common bird, of which four specimens were taken November
2 to 14, two of each sex, all with small gonads; bill black; iris dark brown; feet
black; weight 14 to 15 grams; stomach contents small beetle fragments.
Artomyias fuliginosa J. and E. Verreaux
This dusky brown flycatcher was found to be very numerous in the
Kalinzu; six specimens were collected November 1 to 13; four with enlarged
gonads, the other two unsexed because of damage; bill black; iris dark brown;
feet black; weight 17 to 19 grams; stomach contents beetle fragments. One of
the unsexed birds is subadult, molting into adult plumage, but still has the
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Birds of the Kalinzu Forest
17
upper parts and the breast dotted with white, the abdominal feathers with buffy
grayish white ends, and also with the gape whitish. The birds perch on bare
branches of tall forest trees from which they make frequent sorties to capture
insects. They resemble martins when on the wing.
Fraser ia ocreata ocreata (Strickland)
Common in the Kalinzu Forest; four specimens taken October 30 to
November 5; two of each sex, all in non-breeding state; bill black; iris dark
brown; feet blackish; weight 37.5 to 40 grams; stomach contents beetle frag-
ments. Three of these forest flycatchers were noted as being collected while
they were catching termites on the wing, but the fact remains that only beetles
were found in their stomachs!
Megahyas fiammulata aequatorialis Jackson
Another common Kalinzu species; six specimens taken, three females,
two males, October 28 to November 6; all with small gonads; bill black; iris
red; feet dull brownish pink to maroon; weight 27.5 to 31 grams; stomach
contents beetle fragments, and, in one case, lepidopteran eggs. These birds were
seen feeding in trees and were also netted in the undergrowth in exploited parts
of the forest. On November 8 an immature male, molting into adult plumage,
was collected in the nearby, but lower (3600 feet), Maramagambo Forest.
Bias musicus musicus ( Vieillot)
Fairly common flycatcher. One of each sex collected, October 31 and
November 9 in the Kalinzu, and two others, one of each sex, November 8 in
the Maramagambo Forest; bill black; iris yellow; feet pale yellow; weight 21
to 22 grams; stomach contents beetles and other insect fragments, including
flying insects.
Batis molitor (Hahn and Kuster)
This flycatcher was met with only once, on November 6, when a pair
were seen running along branches of low trees in search of food. Both birds
were in non-breeding state; bill black; iris pale green, lower ring yellow; feet
black; weight 11 and 12 grams; stomach contents small beetles.
Platysteira castanea castanea Fraser
Very common in the Kalinzu Forest; seven examples were taken, October
28 to November 25; gonads large in one, small in the others; bill black; iris
chestnut to dark maroon; feet dull maroon; weight 12 to 16 grams; stomach
contents insect, largely beetle, fragments. The eye wattles in the females seem
to be slightly darker in life than those of the males. These wattle-eyes were
caught largely in nets in the undergrowth, close to the ground, in relatively
unmolested areas of natural forest.
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No. 195
Platysteira blissetti jamesoni (Sharpe)
One male, with enlarged testes, taken October 28 and one female, with a
small ovary, November 13; bill black; iris dark maroon; feet maroon to dull
violet-maroon; weight 10.5 to 11 grams; stomach contents insect fragments,
beetles in one case. Netted near a small stream in natural forest.
Platysteira concreta graueri (Hartert)
This wattle-eye must be very numerous in the Kalinzu Forest, as six
specimens were collected November 4 to 21, three of each sex; one of the
females with much gonadal enlargement, the other individuals with small
gonads. The males illustrate the plumage polymorphism of this species, de-
scribed by Prigogine (1969), as one of them has the entire underparts bright
yellow while the other two have the whole area bright chestnut; the three
females have the chin, throat and breast bright chestnut, the abdomen and
under tail coverts yellow, the two being sharply demarcated; bill black; iris very
dark maroon-brown with a white inner ring; wattles pale yellow-green; feet
dark blue-gray; weight 12.5 to 15 grams; stomach contents small insect, largely
beetle, fragments. Although this wattle- eye has been reported from the Semliki
Valley on the west side of the Congo-Uganda border, and from Kakamega in
western Kenya, we are not aware of any Uganda locality other than Bwamba
and Kalinzu Forests.
Trochocercus nigromitratus (Reichenow)
Two specimens, October 28 and November 13, represent this flycatcher;
bill black; iris dark brown; feet gray to dark blue-gray; weight 10 to 11.5
grams; stomach contents beetle and other insect fragments.
Terpsiphone viridis ferreti (Guerin)
This paradise flycatcher is very com.mon in the Kalinzu Forest; thirteen
specimens were obtained October 28 to November 25. The series illustrates
the polymorphic nature of the species, two of the females have the entire upper
parts, wings and tail dark bluish gray; all the others are reddish brown backed
birds but with much variation in the darkness of the blue-gray underparts,
which range from fairly pale blue-gray to almost blackish blue. The only males
with elongated central rectrices also had enlarged testes; bill dark blue-gray to
pale cobalt-blue, the tip and tomium blackish; iris dark brown; feet blue-gray
to dark blue-gray; eye skin dark blue-gray; weight 12 to 16.5 grams (one female
marked 22 grams) ; stomach contents small beetle and other insect fragments.
One of the dark bluish gray backed females was seen together with, and
apparently following, a red backed, long tailed male (also collected), a good
indication that what we are dealing with here is polymorphism and not a
confusion of two different taxa.
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Birds of the Kalinzu Forest
19
Turdidae
Alethe poUocephala carruthersi Ogilvie-Grant
This thrush was common in the Kalinzu Forest; five specimens were
taken November 1 to 1 3; gonads small in four, large in one; bill black; iris deep
chestnut to dark “soft” brown; feet opaque gray, with or without a pinkish
tone; weight 25 to 34 grams; stomach contents small beetle and other insect
fragments, one snail. The brown-chested alethe was found in both unspoiled
and in heavily exploited parts of the forest.
Erithacus aequatorialis (Jackson)
Fairly common; four specimens taken, October 31 to November 5; all in
non-breeding state; bill black; iris dark brown; feet silvery gray to pale brown-
ish gray; toes yellowish; weight 14 to 19.5 grams; stomach contents insect
fragments, mainly beetles. These akalats were caught in nets in the under-
growth in heavily exploited areas of the forest.
Sphenoeacus mentalis mentalis (Fraser)
One specimen, a female with an enlarged ovary, was taken November 19;
maxilla black above, the tomium and mandible pale gray; iris pale brown; feet
pale gray; weight 36 grams; stomach contents insect fragments. The bird was
shot while it was skulking in scrub grassland at the edge of the forest.
Neocossyphus poensis praepectoralis Jackson
Abundant in the Kalinzu Forest; nine examples were collected November
4 to 21; one female with an enlarged ovary, the other specimens in non-breed-
ing state; bill black; iris dark brown; feet whitish flesh to flesh pink; weight
45 to 60 grams; stomach contents ants and small beetles. These birds were
taken in mist nets in the undergrowth and also shot high up in the forest trees.
Stizorhina fraseri vulpina Reichenow
Fairly common in the Kalinzu; three specimens were taken October 29
to November 8; all with non-breeding gonads; bill black, the base brownish in
a female; iris dark brown; feet pale brown to pinkish brown; weight 37 to 39
grams; stomach contents entirely beetles.
Sylviidae
Schoenicola platyura alexinae (Heuglin)
The great-tailed warbler was met with once, on November 12, when two
males were collected, one with small and one with large testes; bill black above,
pale gray below; iris brown; feet gray with a brownish wash; weight 16.5 and
17 grams; stomach contents small insect fragments.
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No. 195
Prinia leucopogon reichenowi (Hartlaub)
The white-chinned prinia was found to be numerous in the Kalinzu
where six specimens were collected November 1 to 10; gonads ranging from
small to large; bill black; iris chestnut; feet pale pinkish brown; weight 11.5 to
13.5 grams; stomach contents insect fragments. Most of these examples were
netted in the undergrowth in heavily exploited areas of the forest.
Apalis jacksoni jacksoni Sharpe
One specimen, October 28, a male in non-breeding state; bill black; iris
brown; feet flesh-brown; weight 10 grams; stomach contents small flies and
small insect larvae.
Apalis rufogularis nigrescens (Jackson)
This warbler is represented in the collection by two specimens, collected
November 11 and 16; one a male, the other with sex unrecorded; bill black;
iris chestnut to pale brown; feet dull pinkish brown; toes paler; weight 9 and
10 grams; stomach contents small insects and insect larvae. In their dark
brownish dorsal coloration these specimens agree with nigrescens from the
Kibale, Bugoma and Bwamba Forests, and not with the more grayish backed
kigezi of the Impenetrable Forest.
Bathmocercus cerviniventris vulpinus Reichenow
Very common; seven specimens, October 30 to November 21; gonads
small to much enlarged; bill black; iris dark brown; feet dark blue-gray; weight
15.5 to 19.5 grams; stomach contents small insect fragments. The black-faced
rufous warbler was netted largely in the undergrowth in heavily exploited parts
of the forest.
Camaroptera chloronota toroensis (Jackson)
Very common; eight specimens, October 28 to November 23; all in non-
breeding state; bill black above, pale horn to whitish gray below; iris pale
brown to gray-brown; feet pale brown; weight 8 to 13 grams; stomach contents
insect fragments.
Camaroptera brachyura tincta (Cassin)
This little warbler was as numerous as the preceding species; six examples
were taken October 29 to November 21; most were in non-breeding state, but
one male, November 6, had enlarged gonads; bill black; iris pale brown; feet
pale pinkish brown; weight 11 to 12.5 grams; stomach contents insect
fragments.
Sylvietta leucophrys chloronota Hartert
One specimen, a non-breeding female, was taken November 10; bill pale
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Birds of the Kalinzu Forest
21
horn-brown, pinkish at the base; iris chestnut; feet deep pink; weight 10 grams;
stomach contents insect fragments.
Hylia prasina prasina (Cassin)
The green hylia was taken once in the Kalinzu Forest, November 13, and
a second example was collected in the Maramagambo Forest, November 8;
both in non-breeding state; bill black, the tomium and gape yellow; iris dark
brown; feet olive to olive-gray; weight 15 and 16 grams; stomach contents
insect matter.
Laniidae
Tchagra minuta minuta (Hartlaub)
This shrike was not collected in the Kalinzu Forest, but on November 8
two birds, one of each sex, were taken in a grass covered glade in the nearby
Maramagambo Forest; the male had large testes, the female had a small ovary;
bill black; iris pale maroon; feet gray; weight 31 and 36 grams; stomach con-
tents grasshoppers and beetles.
Laniarius liihderi luhderi Reichenow
This bush-shrike was common in the Kalinzu Forest; seven specimens
were taken October 28 to November 10; gonads ranging from small to much
enlarged; bill black; in immature birds it is blackish horn above, paling to pale
blue-gray basally on the mandible; feet gray to bluish gray; weight 41 to 47.5
grams; stomach contents grasshoppers, beetles and other insect remains.
Telophorus bocagei jacksoni (Sharpe)
This shrike is not uncommon in the Kalinzu Forest; two females were
collected on October 30 and November 19; bill black; iris dark brown; feet
dark gray; weight 27 grams each; stomach contents insect larvae.
Lanins coUurio phoenicuroides (Schalow)
This winter visitor from Eurasia is represented by one immature in-
dividual, taken on November 7; bill horn-gray, the tip blackish; iris very dark
brown; feet gray black; weight 26.5 grams; stomach contents small grass-
hoppers. It was shot in secondary growth in one area of forest that had been
exploited.
Lanius mackinnoni Sharpe
Not uncommon; three specimens, October 30, November 7 and 8; all
in non-breeding state; bill black; iris dark brown; feet black; weight 34 to 37
grams; stomach contents caterpillars, grasshoppers and beetle fragments.
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No. 195
Oriolidae
Oriolus brachyrhynchus laetior Sharpe
One adult male of this oriole was taken in the Kalinzu Forest, November
16, and three immature birds were collected in the Maramagambo Forest, on
November 8; bill of adult dull pinkish brown-horn; of immature birds blackish,
pinkish brown basally; iris wine-red in adult male, dull red to dark grayish
brown in the others; feet blue-gray; weight 48 to 56 grams; stomach contents
fruits, small seeds, caterpillars and insect fragments.
Oriolus larvatus percivali Ogilvie-Grant
One adult male with enlarged gonads was taken on October 31, in a mist
net in the undergrowth in an area of heavily exploited forest; bill metallic deep
pink; iris dark crimson; feet dark gray; weight 74 grams; stomach contents
caterpillars.
Paridae
Pams funereus funereus ( Verreaux)
This tit was common in the Kalinzu, where five examples were collected
October 31 to November 2. All were in non-breeding state; bill black; iris red
in four cases, yellow-brown in one; feet slate gray; weight 23 to 27 grams;
stomach contents caterpillars, beetle larvae and small insect remains.
Sturnidae
Poeoptera stuhlmanni (Reichenow)
Two specimens of this starling, a male with large gonads and a female in
non-breeding condition, were taken on October 30 and 31 ; bill black; iris dark
brown; with outer ring yellow; feet black; weight 39 (female) and 44 (male)
grams; stomach contents remains of fruits.
Poeoptera lugubris Bonaparte
Five adult males, November 2 to 14, with gonads ranging from small to
much enlarged; bill black; iris bright yellow; feet black; weight 42 to 43 grams;
stomach contents fruit fragments, seeds, beetles and other small insect remains.
Keith (1968) has recently described a race of this starling, P. 1. webbi,
from the Impenetrable and Kibale Forests, on the basis of greater wing length
and a bluish sheen to the purple gloss of the feathers in the male. He gives the
wing length of males of nominate lugubris as 89 to 96 (average 93.13 mm) , of
webbi as 94 to 99 (average 97.23 mm) . Our five Kalinzu males bridge this
difference as they have wing lengths of 93 to 97 (average 95 mm) . In color
they agree with our series of webbi but there is some slight individual variation
in the amount of blue or purple. No examples of nominate lugubris from
Gabon have been available for comparison and we therefore use a binomial
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Birds of the Kalinzu Forest
23
for our specimens. We cannot avoid the conclusion that webhi is a poorly
marked form, if a valid one.
Onychognathus walleri elgonensis (Sharpe)
Four examples of this race of Waller’s chestnut-winged starling, two
males and one female with large gonads and one female with a small ovary,
were taken November 4 to 14; bill black; iris crimson-red with the inner ring
dark brown; feet black; weight 70 to 82 grams; stomach contents small green
fruits and fruit seeds. These birds were seen feeding in a fruiting Symphonia
tree.
Lamprotoniis splendidus splendidus ( Vieillot)
This starling was fairly common in the Kalinzu Forest; one male and
two females, all with enlarged gonads, were collected on October 31, Novem-
ber 12 and 14; bill black; iris yellow-white to cream; feet black; weight 120
to 135 grams; stomach contents fruits and seeds.
ZOSTEROPIDAE
Zosterops senegalensis toroensis Reichenow
Two specimens of this common white-eye, one of each sex, taken on
November 19, agree with a series of toroensis from the Kibale and Bwamba
Forests; bill black; iris dark brown; feet gray; weight 1 1 grams each; stomach
contents insect fragments.
Nectariniidae
Anthreptes fraseri axillaris (Reichenow)
Common in the Kalinzu; six examples of this sunbird were collected
October 31 to November 18; gonads ranging from small to large; bill black to
dark brown above, pinkish white below; iris brown to reddish brown; feet
olive to olive-gray to gray-brown; weight 10 to 13 grams; stomach contents
insect remains. This species was found chiefly in natural, undisturbed areas of
the forest.
Anthreptes rectirostris tephrolaema (Jardine and Fraser)
This sunbird was found to be very common in the Kalinzu Forest, where
eight specimens were obtained, October 30 to November 19, and one in the
Maramagambo Forest, November 8; of five males four had large gonads, while
of four females none showed gonadal swelling; bill black; iris red-brown to
dark brown; feet black; weight 10 to 12 grams; stomach contents insect frag-
ments, small spiders and bits of fruit. The gray-chinned sunbird was found
both in natural and in heavily exploited areas of the forest.
24
Contributions in Science
No. 195
Anthreptes collaris garguensis Mearns
Our collectors met with this sunbird only on October 31 and November 1;
on each day they obtained a single example, so the bird may not be very
numerous in the Kalinzu; bill black; iris dark brown; feet black; weight 8.5
grams; stomach contents insect remains.
Nectarinia olivacea ragazzn (Salvadori)
The olive sunbird was found to be very numerous in the Kalinzu, and
eleven examples were collected November 2 to 18; gonads ranging from small
to large; bill black above, the base of the mandible dull yellow to pinkish
brown; feet dark brown to black; weight 9 to 12.5 grams; stomach contents
insect fragments.
Nectarinia verticalis viridisplendens (Reichenow)
This sunbird was also very common in the Kalnizu Forest; six males and
one unsexed, but probably female by plumage, were collected November 1 to
10; gonads varying from small to large; bill black; iris dark brown; feet black;
weight 12 to 15 grams (one marked 26.5 grams, probably an error for 16.5
grams) ; stomach contents insect fragments, small insect larvae.
Nectarinia rubescens rubescens (Vieillot)
Common in the Kalinzu; seven specimens October 30 to November 11;
gonads small to large; bill black; iris dark brown; feet black; weight 10.5 to
12.5 grams; stomach contents insect fragments. The green-throated sunbird
was seen chiefly feeding in fairly tall trees.
Nectarinia bouvieri (Shelley)
This sunbird was met with but once, on November 24 when an adult
male with large testes was collected; bill black; iris dark brown; feet black;
weight 10 grams; stomach contents very small insect fragments. The bird was
found in the small scrub-bush of a grass covered hillside at the edge of the
forest.
Nectarinia purpureiventris (Reichenow)
The purple-breasted sunbird, a species primarily of highland forest, was
seen several times in the Kalinzu by Williams, but he was unable to collect one.
Nectarinia kilimensis kilimensis Shelley
One female, ovary enlarged, was taken on November 25; bill black; iris
dark cream; feet black; weight 16 grams; stomach contents insect fragments.
1970
Birds of the Kalinzu Forest
25
Ploceidae
Amblyospiza alhifrons melanota (Heuglin)
One adult male of this heavy-billed weaver in breeding condition was
taken November 13 from a small flock seen in the undergrowth of an ex-
ploited area of the forest; bill horn-black; iris dark brown; feet grayish black;
weight 47 grams; stomach contents seeds.
Ploceus nigricollis nigricollis (Vieillot)
Common in the Kalinzu; three males and four females were collected,
October 31 to November 25, all in non-breeding state; bill black; iris dark
brown; feet dark gray; weight 21.5 to 29 grams; stomach contents small beetles,
green caterpillars and other insect fragments. Several of these specimens were
caught in mist nets in the undergrowth in exploited areas of the forest.
Ploceus melanogaster stephanophorus (Sharpe)
This weaver was very common in the Kalinzu; nine specimens were taken
October 28 to November 24; one male with large gonads, all the others with
small ones; bill black; iris dark brown; feet dark gray; weight 20.5 to 25 grams;
stomach contents insect remains, including beetles and small caterpillars. Four
of the specimens are molting into full adult plumage. A number of these black-
billed weavers were caught in nets in the undergrowth in a heavily exploited
area of the forest, where they came, together with other species, to feed on
flying termites as they emerged from the ground.
Ploceus superciliosus (Shelley)
Not taken in the Kalinzu, but on November 8 one (unsexed) example
was collected in the Maramagambo Forest; bill black above, pale gray below;
iris dark brown; feet pale brown; weight 21 grams; stomach contents grass
seeds.
Ploceus insignis insignis (Sharpe)
This weaver was met with but once, on November 2, when one adult of
each sex, both in non-breeding state, were taken; bill black; iris red-brown in
male, bright brick-red in female; feet pale brown; weight 28 grams each;
stomach contents insect fragments. The birds were searching for food among
the moss and lichen on a small tree.
Malimhus ruhricolUs rubricollis (Swainson)
This malimbe was common in the Kalinzu Forest; four examples were
collected October 31 to November 2; all in non-breeding condition; bill black;
iris dark red-brown; feet black; weight 39 to 45 grams; stomach contents insect
fragments.
26
Contributions in Science
No. 195
Estrildidae
Parmoptila woodhousei jamesoni (Shelley)
The Kalinzu Forest is an additional Uganda locality for this Congolese
flower pecker. One adult male, with enlarged testes, was taken on November
13; bill black; iris very dark red-brown; feet pale huffy brown; weight 10
grams; stomach contents small black ants.
Nigrita fusconota fusconota Fraser
This negro finch was met with but once in the Kalinzu, on November 25,
when a male with large testes and a female in non-breeding state were taken;
bill black; iris dark brown; feet dark gray; weight 8 and 9.5 grams; stomach
contents very small insects.
Nigrita canicapilla schistacea Sharpe
This species was not uncommon in the Kalinzu; four specimens were
taken on October 30 and 31, but then no more were encountered although the
collectors were there for nearly a month; one of our birds was in breeding
condition; the others were not; bill black; iris orange-red; feet dark brown;
weight 19 to 20 grams; stomach contents small seeds and insect fragments.
Nesocharis ansorgei ansorgei (Hartert)
It was something of a surprise to find this little waxbill fairly common in
the Kalinzu Forest; five specimens were taken October 30 to November 7; all
in non-breeding state; bill black, pale gray at base; iris dark brown; feet
blackish; weight 8.5 to 10 grams; stomach contents small crushed grass seeds.
The birds were found in the tall undergrowth in areas of forest that had been
heavily exploited. The Kalinzu Forest is an additional locality to those men-
tioned by the senior author (Friedmann, 1968) in his discussion of this little
weaver finch.
Spermophaga ruficapilla ruficapilla (Shelley)
As in so many west Ugandan forests, this bird was found to be common
in the Kalinzu; six examples were collected, October 29 to November 10; all in
non-breeding condition; bill black with metallic blue wash, the tomium pinkish
orange; iris dark brown; feet dark brown to dark gray-brown; weight 19.5 to
26.5 grams; stomach contents seeds and grit.
Lonchura bicolor poensis (Fraser)
Although this mannikin is not a typical forest bird, the one example
obtained was collected in a mist net in an exploited part of the forest; bill pale
gray; iris dark brown; feet dark gray; weight 10 grams; stomach contents grass
seeds.
1970
Birds of the Kalinzu Forest
27
Fringillidae
Serinus frontalis frontalis Reichenow
This little finch is represented by a single example collected November
12; bill horn brown above, pale horn below; iris dark brown; feet brown;
weight 13 grams; stomach contents insects and small seeds. It was taken in the
scrub undergrowth at the edge of the forest.
Serinus burtoni tanganjicae Granvik
Fairly common in the heavily exploited parts of the Kalinzu Forest. Three
specimens were taken November 4 to 20; gonads small to large; bill horn-
brown, paler below; iris medium to dark brown; feet pale to dark brown;
weight 30 to 32 grams; stomach contents seeds and flower buds.
Literature Cited
Chapin, J. P. 1953-1954. The birds of the Belgian Congo. Pt. III. Amer. Mus. Nat.
Hist. 75A: 1-821; Pt. IV, 75B: 1-846.
Friedmann, H. 1968. The olive weaver-finch, Nesocharis ansorgei ansorgei in
Uganda. Bull. British Ornith. Club 88: 135-128.
Friedmann, H., and J. G. Williams. 1968. Notable records of rare or little known
birds from western Uganda. Rev. Zool. Bot. Afr. 77 : 11-36.
Keith, S. 1968. A new subspecies of Pooeptera lugubris Bonaparte from Uganda.
Bull. British Ornith. Club 88: 119-120.
Keith, S., and A. Twomey. 1968. New distributional records of some East African
birds. Ibis 110: 537-548.
Keith, S., A. Twomey, H. Friedmann, and J. G. Williams. 1969. The avifauna of
the Impenetrable Forest, Uganda. Amer. Mus. Novitates 2389: 1-41.
Prigogine, a. 1969. Polymorphism of the chestnut-bellied wattle-eye, Dyaphoro-
phyia concreta. Ibis 111: 95-96.
Rand, A. L. 1960. African Pynonotidae, in Mayr and Greenway, eds., Check-List of
Birds of the World, 9: 221-300.
White, C. M. N. 1960. A check list of the Ethiopian Muscicapidae (Sylviinae). Pt. I.
Natl. Mus. S. Rhodesia, Bulawayo, Occas. Papers Nat. Sci. 3 : 399-450.
. 1961. A revised check list of African broadbills, pittas, larks, swallows,
wagtails, and pipits. Lusaka, Govt. Printer. 84 p.
. 1962a. A check list of the Ethiopian Muscicapidae (Sylviinae). Pt. II. Natl.
Mus. S. Rhodesia, Bulawayo, Occas. Papers Nat. Sci. 3: 653-738.
. 1962b. Revised check list of African shrikes, orioles, drongos, starlings,
crows, waxwings, cuckoo-shrikes, bulbuls, accentors, thrushes and babblers.
Lusaka, Govt. Printer. 176 p.
. 1963. A revised check list of African flycatchers, tits, tree creepers, sun-
birds, white-eyes, honey eaters, buntings, finches, weavers and waxbills. Lusaka,
Govt. Printer. 218 p.
. 1965. A revised check list of African non-passerine birds. Lusaka, Govt.
Printer. 299 p.
Accepted for publication April 2, 1970
LOS
ANGELES
' COUNTY
MUSEUM
CONTRIBUTIONS
SL IN SCIENCE
iJMBER 196
July 1, 1970
TWO NEW SPECIES OF CATFISHES OF THE GENERA
NANNORHAMDIA AND IMPARALES (FAMILY
PIMELODIDAE) FROM CENTRAL AMERICA
I
By William A. Bussing
Los Angeles County Museum of Natural History •
Los Angeles, California 90007
Exposition Park
'i
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 sep-
arately, and numbers run consecutively regardless of subject matter. Number 1 was
issued January 23, 1957. The series is available to scientific institutions and scien-
tists on an exchange basis. Copies may also be purchased at a nominal price. Inquiries
should be directed to Virginia D. Miller, Los Angeles County Museum of Natural
History, 900 Exposition Boulevard, Los Angeles, California 90007.
Robert J. Lavenberg
Managing Editor
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Manuscripts for the LOS ANGELES COUNTY MUSEUM, CONTRIBU-
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largely with specimens in the Museum’s collections, other technical papers will be
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side the Museum.
Authors proposing new taxa in a CONTRIBUTIONS IN SCIENCE must
indicate that the primary type has become the property of a scientific institution of
their choice and cited by name.
MANUSCRIPT FORM.— (1) The 1964 AIBS Style Manual for Biological
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Virginia D. Miller
Editor
TWO NEW SPECIES OF CATFISHES OF THE GENERA
NANNORHAMDIA AND IMPARALES (FAMILY
PIMELODIDAE) FROM CENTRAL AMERICA
By William A. Bussing^
Abstract: Two new species of pimelodid catfishes, Nanno-
rhamdia lineata and Imparales panamensis, are described from
Costa Rica and Panama respectively. Nannorhamdia is recorded
for the first time in Central America and each species represents
a relict population. The relationships are discussed and habitats
described for each species.
Recent collections by Horace Loftin (1965) in Panama and by Albert
Greenberg in Costa Rica have revealed two undescribed species of small
pimelodid catfishes of the genera Nannorhamdia and Imparales. The former
was previously known only from South America, whereas Miller (1966)
included Imparales sp. in his checklist of Central American fishes. In 1967
Oscar Blanco B. and I obtained additional specimens of the Costa Rican
Nannorhamdia. More specimens of Imparales were collected in 1967 in the
Chucunaque and Balsas drainages of Panama by members of the Battelle
Memorial Institute, Pacific Northwest Laboratory (Battelle NW) as part of
their sub-contractual work for the Interoceanic Sea Level Canal Feasibility
Study. Material from all these sources was used in this study.
Nannorhamdia lineata, new species
Figure 1
Holotype: LACM 30688-1 ( 9, 74.1 mm SL) ) , Costa Rica: Puntarenas
Prov., Quebrada 36 (elev. 80 m) 12 km W of Pueblo Rio Claro at Inter-
american Highway; 5 January 1967; collectors W. Bussing, Oscar Blanco B.
Paratypes: UCR 1 1 1-10 (18, 48.5-71.7 mm SL), same data as holotype;
LACM 30688-2 (3, 51.8-69.9 mm SL), same data as holotype; BMNH
1967.9.29.1-2 (2, 64.0-68.2 mm SL) , same data as holotype; USNM 204694
(2, 58.4 and 69.5 mm SL) , same data as holotype; USNM 194230 ( 1, 57.0
mm SL) Costa Rica: Puntarenas Prov., Rio Salama Nuevo (?) at Interameri-
can Highway; 21 January 1963; collector Albert Greenberg.
Type specimens are deposited in the British Museum (Natural History)
(BMNH) , Los Angeles County Museum of Natural History (LACM) , Museo
de Zoologia, Universidad de Costa Rica (UCR) and the United States National
Museum (USNM).
Comparative Material : Nannorhamdia nemacheir (Eigenmann and
^Departamento de Biologia, Universidad de Costa Rica, San Jose, Costa Rica; and
Research Associate in Ichthyology, Los Angeles County Museum of Natural History.
1
2
Contributions in Science
No. 196
Figure 1. Nannorhamdia lineata, new species, lateral view of holotype LACM
30688-1, 74.1 mm SL; dorsal view of paratype UCR 1 1 1-10, 71.1 mm SL.
Fisher), USNM 121167 (30, 33.8-68.5 mm SL), Venezuela: Rio San Juan
at bridge, tributary to Rio Motatan; 17 and 20 March 1942; collector L. P.
Schultz. UCR 318-1 (6, 30.3-55.1 mm SL) , Colombia : Rio Salado, ca. 14 mile
upstream from junction with Rio Truando; 27 August 1967; collector H.
Loftin. Cetopsorhamdia nasus (Eigenmann and Fisher), USNM 86816 (1,
62.5 mm SL), Bolivia: Tumupasa; December 1921; collector N. E. Pearson.
Diagnosis: A Nannorhamdia with pronounced lateral stripe extending
from the tip of snout to caudal base; short maxillary barbels not passing pec-
toral fin base; subequal caudal lobes; 8-1 1 total gill rakers; 7-9 principal
(branched rays plus one simple ray) anal fin rays; no prominent dorsal blotches
on body contrasting with ground color; all pectoral and dorsal fin elements
unserrated.
The new species is immediately separated from N. guttatus Pearson,
TV. benedettii Fernandez Yepez, TV. stictnotus Fowler and TV. nemacheir Eigen-
mann and Fisher by its lateral stripe and short maxillary barbels; from TV.
guttatus and TV. benedettii it also differs in having fewer anal rays (7-9 vs.
10-12) . From TV. macrocephala Miles it differs in having fewer anal rays
(7-9 vs. 13) and lacking dorsal and pectoral spines with denticulations on the
posterior borders. It differs from TV. schubarti Gomes principally in having
fewer gill rakers (8-1 1 vs. 9-15), shorter maxillary barbels (22.4-26.8 vs. 32.1-
55.5), shorter first dorsal ray (14.3-16.9 vs. 19.4-24.6) and shorter first pec-
toral ray (13.3-15.4 vs. 16.7-20.0) .
The new species is closest to TV. spurrellii of the Rio San Juan drainage.
1970
Two New Species of Catfishes
3
Colombia. The latter is distinguished from N. lineata by its narrower inter-
narial distance, interorbital distance and caudal peduncle depth, much longer
maxillary and mental barbels, higher gill raker count and much more promi-
nent lateral stripe. Mr. Alwyne Wheeler {personal communication, 1968)
reported that the type specimen of N. s purr el Hi is quite round-bodied and does
not seem to have shrunk, thus discounting unnatural proportional differences
due to preservation.
Description: Head slightly depressed, snout bluntly rounded. Narrow
fontanel in middorsal line extending from before eyes to base of occipital
process. Mouth subterminal, snout projecting. Villiform teeth in bands on both
jaws; premaxillary patches slightly wider laterally, but without posteriorly
projecting angles; dentary patches wider medially, very narrow laterally; no
teeth on vomer or palatines. Nostrils wide apart, anterior ones tubular and in
line with bases of maxillary barbels; posterior nostrils with an anterior flap
and lying equidistant between anterior nostril and margin of eye. Maxillary
barbels the longest, one pair lying in a groove along the snout and extending
along the first third or half of the length of pectoral fin; two pairs of mental
barbels, their bases in a straight line, outer pair reaching to or slightly beyond
origin of pectoral fin; inner pair extending three-fourths the distance to pectoral
origin. No ontogenetic variation in barbel length on specimens in the present
size range.
Caudal fin well forked, shortest median rays slightly longer than half
the length of the longest rays; upper lobe slightly longer and more pointed than
lower. First dorsal ray soft, not extended into a filament, margin of fin trun-
cate; origin slightly in advance of pelvic fin origin. Anal fin margin rounded;
origin directly below adipose origin. Adipose fin short, slightly longer than
anal; height about 4.5 times in length. Pelvic fins short, rounded; arising below
third or fourth dorsal ray. First pectoral ray soft, net extended into a filament;
appressed fin not quite reaching a vertical throuigli dorsal origin. Anus between
middle of length of pelvic fins.
Counts: The counts and body proportions in percent of standard length
of the holotype and nine paratypes of N. lineata from Quebrada 36 are re-
corded below, followed by data for the type of TV. spurrellii in parentheses.
Data for the type specimen of TV. spurrellii Regan (BMNH 1913.10.1.41; Rio
Condoto, San Juan drainage, Colombia) were kindly furnished by Alwyne C.
Wheeler of the Zoology Department, British Museum of Natural History.
Proportional measurements of the single specimen from Rio Salama Nuevo,
Costa Rica, which in some cases fall slightly out of the range given below due
to its shrunken condition, are not included.
Dorsal rays i,6; i,5-6 (i,6) . Anal v,8; iv-v,6-8 (ii,6) . Pectoral i,9; i,8-9
(i,9). Pelvic i,5; i,5 (i,5) . Branched caudal rays 15, 15-16 ( 15) . Gill rakers of
holotype 2 -f 7 and 1 + 8, total 9; paratypes 1-2 -1- 7-10, total 8-11 (TV. spur-
rellii 2 -f 10, total 12) .
Comparison of holotype and 16 paratypes of Imparales panamensis and holotype of
I. mariai. Proportions expressed in thousandths of standard length.
4
Contributions in Science
No. 196
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1970
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6
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cottids
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Cynoscion nobilis
Cynoscion reticulatus
1970
Fish Remains from the Palos Verdes Sand
7
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Prionotus stephanophrys
Rhacochilus vacca
1970
Fish Remains from the Palos Verdes Sand
9
I I i
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opercular spine; t = teeth; j = jaw; s = scales * in Roy Kohl collection ** in Humboldt State College collection
10
Contributions in Science
No. 199
inaccessible for paleontological investigation, the fauna seems worthy of
listing.
LACMIP 66
This site was first investigated by museum personnel in 1945. At that
time, and for nearly two decades, there were numerous exposures of fos-
iliferous Palos Verdes Sand on both sides of a dry creek and in eight erosion
gullies that drained into it. These gullies were listed successively as LACMIP
66-1, 66-2, 66-3, etc., and although each was sampled extensively for mollusks,
few fish remains were found. A housing development now covers the entire
mesa south of Newport back bay where LACMIP 66 yielded a rich assemblage
of mollusks (Kanakoff and Emerson, 1959).
Fish remains (otoliths == o, teeth = t, caudal stings = s) gleaned from
the main creek and its tributary gullies by museum personnel were identified
as follows:
Cynoscion nobilis:
Dasyatis dipterurus:
embiotocid:
Genyonemus lineatus:
Heterodontus francisci:
Leptocottus armatus:
Otophidium scrippsi:
Otophidium taylori:
Parophrys vetulus:
Pimelometopon pidchrum:
Se bastes sp.:
Seriphus politus:
66 (lo)
66 ( Is)
66 (lo, 6t)
66-2 (8o); 66-10 (3o)
66-10 (It)
66-2 (lo)
66 (lo); 66-2 (2o)
66 (lo)
66-1 (lo)
66 (17t)
66 (lo)
66 (3o); 66-2 (2o); 66-10 (20)
Kanakoff (1956) noted 11 kinds of sharks and rays from LACMIP 66-2
and five species of teleosts. Ten of the 1 1 elasmobranchs were different species
from the two noted above (i.e., Carcharhinus lamiella [= obscurus\, Car-
charodon carcharias, Cetorhinus maximus, Isurus glaucus {= oxyrinchus\
Lamna ditropis, Myliobatis californicus, Prionace glauca, Sphyrna zygaena,
Triakis semifasciata and Urobatis [= Urolophus] halleri).
LACMIP 68-B
This exposure was near the top of a cliff about 90 feet above the high
tide line in Newport back bay. A housing development, across the bay from
the one covering LACMIP 66, has destroyed this site. Kanakoff (1956)
reported 12 teeth from Carcharhinus lamiella [= obscurus\
LACMIP 136
This 14- to 22-foot thick bed of Palos Verdes Sand was rich in mollusks
but contained few fish remains. It was in the cliff near the top of the mesa
that lies south of Newport back bay. The housing development that covers
1970
Fish Remains from the Palos Verdes Sand
11
LACMIP 66 also destroyed this locality. About 200 pounds of fossiliferous
matrix yielded remains from the following six elasmobranchs and five teleosts
(r = gill raker) :
Cetorhinus maximus:
(It)
Dasyatis dipterurus:
(Is)
Galeorhinus zyopterus:
(3t)
Genyonemus lineatus:
(5o)
Heterodontus francisci:
(It)
Leptocottus armatus:
do)
Myliobatis californiciis:
(It)
Otophidium scrippsi:
(3o)
Pimelometopon pulchrum:
(2t)
Porichthys myriaster:
(skull with 2o)
Urolophus halleri:
(Is)
Newport-Corona del Mar Shopping Center
During construction of a vast shopping center east of Pacific Coast High-
way and between Newport Beach and Corona del Mar, several beach deposits
of Palos Verdes Sand were exposed. Badly eroded shells of the bean clam,
Donax gouldii, comprised the bulk of the mollusk remains in these beds, but
shells of Pismo clams, Tivela stultorum, moon snails, Polinices spp., whelks,
Nassarius spp. and olives, Olivella biplicata, were also common. A 50-pound
field sample from one of these beds yielded six badly eroded otoliths from
Genyonemus lineatus and one from Seriphus politus. These strata are now
under parking lots and an assortment of buildings.
Northern California Site
Crannell Road
This locality was exposed during highway construction north of Areata
where Crannell Road intersects U.S. Highway 101. Excavation of the hill
at the southeast corner of this road junction revealed a three foot thick,
highly fossiliferous layer of relatively coarse sand beneath 150 feet of muddy
silt containing numerous thin beds and lenses of marine fossils. All of the
fish remains from this locality (Tables 1 and 2) were recovered from the
sandy matrix at the base of the hill, at an elevation of approximately 20 feet
(Crannell Quadrangle, Calif., 1.5' series, U.S.G.S. 1966).
Numerous students from Humboldt State College have sampled this
site extensively and frequently, as have amateur and professional paleontolo-
gists. I have had access to the collections of three students, Lloyd Barker,
Jack Hopkins and Roy Kohl, and two of the species I have included (Platich-
thys stellatus and Sebastes serranoides) are based upon otoliths in Roy Kohl’s
collection, while the soupfin shark {Galeorhinus zyopterus) record is based
upon a tooth in Lloyd Barker’s collection. The Pacific halibut (Hippoglossus
12
Contributions in Science
No. 199
stenolepis) is represented by a mandible in a collection at Humboldt State
College.
In all, the 700 to 800 pounds of matrix that I examined from this site
yielded 724 otoliths and otolith fragments, 137 teeth including four species
of teleosts not represented by otoliths, plus an assortment of vertebrae, skate
bucklers and wing spines and miscellaneous bony fragmentia (Tables 1 and
2).
Species Accounts
Elasmobranchs
Alopias vulpinus ( Bonnaterre ) —common thresher
Teeth of the common thresher were found only at Playa del Rey (Fitch
1964, 1966).
Total material : 3 teeth.
Carcharhinus spp.— requiem sharks
Although Kanakoff ( 1956) reported teeth of C. lamiella [= obscurus]
from Playa del Rey and two Newport Beach localities, I prefer to list them
simply as Carcharhinus spp. until Carcharhinus taxonomy is better under-
stood than at present. Carcharhinus teeth were found at Playa del Rey (Fitch
1964, 1966), San Pedro, and Newport Beach (Table 1) ; at least two species
are represented.
Total material : 109 teeth.
Carcharodon carcharias ( Linnaeus ) —white shark
C. carcharias has been reported previously from Playa del Rey (Kana-
koff, 1956; Fitch, 1964) and Newport Beach (Kanakoff, 1956). A white
shark tooth also was found at San Pedro (Table 1).
Total material: 10 teeth and 4 vertebrae.
Cetorhinus maximus ( Gunnerus ) —basking shark
Kanakoff ( 1956) reported a Cetorhinus tooth from LACMIP 66-2 at
Newport Beach. Since then I have found single gill rakers in material from
the 700 block North Pacific Ave., San Pedro, and from LACMIP 136 at
Newport Beach (Table 1).
Total material : 1 tooth and 2 gill rakers.
Dasyatis dipterurus (Jordan and Gilbert) —diamond stingray
Kanakoff ( 1956) identified a caudal sting from LACMIP 77 at Wilming-
ton, Calif., as being from this species, and Fitch ( 1966) reported two D.
dipterurus teeth from Playa del Rey. Eight additional teeth and a sting have
been found since then at San Pedro (700 block North Pacific and the In-
cinerator site) and at two Newport Beach localities, LACMIP 66 and 136
(Table 1).
Total material: 10 teeth and 2 caudal stings.
1970
Fish Remains from the Palos Verdes Sand
13
Galeorhinus zyopterus Jordan and Gilbert— soupfin shark
The eight teeth that Kanakoff (1956) identified as being from Triakis
semifasciata proved to be from Galeorhinus. Fitch ( 1964, 1966) subsequently
reported soupfin shark teeth from Playa del Rey, and included the seven
that Kanakoff had misidentified. Galeorhinus teeth were later found in
several San Pedro deposits, at Newport Beach and at Crannell Road (Table
1).
Total material: 27 teeth, including one in the Lloyd Barker collection.
Heterodontus francisci (Girard)— horn shark
Kanakoff (1956) reported finding four dorsal spines from Heterodontus
at Newport Beach (LACMIP 66-2), and Fitch (1966) noted one tooth
from LACMIP 59 at Playa del Rey. Seven additional teeth have been found
at San Pedro and Newport Beach localities in recent years (Table 1 ).
Total material: 8 teeth and 4 dorsal spines.
Isurus oxyrinchus Rafinesque— mako
Kanakoff (1956) reported teeth from /. glaucus [= oxyrinchus^ in de-
posits at Newport Beach (LACMIP 66-2) and San Pedro (LACMIP 131),
and both Kanakoff (1956) and Fitch (1964, 1966) reported them from Playa
del Rey (LACMIP 59). Only two additional mako teeth have been found
in Palos Verdes Sand subsequent to these reports, and both of these were
from the LACMIP 131 locality (Table 1).
Total material: 22 teeth.
Lamna ditropis Hubbs and Follett— salmon shark
Kanakoff (1956) noted five salmon shark teeth from the Palos Verdes
Sand at Newport Beach (LACMIP 66-2), and reported that it is “strictly
northern in distribution.” Actually, L. ditropis inhabits oceanic waters both
north and south of southern California, so Newport Beach can not be con-
sidered as being outside of its normal range, latitudinally. A salmon shark
tooth found in Palos Verdes Sand at San Pedro (LACMIP 131) has not been
reported previously.
Total material: 6 teeth.
Myliobatis californicus Gill— bat ray
Bat ray teeth are large and very distintive, so they are easily seen in
fossiliferous matrix. Their abundance in southern California Pliocene and
Pleistocene deposits, especially Palos Verdes Sand (Table 1), reflects the
ease with which they can be found. Kanakoff (1956) reported 160 M. cali-
fornicus teeth from LACMIP 131 at San Pedro, and caudal stings from
Playa del Rey and Newport Beach (LACMIP 66-2). I reported finding teeth
of M. californicus at Playa del Rey (Fitch, 1964, 1966) and subsequently
14
Contributions in Science
No. 199
have found them in fossiliferous Palos Verdes Sand at San Pedro and New-
port Beach (Table 1).
Total material: 613 teeth and 8 stings.
N otorynchus maculatus Ayres— sevengill shark
N. maculatus previously was reported from Play a del Rey by Kanakoff
(1956) and Fitch (1964) based upon 12 teeth. A single tooth found at San
Pedro (700 block, North Pacific Ave.) has not been noted previously. The
range of N. maculatus has generally been given as northern British Columbia
to southern California, but there are now several records of capture from
Baja California as far south as Pta. San Carlos.
Total material: 13 teeth.
Prionace glauca (Linnaeus)— blue shark
Blue shark teeth are rare in southern California fossil deposits, but Kana-
koff (1956) reported finding eight at Playa del Rey and one at Newport
Beach (LACMIP 66-2). In my sampling of the Palos Verdes Sand, I have
found but a single tooth from P. glauca and that was in screenings from
LACMIP 131 at San Pedro.
Total material: 10 teeth.
Raja spp.— unidentified skates
I previously reported Raja from Palos Verdes Sand based upon eight
vertebrae and seven bucklers and “wing” spines (Fitch, 1964, 1966). Subse-
quent sampling at San Pedro has turned up five additional bucklers and 13
teeth, but Raja remains have not been found at Newport Beach. In northern
California (Crannell Road), skate remains were abundant, and I was able
to find 1 14 teeth and 1 1 wing spines and bucklers (dermal denticles) in the
material I examined. Lloyd Barker’s collection contains 16 teeth and five
dermal denticles.
Total material : 143 teeth, 28 dermal denticles and 8 vertebrae.
Rhizoprionodon longurio (Jordan and Gilbert)— Pacific sharpnose shark
R. longurio was previously reported from Playa del Rey (as Scoliodon
longurio) by Fitch ( 1964) who had a single tooth from there. No additional
remains from this species have been found in any exposure of Palos Verdes
Sand.
Total material: 1 tooth.
Sphyrna spp. —unidentified hammerheads
Kanakoff ( 1956) reported finding four teeth from Sphyrna zygaena at
Playa del Rey and one at Newport Beach (LACMIP 66-2), but in my reports
on the Playa del Rey fauna (Fitch 1964, 1966), I did not attempt to determine
from what species of Sphyrna the nine teeth and 17 vertebrae came. Four
1970
Fish Remains from the Palos Verdes Sand
15
hammerhead teeth that I have obtained from deposits of Palos Verdes Sand
at San Pedro (Table 1) appear to be from a single species, but I do not feel
qualified to place a specific name on them. Gilbert (1967) reports only two
species of Sphyrna from California during modern times, S. tiburo and S.
zygaena.
Total material: 18 teeth and 17 vertebrae.
Squatina californica Ayres— Pacific angel shark
Teeth, vertebrae and dermal denticles of S. californica have turned up in
Palos Verdes Sand at Playa del Rey (Fitch 1964, 1966) and San Pedro
(Table 1), but no identifiable angel shark remains have been found at New-
port Beach, as yet.
Total material: 23 teeth, 8 vertebrae, and 1 dermal denticle.
Triakis semifasciata Girard— leopard shark
Although Kanakoff ( 1956) reported finding eight teeth of Triakis at
Playa del Rey and Newport Beach, a critical comparison of his material with
teeth from living sharks revealed that they were actually from Galeorhinus
zyopterus, a close relative. Deposits of Palos Verdes Sand at San Pedro and
Newport Beach have yielded Triakis teeth, however (Table 1).
Total material: 5 teeth.
Urolophus halleri Cooper— round stingray
Several hundred caudal sting fragments have been identified as being
from round stingrays (Kanakoff, 1956; Fitch, 1964, 1966), and subsequent
sampling of the Palos Verdes Sand has turned up additional stings (Table 1),
but so far no other remains from Urolophus have been found. The jaws of an
adult round stingray contain more than 1,000 teeth, but these are so small
that they easily pass through a 20-mesh screen, and I have not examined
very much residue that is finer than what is retained by 20-mesh screens.
Total material: 246 stings and fragments of stings.
Unidentified elasmobranchs
Most of the deposits of Palos Verdes Sand that I have examined have
contained a few elasmobranch remains that are unidentifiable for one reason
or another (Fitch, 1964, 1966; Table 1). Teeth sometimes are too fragmented
or eroded to identify, and too little is known about vertebrae, wing spines
and bucklers to assign reliable specific names.
Total material: 27 vertebrae, 3 teeth, and 2 bucklers.
Teleosts
Allosmerus elongatus ( Ayres ) —whitebait smelt
Allosmerus elongatus has been reported as ranging from the Strait of
Juan de Fuca, Washington to San Pedro, but its occurrence south of Morro
16
Contributions in Science
No. 199
Bay is subject to doubt. It seldom is taken in the surf zone or at depths
greater than about 300 feet, but throughout much of this area, particularly
north of San Francisco, whitebait smelt are extremely abundant during most
years. The species is reported to attain a length of about nine inches, but of
many thousands that I have seen over the past 20 years, none has exceeded
six inches. The 9-inch fish probably was misidentified. Eleven otoliths from
the Crannell Road deposit (four in the Lloyd Barker collection) constitute
the only fossil record for the species. Sagittae of a large adult will exceed
3.5 mm in length.
Total material: 11 otoliths 2.7 to 3.6 mm (not figured).
Ammodytes hexapterus Pallas— Pacific sand lance
Pacific sand lance otoliths have been found in several Pliocene and Pleis-
tocene deposits in California (Fitch, 1968, and unpublished data), but until
now, they have not been reported from Palos Verdes Sand. If otoliths of this
Figure:!. Fish otoliths found in various deposits of “Palos Verdes Sand.” Lengths
(in mm) are given for each otolith; notations are made regarding its position in the
skull (left or right); otolith condition is noted if imperfect; and locality of fossil
deposit is given. All otoliths in this figure are sagittae, and all views are inner faces,
a. Spirinchus thaleichthys, 2.7, /, Crannell Road; b. Hypomesus pretiosus, 3.0, r,
Crannell Road; c. Spirinchus starksi, 3.5, r, Crannell Road; d. Oncorhynchus
tshawytscha, 11.8, r, Crannell Road; e. Anchoa compressa, 1.9, r, Playa del Rey;
f. Engraulis mordax, 4.0, /, V.T. Bridge; g. Argentina sialis, 3.0, r, Playa del Rey;
h. Sy nodus lucioceps, 3.0, r, Playa del Rey; i. Clupea pallasi, 3.5, r, Crannell Road;
j. Stenobrachius leucopsarus, 2.0, /, Playa del Rey; k. Symbolophorus calif orniensis,
4.4, r, Playa del Rey; 1. Electrona rissoi, 3.6, r, LACMIP 131, badly eroded;
m. Diaphus theta, 2.2, r, Crannell Road. Photographs by Jack W. Schott.
1970
Fish Remains from the Palos Verdes Sand
17
northern species had been found in southern California exposures of Palos
Verdes Sand, it could be deemed “unexpected” to say the least, but their
presence at the Crannell Road site is well within their normal present day
range.
Total material: 4 otoliths (Fig. 6/).
Amphistichus koelzi (Hubbs)— calico surfperch
The calico surfperch ranges from Cape Flattery, Washington, to Santo
Tomas Point, Baja California. It is a typical inhabitant of the sandy beach
surf zone, but occasionally strays into depths of 60 to 75 feet. The species
attains a length of about 10 inches and a pound in weight. Remains of A.
koelzi have not been found in any other deposits. Sagittae of a large adult
will exceed 7.0 mm in length.
Total material: 1 otolith 4.5 mm long (Fig. 5b).
Amphistichus rhodoterus (Agassiz)— redtail surfperch
The redtail surfperch ranges from Cape Flattery, Washington to Mon-
terey Bay. It also is a typical surf zone inhabitant that occasionally strays
into deeper water (to at least 60 feet). The species attains a length of 16
inches and nearly four pounds in weight. Remains of A. rhodoterus have
not been found in any other deposit; the Crannell Road exposure lies well
within their present-day range. The sagittae of a large adult will exceed
10.0 mm in length.
Total material: 1 partially digested otolith 6.6 mm long (Fig. 5d).
Anarrhichthys ocellatus Ayres— wolf -eel
The wolf-eel ranges from southeastern Alaska to Imperial Beach, Cali-
fornia, but is rare south of Point Conception. It prefers the rocky subtidal
and adjacent offshore shallow areas, but has been captured as deep as 400
feet. It is reported to attain eight feet, but the largest authentic record in
recent years was six feet eight inches and 40 pounds. Otoliths of this species
have not been found, but A. ocellatus has large, very distinctive teeth and
two of these were recovered from the Crannell Road deposit.
Total material: 2 teeth, including one in the Lloyd Barker collection.
Anchoa compressa (Girard ) — deepbody anchovy
Otoliths of A. compressa previously were reported from Playa del Rey
(Fitch, 1966), and no additional remains have been found.
Total material: 4 otoliths (Fig. \e).
Anisotremus davidsonii (Steindachner)— sargo
Fitch ( 1964) reported a sargo otolith from Playa del Rey that previously
had been noted by Kanakoff (1956); no additional remains of A. davidsonii
have been found.
Total material: 1 otolith (Fig. 4/).
18
Contributions in Science
No. 199
Argentina sialis Gilbert— Pacific argentine
Fitch (1966) reported a Pacific argentine otolith from Playa del Rey;
no additional remains have been found.
Total material: 1 otolith (Fig. 1^).
Atheresthes stomias (Jordan and Gilbert)— arrowtooth flounder
Otoliths of A. stomias have been found in Pliocene and Pleistocene
deposits of southern California (Fitch, 1968; and unpublished data) but
never in Palos Verdes Sand. Their teeth are large and have distinctive dart-
shaped tips. Two jaw teeth of A. stomias were recovered from the Crannell
Road deposit.
Total material: 2 teeth: one in the Lloyd Barker collection.
Atherinops affinis (Ayres)— topsmelt
The topsmelt previously was reported from Playa del Rey based upon
a single otolith (Fitch, 1964); one additional topsmelt otolith has been found
in a Palos Verdes Sand deposit in southern California (Table 2).
Total material: 2 otoliths (Fig. 4e).
Atherinopsis californiensis Girard— jacksmelt
Jacksmelt otoliths (4) previously were reported from Playa del Rey
(Fitch, 1964, 1966). Subsequent sampling has yielded two additional oto-
liths from Palos Verdes Sand in the San Pedro area (Table 2).
Total material: 6 otoliths (Fig. 4/).
Bairdiella icistia (Jordan and Gilbert) —bairdiella
The bairdiella ranges from the head of the Gulf of California to south-
ern Mexico, and it is one of the species that was successfully introduced into
Salton Sea. It is an inhabitant of relatively shallow (to about 150 feet) near-
shore waters in non-rocky habitat. The species seldom reaches a foot in
length or a pound in weight. Sagittae of a large adult will exceed 1 1.0 mm in
length. A single broken otolith was found in one of the San Pedro deposits
(Table 2).
Total material: 1 otolith 3.7 mm long ( Fig. 5q).
Calamus brachysomus (Lockington)— Pacific porgy
The pacific porgy has been recorded from Oceanside (twice) to 150
miles south of Lima, Peru, but it apparently does not spawn north of Sebas-
tian Viscaino Bay, Baja California. Young fish are abundant in shallow
water where the bottom is sandy or firm sandy mud whereas adults prefer
somewhat deeper water in the same type of habitat. A large individual will
exceed 20 inches in length and four pounds. Their flat-topped round molars
1970
Fish Remains from the Palos Verdes Sand
19
are easily recognized, and one of these was found in a Palos Verdes Sand
deposit at San Pedro (Table 2).
Total material: 1 tooth.
Chitonotus pugetensis (Steindachner)— staghorn sculpin
Staghorn sculpin otoliths have been found in a number of Pliocene and
Pleistocene deposits in southern California including Playa del Rey (Fitch,
1964; 1966). Aside from the Playa del Rey locality, the only other Palos
Verdes Sand exposure to yield C. pugetensis otoliths was in the San Pedro
area (Table 2).
Total material: 12 otoliths (Fig. 6/).
Citharichthys sordidus (Girard)— Pacific sanddab
Pacific sanddab otoliths have been found in many fossil deposits through-
out California, often in great numbers, and their occurrence in Palos Verdes
Sand has been reported at Playa del Rey (Fitch, 1964; 1966). Subsequent
sampling of Palos Verdes Sand has yielded C. sordidus otoliths in all areas
except Newport Beach (Table 2). Of 13 otoliths seen from the Crannell
Road deposit, three are in the Lloyd Barker collection.
Total material: 59 otoliths (Fig. 3>b).
Citharichthys stigmaeus Jordan and Gilbert— speckled sanddab
Speckled sanddab otoliths have been found in more deposits and more
abundantly than those of C. sordidus. Previously they have been reported
in Palos Verdes Sand at Playa del Rey (Fitch, 1964; 1966), but their occur-
rence in the San Pedro area and at Crannell Road (Table 2) has not been
noted previously. They have not been found in the Newport Beach exposures
of Palos Verdes Sand.
Total material: 504 otoliths (Fig. 3a).
Citharichthys xanthostigma Gilbert— longfin sanddab
Although longfin sanddab otoliths have been reported from the Pleisto-
cene of southern California (Fitch, 1968), and have been found in many
other Pliocene and Pleistocene deposits (unpublished data), only one expo-
sure of Palos Verdes Sand contained some (Table 2).
Total material: 4 otoliths (Fig. 3c).
Clupea pallasi Valenciennes— Pacific herring
Pacific herring otoliths have been found in several Pliocene and Pleis-
tocene deposits of southern California (Fitch, 1967, 1968; unpublished data),
but five sagittae from Crannell Road (Table 2, plus one in the Lloyd Barker
collection) represent the first record for the species from Palos Verdes Sand.
Total material: 5 otoliths (Fig. 1/).
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Figure 2. Fish otoliths found in various deposits of “Palos Verdes Sand.” Lengths
(in mm) are given for each otolith; notations are made regarding its position in the
skull (left or right); otolith condition is noted if imperfect; and locality of fossil
deposit is given. All otoliths in this figure are sagittae, and all views except Theragra
are inner faces, a. Porichthys myriaster, 13.0, /, LACMIP 131; b. Porichthys notatus,
6,0, r, V.T. Bridge; c. Lepophidium negropinna, 10.2, r, V.T. Bridge; d. Microgadus
proximus, 12.0, r, Crannell Road; e. Coelorhynchus scaphopsis, 7.9, 1, Playa del Rey;
f. Otophidium taylori, 7.9, I, V.T. Bridge; g. Theragra chalco gramma, 5.1, /, V.T.
Bridge, badly eroded and posterior end missing; h. Merluccius product us, 19.6, /,
Crannell Road; i. Otophidium scrippsi, 7.8, I, V.T. Bridge. Photographs by Jack W.
Schott,
Coelorhynchus scaphopsis (Gilbert)— Gulf rattail
Otoliths of the Gulf rattail have been found only in Palos Verdes Sand.
The species was previously reported by Fitch ( 1966) for Playa del Rey
based upon a single otolith. A second otolith subsequently was found in a
San Pedro exposure (Table 2).
Total material: 2 otoliths (Fig. 2e).
Cymatogaster aggregata Gibbons— shiner perch
Shiner perch otoliths previously have been reported from Palos Verdes
Sand at Playa del Rey (Fitch, 1964, 1966). Subsequently they have been
found at all San Pedro localities and Crannell Road, but not at Newport Beach
1970
Fish Remains from the Palos Verdes Sand
21
(Table 2). Lloyd Barker’s colleetion contained 17 otoliths from C. aggre-
gata.
Total material: 135 otoliths (Fig. 5/).
Cy noscion nobilis (Ayres)— white seabass
Five white seabass otoliths previously were reported from Palos Verdes
Sand at Playa del Rey (Kanakoff, 1956; Fitch, 1964). A single otolith sub-
sequently was found by museum personnel in one of the Newport Beach
localities (LACMIP 66; Table 2).
Total material: 6 otoliths (Fig. 5p).
Cynoscion reticulatus (Giinther)— striped corvina
Fossil remains of this southern species have been found only in Palos
Verdes Sand. They have been reported from Playa del Rey based upon 5 oto-
liths (Fitch, 1964, 1966), and six additional otoliths were found in two
deposits at San Pedro (Table 2).
Total material: 11 otoliths (Fig. 5r).
Diaphus theta Eigenmann and Eigenmann— California headlightfish
Otoliths of this mesopelagic lanternfish have been found in many Plio-
cene and Pleistocene deposits in southern California (Fitch, 1968, and un-
published data), but the single sagitta from the Crannell Road locality (Fitch,
1969a; Table 2) is the only record of D. theta from Palos Verdes Sand.
Total material: 1 otolith (Fig. Im).
Electrona rissoi (Cocco)— chubby flashlightfish
Otoliths of E. rissoi have been found in Pliocene (Fitch and Reimer,
1967) and Pleistocene deposits (Fitch, 1968), including Palos Verdes Sand
at San Pedro (Fitch, 1969a).
Total material: 1 otolith (Fig. 1/).
Embiotoca cf. jacksoni Agassiz— black perch
Otoliths of E. jacksoni have been found in a Pliocene deposit near San
Diego (unpublished data), but the single badly worn otolith from Palos
Verdes Sand at San Pedro (Table 2) represents the only record (a doubtful
one) from Pleistocene.
Total material: 1 badly eroded otolith (Fig. 5/).
Engraulis mordax Girard— northern anchovy
Northern anchovy otoliths were abundant in Palos Verdes Sand at
Playa del Rey (Fitch, 1964; 1966), and were present at two San Pedro
localities and Crannell Road, but none has been found at Newport Beach
(Table 2). The fossil record of E. mordax in California is unbroken over
the 10 million year span since the beginning of the Pliocene, including the
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most recent 7,000 years of the earth’s history as divulged by investigating
Indian middens (Fitch, 1969b).
Total material: 280 otoliths including one in the Lloyd Barker collection
(Fig. 1/).
Eopsetta jordani (Lockington)— petrale sole
Otoliths of E. jordani have been found in several Pleistocene deposits
(Fitch, 1967, and unpublished data), but they have not previously been
reported from Palos Verdes Sand. A single sagitta was found in the Crannell
Road deposit (Table 2), and three others are in the Lloyd Barker collection.
Total material: 4 otoliths (Fig, 3g).
Genyonemus lineatus (Ayres)— white croaker
Otoliths of G. lineatus were among the five most abundant sagittae at
Playa del Rey (Fitch, 1964, 1966), and have turned up at all other localities
where Palos Verdes Sand has been sampled (Kanakoff, 1956; Table 2).
Although white croakers have been reported as far north as Vancouver Island,
their occurrence north of San Francisco is spotty (Miller and Gotshali,
1965) and depends a great deal upon warm oceanic temperatures. The
sagittae in the Crannell Road deposit (including 2 in the Lloyd Barker
collection) were from large adults.
Total material: 740 otoliths (Fig. 51).
Glyptocephalus zachirus Lockington— rex sole
Rex sole remains have been found in an assortment of Pliocene and
Pleistocene deposits in California (Fitch, 1967, 1968, and unpublished data)
but not previously in Palos Verdes Sand. No otoliths of G. zachirus were
found in any of the southern California localities (Table 2) but two (one in
the Roy Kohl collection) were present in Crannell Road material examined.
Total material : 2 otoliths (Fig. 3/).
Hippoglossus stenolepis Schmidt— Pacific halibut
The Pacific halibut ranges from the Bering Sea to Point Piedras Blancas,
California, in depths of 60 to 3,600 feet. Males are known to reach a weight
of 123 pounds, and females, 495. Otoliths of H. stenolepis have never been
found in any fossil deposit, but in the Humboldt State College collection
there is a lower jaw of a fairly large individual that came from the Crannell
Road deposit.
Total material: 1 lower jaw in Humboldt State College collection.
Hyperprosopon anale Agassiz— spotfin surfperch
The spotfin surfperch is an inhabitant of the surf zone along sandy,
outer coast beaches between Seal Rock, Oregon, and Blanca Bay, Baja Cali-
fornia. Individuals occasionally are caught in depths of 60 feet or more
1970
Fish Remains from the Palos Verdes Sand
23
Figure 3. Fish otoliths found in various deposits of “Palos Verdes Sand.” Lengths
(in mm) are given for each otolith; notations are made regarding its position in the
skull (left or right); otolith condition is noted if imperfect; and locality of fossil
deposit is given. All otoliths in this figure are sagittae, and all views are inner faces,
a. Citharichthys stigmaeus, 2.8, I, V.T. Bridge; b. Citharichthys, sordidus, 6.7, /,
LACMIP 131; c. Citharichthys xanthostigma, 3.6, r, LACMIP 131; d. Paralichthys
californicus, 8.2, r, LACMIP 131; e. Pleuronichthys ritteri, 3.2, I, Playa del Rey;
f. Lyopsetta exilis, 3.5, r, Playa del Rey; g. Eopsetta jordani, 6.0, /, Crannell Road;
h. Parophrys vetuliis, 6.3, I, Crannell Road; i. Microstomus pacificus, 5.4, r, Cran-
nell Road; j. Glyptocephalus zachirus, 2.0, I, Crannell Road, badly eroded;
k. Platichthys stellatus, 9.5, I, Crannell Road; 1. Isopsetta isolepis, 6.3, r, Crannell
Road. Photographs by Jack W. Schott.
but they are not abundant at any depth. The largest fish reported was eight
inches long and weighed just five ounces. Their otoliths have been found in
several Pliocene and Pleistocene deposits (unpublished data), but only at
Crannell Road have they turned up in Palos Verdes Sand. One of seven
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otoliths from this site is in the Roy Kohl collection. Sagittae of a large adult
will exceed 6.5 mm in length.
Total material: 7 otoliths from 2.5 to 5.2 mm long (Fig. 5^).
Hyperprosopon ellipticum (Gibbons)— silver surfperch
The silver surfperch ranges from Clallam County, Washington, to Point
Dume, California. It prefers the surf zone along sandy, outer coast beaches,
but also frequents rocky subtidal pools and occasionally strays into depths
of 60 feet or more. They are known to attain a length of 10.5 inches and a
weight of about one-quarter pound. The two otoliths from the Crannell
Road deposit (Table 2) represent the only fossil record for this species.
Sagittae of a large adult will exceed 6.0 mm in length.
Total material: 2 otoliths 3.9 to 4.5 mm long (Fig. 5^).
Hypomesus pretiosus (Girard)— surf smelt
The surf smelt ranges from Prince William Sound, Alaska, to Long
Beach, California but it is rarely seen south of Morro Bay. It is an inhabitant
of sandy beaches along the outer coast, where it spawns during daylight
hours in the breaking surf and strays into depths of 60 feet or more when
not spawning. The species is known to reach a length of 10 inches and a
weight of about three ounces. Only one genus (Mallotus) in family Osmeridae
has a fossil record according to McAllister (1963). H. pretiosus is but one
of four osmerids from which otoliths were found in the Crannell Road deposit.
In addition to the 1 8 that I found, there are three in the Lloyd Barker collec-
tion. Sagittae of a large adult will exceed 5.0 mm in length.
Total material: 21 otoliths 2.6 to 3.4 mm long (Fig. \b).
Icelinus quadriseriatus ( Lockington ) — yellowchin sculpin
The yellowchin sculpin previously was reported from Palos Verdes Sand
at Playa del Rey based upon 10 otoliths (Fitch, 1966). No additional otoliths
have turned up since then.
Total material: 10 otoliths (Fig. 6k).
Icelinus tenuis Gilbert— spotfin sculpin
Otoliths of the spotfin sculpin have been found in a number of Pliocene
and Pleistocene deposits in southern California (Fitch, 1967, 1968, and
unpublished data), but they have not been reported previously from Palos
Verdes Sand. Two otoliths from one of the San Pedro sites were identified
as from this species (Table 2).
Total material: 2 otoliths (Fig. 6/).
Isopsetta isolepis ( Lockington ) — scaly-fin sole
The scaly-fin sole ranges from Alaska to southern California, but it is
rare south of about Morro Bay. It inhabits areas where the bottom is sandy
or sandy mud, and prefers depths shallower than about 150 feet although it
1970
Fish Remains from the Palos Verdes Sand
25
has been found deeper than 300. It is known to attain a length of 22 inches,
but no weights are available for such large individuals. The Crannell Road
site is the only locality where fossil remains of /. isolepis have been found.
In addition to the 21 otoliths that I found, I have seen two in the Lloyd
Barker collection and one in Roy Kohl’s. Sagittae of a large adult will exceed
9.0 mm in length.
Total material: 24 otoliths 1.4 to 6.3 mm long (Fig. 3/).
Lepidogobius lepidus (Girard)— bay goby
The bay goby has been recorded from Palos Verdes Sand at Playa del
Rey (Fitch, 1964, 1966), but the 28 otoliths obtained from San Pedro sites
(Table 2) have not been noted previously. This species has not been found
at Newport Beach or Crannell Road (Table 2).
Total material: 37 otoliths (Fig. 6g).
Lepophidium negropinna Hildebrand and Barton— giant cusk-eel
Remains of L. negropinna, a locally extinct southern species, have been
found only in the Palos Verdes Sand of southern California. The two sagit-
tae from Playa del Rey were reported by Fitch (1964), but the two from
San Pedro sites (Table 2) have not been previously noted.
Total material: 4 otoliths (Fig. 2c).
Leptocottus armatus Girard— staghorn sculpin
Staghorn sculpin remains have been found in several southern Cali-
fornia Pliocene and Pleistocene deposits (Fitch, 1967, 1968, and unpub-
lished data), but until now, only an opercular spine had been reported from
Palos Verdes Sand (Kanakoff, 1956). There were no otoliths of L. armatus
at Playa del Rey and few at San Pedro and Newport Beach sites (Table 2),
but the Crannell Road deposit yielded 13 otoliths and four opercular spines
(plus four otoliths and three spines in the Lloyd Barker collection and five
otoliths in Roy Kohl’s).
Total material: 22 otoliths and 7 opercular spines (Fig. 6m).
Lethops connectens Hubbs— halfblind goby
Otoliths of this species have been found in two southern California
Pleistocene deposits (Fitch, 1968, unpublished data), but not in Palos Verdes
Sand south of Monterey, which is the present-day northern limit of their
range. The Crannell Road site, several hundred miles north of their known
range, yielded seven otoliths of Lethops (Table 2).
Total material: 7 otoliths (Fig. 6h).
Leuresthes tenuis (Ayres)— grunion
Fossil grunion otoliths have been found only at Playa del Rey whence
they have been reported by Fitch (1964, 1966).
Total material: 10 otoliths (Fig. Ad).
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Figure 4. Fish otoliths found in various deposits of “Palos Verdes Sand.” Lengths
(in mm) are given for each otolith; notations are made regarding its position in the
skull (left or right); otolith condition is noted if imperfect; and locality of fossil
deposit is given. All otoliths in this figure are sagittae, and all views are inner faces:
a. Trachuriis symmetricus, 6.7, /, Playa del Rey, badly eroded and rostrum missing;
b. Paralabrax sp. 7.8, r. Play del Rey, anterior half missing; c. Symphurus atri-
cauda, 2.5, r, Playa del Rey; d. Leuresthes tenuis, 2.9, /, Playa del Rey; e. Atherinops
affinis, 3.1, /, V.T. Bridge, badly eroded and anterior tip missing; f. Atherinopsis
californiensis, 5.8, I, LACMIP 131; g. Xenistius calif orniensis, 9.0, /, Playa del Rey,
rostrum tip missing; h. Sphyraena argentea, 10.7, I, Playa del Rey, badly eroded
and anterior end missing; i. Anisotremus davidsonii, 9.8, r, Playa del Rey, eroded
and rostrum tip missing. Photographs by Jack W. Schott.
Lycodopsis pacifica ( Collett ) — blackbelly eelpout
Although sagittae of L. pacifica have been found in numerous Pliocene
and Pleistocene deposits of California (Fitch, 1967, 1968, and unpublished
data), only the Crannell Road site among the Palos Verdes Sand localities
investigated (Table 2) contained an otolith.
Total material: 1 otolith (Fig. 6o).
Lyopsetta exilis (Jordan and Gilbert)— slender sole
Slender sole otoliths are abundant in many Pliocene and Pleistocene
deposits, but the single otolith reported from Playa del Rey (Fitch, 1964) is
the only evidence of the species in Palos Verdes Sand.
Total material: 1 otolith (Fig. 3/).
Menticirrhus undulatus (Girard)— California corbina
Aside from the six otoliths reported from Playa del Rey (Fitch, 1964,
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Fish Remains from the Palos Verdes Sand
27
1966), only one additional sagitta has been found in deposits of Palos Verdes
Sand (Table 2).
Total material: 7 otoliths (Fig. 5k).
Merluccius productus (Ayres)— Pacific hake
Otoliths of Pacific hake (initially reported by Kanakoff, 1956) have
been present in almost every Pliocene and Pleistocene deposit that I have
investigated (Fitch, 1969b), but most have been from small fish. Except for
the Crannell Road deposit, where all five hake otoliths (Table 2) were from
large fish, the Palos Verdes Sand localities have yielded few large sagittae.
Twenty otoliths from Crannell Road in the Lloyd Barker and Roy Kohl col-
lections also are mostly from large fish.
Total material: 61 otoliths (Fig. 2h).
Microgadus proximus (Girard)— Pacific tomcod
Pacific tomcod otoliths have been found in numerous Pliocene and
Pleistocene deposits of California (Fitch, 1967, 1968, and unpublished data;
Fitch and Reimer, 1967), but only at the Crannell Road site are they abun-
dant. This deposit, the only Palos Verdes Sand to contain them, yielded 411
sagittae to my efforts (Table 2), and 623 others to Lloyd Barker and Roy
Kohl.
Total material: 1,034 otoliths (Fig. Id).
Micropogon ectenes Jordan and Gilbert— berrugato
Fossil otoliths from this locally extinct southern species have been found
only in Palos Verdes Sand: two at Playa del Rey (Fitch, 1964) and one at
San Pedro (Table 2).
Total material: 3 otoliths (Fig. 5m).
Microstomus pacificus ( Lockington ) —Dover sole
Sagittae of M. pacificus have been rare in the Pliocene and Pleistocene
deposits that I have examined. Prior to finding two at the Crannell Road
locality (Table 2), only Timms Point Silt had yielded an otolith (Fitch,
1968). An additional Dover sole otolith was seen in the Lloyd Barker col-
lection.
Total material: 3 otoliths (Fig. 3i).
Oncorhynchus tshawytscha (Walbaum)— King salmon
King salmon have been taken in the Pacific Ocean from San Diego to
the Bering Sea and south, on the Asiatic side, to Japan. They are seldom
seen or caught south of Morro Bay, however. They are known to attain
lengths of at least five feet and a weight of 126.5 pounds, but a 50-pounder
is rare enough to generate publicity when brought ashore. Aside from a very
questionable record of O. tshawytscha from the Pleistocene of Oregon, based
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upon fragments of jaws, teeth and vertebrae (Uyeno and Miller, 1963), the
otolith from the Crannell Road locality (Table 2) represents the only fossil
occurrence of this important anadromous fish. Sagittae from a large adult
king salmon will exceed 15.0 mm in length.
Total material: 1 otolith 11.8 mm long (Fig. \d).
Ophiodon elongatus Girard— lingcod
The lingcod ranges from northwestern Alaska to San Carlos anchorage,
Baja California, but is not abundant south of Point Conception except on a
few offshore banks. It appears to prefer rocky bottom areas and depths shal-
lower than about 350 feet, but many are caught on sandy or sandy-mud
bottoms, and to depths of 2,700 feet. Lingcod reportedly attain lengths ex-
ceeding five feet and weights over 70 pounds, but 40-pound fish are rare off
California. The strong conical jaw teeth of large O. elongatus are distinctive
among California’s marine fishes, and 1 1 of the teleost teeth I found in the
Crannell Road deposit had come from this species. An additional 12 lingcod
teeth were seen in the Lloyd Barker collection.
Total material: 23 jaw teeth.
Ophioscion sp.— sciaenid
Several species of Ophioscion, smallish sciaenids that seldom attain a
foot in length, are inhabitants of the tropical eastern Pacific. In this genus, two
of the three pairs of otoliths (sagittae and lapilli) are almost identical in
size. Except among ostariophysans, teleost lapilli from a given fish are almost
invariably microscopic compared with sagittae from the same individual.
Thus, the lapilli of Ophioscion are easily identified to genus by their size and
configuration, but I have been unable to find characters among several species
that will permit a specific determination. O. scierus has been noted at Mag-
dalena Bay on the outer coast of Baja California during historic times, but
this is not sufficient reason to name the lapillus found in Palos Verdes Sand
at San Pedro.
Total material: 1 lapillus 6.6 mm long (Fig. 5n).
Otophidium scrippsi Hubbs— basketweave cusk-eel
Otoliths of O. scrippsi have been among the most abundant teleost re-
mains in many southern California Pliocene and Pleistocene deposits (Fitch,
1964, 1966, and unpublished data; Fitch and Reimer, 1967; Kanakoff, 1956).
They were found at all Palos Verdes Sand localities except Crannell Road
(Table 2).
Total material: 275 otoliths (Fig. 2i).
Otophidium taylori (Girard)— spotted cusk-eel
Otoliths of O. taylori, an inhabitant of more northerly and deeper water
than O. scrippsi, have been found in even more Pliocene and Pleistocene
deposits than O. scrippsi, but were missing at Crannell Road (Table 2). The
1970
Fish Remains from the Palos Verdes Sand
29
first fossil occurrence of these was noted by Kanakoff (1956),
Total material: 185 otoliths (Fig. 2/).
Paralabrax spp.—sand or kelp bass
Three Paralabrax otoliths, reported from Palos Verdes Sand at Playa del
Rey by Fitch (1966), are the only serranid remains found in the Pliocene
or Pleistocene of southern California to date.
Total material: 3 otoliths (Fig. 4b).
Paralichthys californicus (Ayres) —California halibut
California halibut otoliths have been found only at Signal Hill (Fitch,
1967), Playa del Rey (Fitch, 1964, 1966) and two San Pedro sites (Table 2).
Total material: 10 otoliths (Fig. 3d).
Parophrys vetulus Girard— English sole
Otoliths of P. vetulus have been found in many Pliocene and Pleistocene
deposits including Palos Verdes Sand (Fitch, 1964, 1966, 1968; Fitch and
Reimer, 1967). None was found in Palos Verdes Sand at San Pedro, but one
turned up at Newport Beach and 14 at Crannell Road (Table 2). An addi-
tional 12 were seen in the Lloyd Barker collection.
Total material: 31 otoliths (Fig. 3h).
Phanerodon furcatus Girard— white seaperch
Otoliths of P. furcatus have not been found often, nor have they been
abundant (unpublished data). I have found only one in Palos Verdes Sand
(Crannell Road, Table 2), although the Lloyd Barker and Roy Kohl collec-
tions also contain white seaperch otoliths (3) from Crannell Road. The nine
otoliths from the Playa del Rey locality that I reported as Phanerodon (Fitch,
1964) were misideetified. A careful re-examination of these has revealed that
seven of the nine were from Xenistius calif orniensis and the other two (small
and badly eroded) from Trachurus symmetricus.
Total material: 13 otoliths (Fig. 5c).
Pimelometopon pulchrum (Ayres)— California sheephead
The distinctive jaw and pharyngeal teeth of P. pulchrum have been
found in many fossil deposits, but to date, no sheephead otoliths have turned
up. Teeth previously were reported from Palos Verdes Sand at Newport Beach
(Kanakoff, 1956), and Playa del Rey (Fitch, 1964), but those from San
Pedro (Table 2) are being noted for the first time. None has been found at
Crannell Road.
Total material: 28 teeth.
Platichthys stellatus (Pallas)— starry flounder
The starry flounder ranges throughout the north Pacific, having been
recorded south to Santa Barbara in the eastern Pacific, but generally north
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of Morro Bay. They prefer sandy or sandy mud bottoms and have been caught
in fresh, brackish, and salt water. In the ocean, they are most abundant in
depths shallower than 200 feet, but some individuals have been caught five
times that deep. They are reported to reach a length of three feet and a weight
of 20 pounds, but few are seen that exceed 15. A single otolith in the Roy Kohl
collection, plus five stellate scales (two in Lloyd Barker’s collection) are the
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Fish Remains from the Palos Verdes Sand
31
Figure 5. Surfperch and croaker otoliths found in various deposits of “Palos Verdes
Sand.” Lengths (in mm) are given for each otolith; notations are made regarding
its position in the skull (left or right); otolith condition is noted if imperfect; and
locality of fossil deposit is given. All otoliths in this figure except Ophioscion (a
lapillus) are sagittae, and all views are inner faces, a. Rhacochiliis vacca, 8.1, r,
LACMIP 131, badly eroded and portions missing from both ends; b. Amphistichus
koelzi, 4.5, r, V.T. Bridge, portions missing from rostrum and posterior end;
c. Phanerodon furcatus, 8.6, I, Crannell Road; d. Amphistichus rhodoterus, 6.6, I,
Crannell Road, badly eroded; e. Hyperprosopon ellipticum, 3.9, /, Crannell Road;
f. Embiotoca cf. jacksoni, 6.0, /, V.T. Bridge, badly eroded; g. Hyperprosopon ancde,
5.2, /, Crannell Road; h. Roncador stearnsi, 9.4, /, LACMIP 131, badly eroded;
I. Cymatogaster aggregata, 6.0, /, V.T. Bridge; j. Seriphus politus, 7.9, I, V.T.
Bridge; k. Menticirrhiis iindidatus, 5.1, r, Playa del Rey; 1. Genyonemus lineatus,
II. 5, I, Crannell Road; m. Micropogon ectenes, 7.8, 1, V.T. Bridge; n. Ophioscion
sp., 6.6, r, LACMIP 131; o. Umbrina roncador, 10.9, r, LACMIP 131; p. Cy noscion
nobilis, 21.5, r, Playa del Rey; q. Bairdiella icistia, 3.7, r, LACMIP 131, ventral
margin missing; r. Cynoscion reticulatiis, 8.7, /, V.T. Bridge. Photographs by
Jack W. Schott.
only fossil record of this species. All of these remains are from the Crannell
Road deposit. Sagittae of a large adult will exceed 12.0 mm in length.
Tot3l material: 1 otolith 9.5 mm long (Fig. 3Ar) and 5 scales.
Pleuronichthys ritteri (Starks and Morris)— spotted turbot
The single otolith of P. ritteri that I reported from Playa del Rey (Fitch,
1964) is the only known fossil record of this species.
Total material: 1 otolith (Fig. 3c).
Porichthys myriaster Hubbs and Schultz— specklefin midshipman
Otoliths of P. myriaster have been found in every Palos Verdes Sand
locality, except Crannell Road, and were present in several other Pliocene
and Pleistocene exposures in southern California (Fitch and Reimer, 1967;
Fitch, unpublished data). In addition to the otoliths listed here (Table 2),
the Los Angeles County Museum collections contain a disarticulated, but
nearly complete, skull including otoliths that came from a Newport Beach
exposure of Palos Verdes Sand (LACMIP 136).
Total material: 47 otoliths (Fig. 2a) and a disarticulated skull.
Porichthys notatus Girard— plainfin midshipman
Otoliths of P. notatus are one of the five most abundant teleost remains
in Palos Verdes Sand (Table 2) having been reported from Playa del Rey
(Fitch, 1964, 1966), but not previously from San Pedro or Crannell Road.
They are common to extremely abundant in numerous other Pliocene and
Pleistocene exposures throughout California (Fitch, 1967, 1968, and un-
published data; Fitch and Reimer, 1967). Two of the three I have seen from
Crannell Road are in Lloyd Barker’s collection.
Total material: 504 otoliths (Fig. 2b).
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Prionotus riiscarius Gilbert and Starks— shortfin searobin
P. ruscarius has never been caught north of Magdalena Bay, Baja Cali-
fornia; it ranges from there south to Panama. It appears to prefer moderate
depths, possibly 20 to 150 feet, in areas of sandy or sandy mud bottom. Short-
fin searobins are reported to attain a foot in length and possibly a pound in
weight. A single otolith in Palos Verdes Sand at San Pedro (Table 2) is the
only known fossil record of this species. Sagittae of a large adult will exceed
7.5 mm in length.
Total material : 1 otolith 6.6 mm long (Fig. 6p).
Prionotus stephanophrys Lockington— lumptail searobin
In a previous report (Fitch, 1964), I erroneously identified an otolith
P. stephanophrys as Sebastodes {= Sebastes\ aurora. Subsequently (Fitch,
1966), I noted two additional lumptail searobin otoliths from Playa del Rey,
but the sagitta from San Pedro (Table 2) has not been reported until now.
Fossil remains of P. stephanophrys are known only from Palos Verdes Sand.
Total material: 4 otoliths (Fig. 6n).
Rhacochilus vacca (Girard)— pile perch
Pile perch remains have been found in several Pliocene and Pleistocene
deposits in southern California (Fitch, 1967, and unpublished data) , but
until now they have not been reported from Palos Verdes Sand. Their massive,
straight sided pharyngeal teeth are very distinctive, and two of these were
found at the Crannell Road site and one at San Pedro (Table 2). The only
pile perch otolith in Palos Verdes Sand came from one of the San Pedro
localities.
Total material: 1 otolith (Fig. 5a) and 3 pharyngeal teeth.
Roncador stearnsi (Steindachner) — spotfin croaker
Spotfin croaker otoliths have been found only in the Signal Hill Pliocene
(Fitch and Reimer, 1967), and in Palos Verdes Sand (Fitch, 1964; Table 2).
Total material : 22 otoliths (Fig. 5h).
Sebastes diploproa (Gilbert)— splitnose rockcod
Sebastes jordani ( Gilbert) — shortbelly rockcod
Sebastes paucispinis Ayres— bocaccio
Sebastes rhodochloris (Jordan and Gilbert) —swordspine rockcod
Otoliths from these four species of Sebastes have all been reported (as
Sebastodes) from Palos Verdes Sand at Playa del Rey (Fitch, 1964). All
except S. paucispinis were represented by a single otolith (there were 6 of
paucispinis) , and all except paucispinis have been found in other Pliocene
1970
Fish Remains from the Palos Verdes Sand
33
and Pleistocene deposits of southern California (unpublished data), but not
in other exposures of Palos Verdes Sand (Table 2).
Total material: 6 otoliths of S. paucispinis and 1 each of the other three species
(Figs. 6b to 6e).
Se bastes serranoides (Eigenmann and Eigenmann ) —olive rockcod
The olive rockcod ranges from Crescent City to Cape Colnett, Baja
California, in depths to 480 feet, but it is rare north of San Francisco. The
species is reported to attain a length of 24 inches and a weight of seven to
eight pounds, but no fish of that size has been measured or weighed. A single
otolith found by Roy Kohl at the Crannell Road deposit is the only known
fossil record of this species. Sagittae of a large adult will exceed 20.0 mm in
length.
Total material : 1 otolith 21.0 mm long (Fig. 6a).
Seriphus politus Ayres— queenfish
Queenfish otoliths have been abundant in many Pliocene and Pleistocene
deposits throughout southern California (Fitch, 1967, 1968, and unpublished
data; Fitch and Reimer, 1967) and were particularly common in Palos Verdes
Sand at Playa del Rey (Fitch, 1964, 1966). Subsequently, otoliths of S. politus
have been found in all other exposures of Palos Verdes Sand including Cran-
nell Road (Table 2). The single otolith from the Crannell Road deposit was
tiny and badly eroded, however.
Total material : 514 otoliths (Fig. 5/).
Sphyraena argentea Girard— California barracuda
Otoliths of S. argentea have been found in two Pliocene deposits of
southern California (unpublished data), but the single badly eroded otolith
from Palos Verdes Sand at Playa del Rey (Fitch, 1964) is the only record
of the species in the Pleistocene.
Total material: 1 otolith (Fig. Ah).
Spirinchus starksi (Fisk)— night smelt
Otoliths of S. starksi have been found in several Pleistocene deposits of
southern California (Fitch, 1967, and unpublished data), but until now, they
have not been reported from Palos Verdes Sand. The material I examined
from Crannell Road contained 124 night smelt otoliths and many others
were seen in the Lloyd Barker and Roy Kohl collections.
Total material: 177 otoliths (Fig. Ic).
Spirinchus thaleichthys (Ayres)— longfin smelt
The longfin smelt ranges from Hinchinbrook Island, Prince William
Sound, Alaska, to San Francisco. It spawns in fresh water, but when not
spawning, it is found in shallow to moderate depths outside the surf zone
along the open coast. It is known to attain a length of just over six inches, and
34
Contributions in Science
No. 199
Figure 6. Fish otoliths found in various deposits of “Palos Verdes Sand.” Lengths
(in mm) are given for each otolith; notations are made regarding its position in the
skull (left or right); otolith condition is noted if impeirfect; and locality of fossil
deposit is given. All otoliths in this figure are sagittae, and all views are inner faces,
a. Sebastes sermnoides, 21.0, I, Crannell Road; b. Sebastes rhodochloris, 7.0, r,
Playa del Rey; c. Sebastes diploproa, 12.9, /, Playa del Rey; d. Sebastes paucispinis,
7.3, /, Playa del Rey, badly eroded and pieces missing from both ends; e Sebastes
jordani, 6.4, /, Playa del Rey; f. Ammodytes hexapterus, 3.0, r, Crannell Road;
g. Lepidogobius lepidus, 2.3, /, V.T. Bridge; h. Lethops connectens, 1.4, /, Crannell
Road; i. Zaniolepis latipinnis, 3.3, /, Playa del Rey, badly eroded; j. Icelinus tenuis,
3.8, /, V.T. Bridge; k. Icelinus quadriseriatus, 2.8, /, Playa del Rey, badly eroded;
1. Chitonotus pugetensis, 4.6, I, Playa del Rey; m. Leptocottus armatus, 7.5, I, Cran-
nell Road; n. Prionotus stephanophrys, 6.8, r, Playa del Rey; o. Lycodopsis pacifica,
3.2, /, Crannell Road; p. Prionotus ruscarius, 6.6, I, LACMIP, 131, badly eroded.
Photographs by Jack W. Schott.
1970
Fish Remains from the Palos Verdes Sand
35
a weight of slightly more than one ounce. The five otoliths I found in the
Crannell Road deposit constitute the only known fossil record for this species.
Sagittae of a large adult will exceed 4.0 mm in length.
Total material: 5 otoliths 2.7 to 3.8 mm long (Fig. la).
Stenohrachiiis leucopsarus (Eigenmann and Eigenmann)— northern lampfish
The single otolith of S. leucopsarus from Playa del Rey (Fitch, 1966) is
still the only record of this species from Palos Verdes Sand although its oto-
liths have been present in many other Pliocene and Pleistocene deposits and
abundant in some (Fitch, 1967, 1968, and unpublished data).
Total material: 1 otolith (Fig. 1/).
Symbolophorus californiensis (Eigenmann and Eigenmann)— California lan-
ternfish
Otoliths from S. californiensis have been found in several Pliocene and
Pleistocene deposits in southern California including Palos Verdes Sand at
Playa del Rey and Crannell Road (Fitch, 1964, 1969a). A single otolith also
was found at one of the San Pedro sites (Fitch, 1969a; Table 2). The sagitta
from Crannell Road is broken, and only the posterior half was found, but
there is no question regarding its identification.
Total material: 3 otoliths (Fig. Ik).
Symphurus atricauda (Jordan and Gilbert)— California tonguefish
Two additional otoliths of S. atricauda have been found in Palos Verdes
Sand at San Pedro (Table 2) since I first reported the species at Playa del Rey
(Fitch, 1966). These 15 otoliths constitute the only fossil record of the
species.
Total material: 15 otoliths (Fig. 4c).
Sy nodus lucioceps (Ayres)— California lizardfish
The California lizardfish has been reeorded from San Francisco Bay to
Cape San Lucas and throughout much of the Gulf of California. Larvae and
young individuals up to about three inches live in the upper water layers,
often over deep water many miles from shore. Adults live at or near the
bottom in sandy areas, usually in depths of 60 to 150 feet but sometimes
shallower or deeper. The largest individual known was 25 Vs inches long and
weighed just over four pounds. A single otolith from S. lucioceps was found
at Playa del Rey; it was erroneously identified as Occa verrucosa in a previous
report (Fitch, 1966). In Occa the cauda (posterior portion of the sulcus or
groove on the inner face of the otolith) terminates farther forward than it
does in Synodus, and this differentiating character sometimes is difficult to
distinguish in eroded, worn, or broken otoliths. Sagittae of a large adult S.
lucioceps will exceed 8.0 mm.
Total material: 1 otolith 3.0 mm long (Fig. 1/r).
36
Contributions in Science
No. 199
Theragra chalcogramma (Pallas) —walleye pollock
Four walleye pollock otoliths from Early Pleistocene in the cold-water
Timms Point Silt (Fitch, 1968) and a badly eroded, broken fragment from
the Palos Verdes Sand at San Pedro (Table 2) constitute the only fossil rec-
ord for this northern species. The only reasonable explanation for its presence
in a warm-water deposit would entail predation offshore in deep (cold) water,
erosion by digestive action in the alimentary tract of the predator and excre-
tion in shallow (warm) water several hours after ingestion. Otoliths of prey
species often are broken while being eaten, and erosion of all exposed surfaces
including concave areas can be accomplished only by digestive action.
Total material: 1 eroded otolith fragment (Fig. Ig).
Trachurus symmetricus (Ayres) —Pacific jack mackerel
The three otoliths reported from Playa del Rey (Fitch, 1966) and two
that I misidentified as Phanerodon furcatus (Fitch, 1964) constitute the only
record of T. symmetricus from Palos Verdes Sand although jack mackerel
sagittae have been present in several other Pliocene and Pleistocene deposits
(Fitch, 1967, 1968, and unpublished data).
Total material: 5 otoliths (Fig. Aa).
Umbrina roncador Jordan and Gilbert— yellowfin croaker
Remains of U. roncador have been found only in the San Diego Fm.
(Pliocene) and in Palos Verdes Sand. The 17 otoliths from Playa del Rey
(Table 2) have been reported (Fitch, 1964, 1966), but those from the San
Pedro site have not been noted previously.
Total material: 23 otoliths (Fig. So).
Xenistius californiensis (Steindachner) — salema
The salema ranges from Monterey Bay to Peru, but it seldom is present
off California except during warm water years and even then it usually is not
abundant. It is an inhabitant of shallow water (to perhaps 60 feet) along the
outer coast, but sometimes enters bays and lagoons. Salemas are reported to
attain a length of 10 inches, but no weights are available for fish that size.
Seven otoliths from Playa del Rey, erroneously identified as Phanerodon fur-
catus in my earliest report on that site (Fitch, 1964), constitute the only fossil
record of this species. When in perfect condition, sagittae of adult X. califor-
niensis differ from P. furcatus in many respects, but the two best characters on
the worn fossils (i.e., posterior otolith taper and angle of downward flexure
of the cauda) were overlooked in making my earlier determinations. Sagittae
of a large adult will exceed 9.0 mm in length.
Total material: 7 otoliths 6.6 to 9.7 mm long (Fig.4g).
Zaniolepis latipinnis Girard— longspine combfish
The single otolith of Z. latipinnis reported from Palos Verdes Sand at
Playa del Rey (Fitch, 1966) is still the only fossil record of the species.
Total material: 1 otolith (Fig. 6/).
1970
Fish Remains from the Palos Verdes Sand
37
Unidentified teleosts
Many otoliths are so badly broken or worn that they cannot be identified,
even to family. Many others can be placed in a particular family but cannot
be identified as to genus, and still others can be given generic, but not specific,
names. Juvenile rockcod otoliths (Sebastes spp.) are almost impossible to
assign to species, yet they are easy to distinguish from Sebastolobus and Scor-
paena, the other two commonest genera in the family.
Forty-three otoliths found in various exposures of Palos Verdes Sand
were too broken or worn to assign any taxon (Table 2). Twenty-three otoliths
and nine teeth could be placed in three families (atherinids: 11 otoliths; cot-
tids: 2 otoliths; and embiotocids: 10 otoliths and 9 teeth) but could not be
assigned to genera within these families. Finally, 131 otoliths unquestionably
belonged to four genera {Citharichthys, Otophidium, Porichthys and Sebas-
tes) but could not be identified to species. For the first three named genera
no additional species could have resulted, but if additional Sebastes otoliths
could have been identified, it could have increased the faunal list for Palos
Verdes Sand. As an example, at Crannell Road only one species of Sebastes
could be named, but the 20 unnamed otoliths (six in Lloyd Barker’s collec-
tion) were unquestionably from at least four additional species. Some, if not
all of these, very likely were different from the four named species found in
the Play a del Rey area (Table 2).
Total material: 197 otoliths and 9 teeth.
Discussion
Three of the 18 (at least) kinds of sharks, skates and rays reported from
Palos Verdes Sand of southern California (Table 1) are “southern” species.
Rhizoprionodon has been recorded only once north of Mexico, but Carch-
arhinus and Sphyrna are observed or caught in our coastal waters during most
periods when ocean temperatures are considerably warmer than normal
(Radovich, 1961). The other 15 kinds of elasmobranchs are year around resi-
dents of the ocean off California, but some of these are not present during all
months and others are never abundant.
Since 10 of the 59 teleosts from the southern Californian deposits of
Palos Verdes Sand are southern species and three are mesopelagics, one would
not expect to capture these in routine netting operations or inside the 100-
fathom curve. The 46 remaining species are year around residents, however,
and all but three of these {Pimelometopon, Sebastes rhodochloris and The-
ragra) were components of beach seine and trawl catches made off southern
California during recent years (Carlisle, Schott and Abramson, 1960; Car-
lisle, 1969; Table 3). Only The ragra, of the three not captured in these
netting operations, is not normally taken south of Point Conception. Pimelo-
metopon and Sebastes rhodochloris are common in offshore rocky habitat
which is unsuitable for trawling.
38
Contributions in Science
No. 199
Six of the 10 southern teleosts {Bairdiella, Cy noscion reticulatus, Lepo-
phidium, Micropogon, Ophioscion and Prionotus ruscarius) have not been
captured within several hundred miles of California during modern times
(Table 4). Of the remaining four, Xenistius and Prionotus stephanophrys
have been noted on many occasions, whereas, neither Calamus nor Coelo-
rhynchus has been reported north of Mexico more than two or three times.
Table 3
Teleosts reported from Palos Verdes Sand which comprised
portions of beach seine and trawl catches
Method and depth of capture
Beach seine only ^
< 1 0 feet
Beach seine and trawC’ “
Maximum depth (in feet)
noted
Trawl only 2
Minimum depth
(in feet) noted
Amphistichus koelzi Citharichthys sordidus
528
Argentina sialis
120
Anchoa compressa
Citharichthys stigmaeus
540
Chitonotus
pugetensis
126
Anisotremus
davidsonii
Cymatogaster aggregata 346
Citharichthys
xanthostigma
60
Atherinops affinis
Embiotoca jacksoni
60
Icelinus
quadriseriatus
60
Atherinopsis
calif orniensis
Engraulis mordax
600
Icelinus tenuis
186
Cynoscion nobilis
Genyonemus lineatus
426
Lyopsetta exilis
60
Leptocottus armatus
Lepidogobius lepidus
300
Merluccius
productus
360
Leuresthes tenuis
Paralabrax spp.
120
Otophidium scrippsi
60
Menticirrhus
undulatus
Paralichthys
californicus
436
Otophidium toy lor i
60
Roncador stearnsi
Pleuronichthys r Uteri
285
Parophrys vetulus
60
Sphyraena argentea
Porichthys myriaster
387
Porichthys notatus
60
Umbrina roncador
Rhacochilus vacca
60
Sebastes diploproa
180
Seriphus politus
126
Sebastes jordani
378
Symphurus atricauda
563
Sebastes paucispinis
Sy nodus lucioceps
120
60
Trachurus symmetricus
600
Zaniolepis latipinnis
118
^ from Carlisle, Schott and Abramson ( 1960)
■ from Carlisle ( 1969)
1970
Fish Remains from the Palos Verdes Sand
39
Table 4
Fish species whose fossil record is exclusively in
deposits of Palos Verdes Sand
Southern Californian exposures
Crannell Road (Northern California)
Alopias vulpinus
A llosmerus elongatus
Dasyatis dipterurus
Arnphistichus rhodoterus
Notorynchus maculatus
Anarrhichthys ocellatus
Rhizoprionodon longurio^
Hippoglossus stenolepis
Arnphistichus koelzi
Hyperprosopon ellipticum
Anchoa compressa
Hypomesus pretiosus
Anisotremus davidsonii
Isopsetta isolepis
Argentina sialis
Onocorhynchus tshawytscha
Bairdiella icistia"^^
Ophiodon elongatus
Calamus brachysomus*
Platichthys stellatus
Coelorhynchus scaphopsis*
Sebastes serranoides
Cynoscion reticulatus* i
Lepophidium negropinna'*^
Leuresthes tenuis
Menticirrhus undulatus
Micropogon ectenes^i
Ophioscion sp."'f
Paralabrax sp.
Pleuronichthys ritteri
Prionotus ruscarius*^
Prionotus stephanophrys^
Symphurus atricauda
Synodus lucioceps
Xenistius californiensis'^
Zaniolepis latipinnis
Spirinchus thaleichthys
* southern species
t locally-extinct southern species
In our area, Pionotus and Coelorhynchus are found in depths exceeding 60
feet, but Xenistius and Calamus reside in relatively shallow near-shore waters.
Five of the 37 teleosts found in the northern California equivalent of
Palos Verdes Sand (Crannell Road) are southern species for that latitude
(i.e., Genyonemus, Hyperprosopon anale, Lethops, Sebastes serranoides and
Symbolophorus) , but only Lethops has not been reported that far north (by
several hundred miles) during modern times. Except for the two mesopelagics
{Diaphus and Symbolophorus) , all of the fishes identified from the Crannell
Road deposit can be captured at depths shallower than 120 feet. Diaphus has
40
Contributions in Science
No. 199
been netted off northern California in 35 fms (210 feet), but Symbolophorus
generally inhabits waters outside the continental shelf.
In my investigations of Plio-Pleistocene fishes of California I have iden-
tified the remains of 23 kinds (at least) of elasmobranchs and 150 species
of teleosts. Some species (e.g., Citharichthys, Genyonemus, Porichthys, Seri-
phus, etc.) have turned up in almost every deposit I have examined, but others
have been in only one stratum, or at one particular site to the exclusion of
all others. Thirty-seven fishes, including four elasmobranchs, are found ex-
clusively in Palos Verdes Sand, including the northern Californian equivalent
(Table 4), but only the 1 1 southern forms could be considered indicator
species for this period of the Pleistocene.
Critical examination of additional deposits of Palos Verdes Sand in
northern California undoubtedly would lengthen the list of species (Table 4)
exclusive to this stratum, but examination in northern California and Oregon
of Pliocene and Pleistocene deposits representing other time periods would
shorten the Crannell Road list of “exclusives” by possibly 50 percent.
Acknowledgments
My work with fossil otoliths has been supported by research grants
(GB-1244 and GB-6490) from the National Science Foundation. In addition
to the numerous student workers who previously were acknowledged for their
help with fossiliferous material from Playa del Rey, I wish to thank Lloyd
Barker, Dan Gotshall, Percy Hollister, Jack Hopkins, Roy Kohl and William
Zinnsmeister. These six individuals accompanied me in the field, collected
material at my request, permitted me to examine and borrow specimens in
their collections and were helpful in several other phases of investigation.
Richard A. Fitch washed, screened and sorted much of the fossiliferous ma-
trix; Shelton P. Applegate checked my elasmobranch identifications for accu-
racy; and Robert J. Lavenberg supplied helpful advice and comparative
material (otoliths) throughout the investigation and reviewed the completed
manuscript.
Jack W. Schott took the excellent otolith photographs, Larry Reynolds
applied his photo processing skills on the otolith prints and Mrs. Loretta
Proctor typed the final draft of the manuscript.
Literature Cited
Arnold, Ralph. 1903. The paleontology and stratigraphy of the marine Pliocene
and Pleistocene of San Pedro, California. Mem. Calif. Acad. Sci. 3:1-420.
Carlisle, John G., Jr. 1969. Results of a six-year trawl study in an area of heavy
waste discharge: Santa Monica Bay, California. Calif. Fish and Game
55(l):26-46.
Carlisle, John G., Jr., J. W. Schott, and N. J. Abramson. 1960. The barred
surfperch {Amphistichus argenteus Agassiz) in southern California. Calif.
Dept. Fish and Game, Fish Bull. 109: 1-79.
1970
Fish Remains from the Palos Verdes Sand
41
Fanale, F. P., and O. a. Schaeffer. 1965. Helium-uranium ratios for Pleistocene
and Tertiary fossil aragonites. Science 149(3681 ) :312-317.
Fitch, John E. 1964. The fish fauna of the Playa del Rey locality, a southern
California marine Pleistocene deposit. Los Angeles Co. Mus., Contrib. in Sci.
82:1-35.
1966. Additional fish remains, mostly otoliths, from a Pleistocene deposit
at Playa del Rey, California. Los Angeles Co. Mus., Contrib. in Sci. 119:1-16.
1967. The marine fish fauna, based primarily on otoliths, of a Lower
Pleistocene deposit at San Pedro, California (LACMIP 332, San Pedro Sand).
Los Angeles Co. Mus., Contrib. in Sci. 128:1-23.
1968. Otoliths and other fish remains from the Timms Point Silt (early
Pleistocene) at San Pedro, California. Los Angeles Co. Mus., Contrib. in Sci.
146:1-29.
1969a. Fossil lanternfish otoliths of California, with notes on fossil
Myctophidae of North America. Los Angeles Co. Mus., Contrib. in Sci.
173:1-20.
1969b. Fossil records of certain schooling fishes of the California Cur-
rent System. Calif. Mar. Res. Comm., CalCOFI Rept. 13:71-80.
Fitch, John E., and Roger D. Reimer. 1967. Otoliths and other fish remains from
a Long Beach, California, Pliocene deposit. So. Calif. Acad. Sci., Bull.
66(2):77-91.
Gilbert, Carter R. 1967. A revision of the hammerhead sharks (Family Sphyr-
nidae). U.S. Natl. Mus., Proc. 1 19:1-88.
Jordan, D. S., and Harold Hannibal. 1923. Fossil sharks and rays of the Pacific
slope of North America. So. Calif. Acad. Sci., Bull. 22:27-63, 65-68.
Kanakoff, George P. 1956. Fish records from the Pleistocene of southern Cali-
fornia in the collections of the Los Angeles County Museum. So. Calif. Acad.
Sci., Bull. 55:47-49.
Kanakoff, George P., and W. K. Emerson. 1959. Late Pleistocene invertebrates
of the Newport Bay area, California. Los Angeles Co. Mus., Contrib. in Sci.
31:1-47.
McAllister, D. E. 1963. A revision of the smelt family, Osmeridae. Natl. Mus.
Canada, Bull. 191:1-53.
Miller, Daniel J., and Daniel Gotshall. 1965. Ocean sportfish catch and effort
from Oregon to Point Arguello, California, July 1, 1957-June 30, 1961. Calif.
Dept. Fish and Game, Fish Bull. 130: 1-130.
Radovich, John. 1961. Relationships of some marine organisms of the northeast
Pacific to water temperatures, particularly during 1957 through 1959. Calif.
Dept. Fish and Game, Fish Bull. 112: 1-62.
Uyeno, Teruya, and Robert R. Miller. 1963. Summary of late Cenozoic fresh-
water fish records for North America. Univ. Mich. Mus. Zook, Occ. Pap.
631:1-34.
Valentine, James W. 1961. Paleoecologic molluscan geography of the Californian
Pleistocene. Univ. Calif. Publ. Geol. Sci. 34(7) : 309-442.
Accepted for publication June 11, 1970
■
ii
ll
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' ANGELES
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1 MUSEUM
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CONTRIBUTIONS
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A NEW SPECIES OF BATRACHOSEPS (SLENDER SALAMANDER)
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Editor
A NEW SPECIES OF BATRACHOSEPS
(SLENDER SALAMANDER)
FROM THE DESERT OF SOUTHERN CALIFORNIA
By Arden H. Brame, Jr.^
Abstract: A new species of Batmchoseps is described from
the lower desert slopes of the Santa Rosa Mountains. This species
occurs in a small canyon tributary of Deep Canyon beneath
limestone sheeting where there is perennial water seepage and
below talus on the canyon floor during the wetter months. The
closest relative of this desert slender salamander is Batrachoseps
stebbinsi with which it agrees by having large feet, low numbers
of trunk vertebrae, and a reduced phalangeal formula of 0-2-3 -2.
The new species, a broadheaded salamander, is primitive, and
probably is a fragment of a once widely distributed form. Batra-
choseps stebbinsi, B. pacificus and B. simatus possibly represent
other fragments of the same ancestral population.
In the most recent review of the genus, Brame and Murray (1968),
described three new species of Batrachoseps and elevated B. major to a full
species. In addition, they compared the morphology and ecology of each of
the then known species. They indicated that Batrachoseps wrighti is the most
primitive species and is restricted to derivatives of Arcto-Tertiary plant com-
munities of north-central Oregon. They considered Batrachoseps relictus to
be close to the ancestral form that gave rise to all of the species in California
and northern Baja California, and showed it to occur in four disjunct regions:
(1) the Santa Lucia Range of Monterey and San Luis Obispo counties, (2)
1,000 to 8,000 feet elevation on the western slopes of the Sierra Nevada, (3)
Santa Cruz Island off Santa Barbara County, and (4) 6,000 to 8,000 feet ele-
vation in the San Pedro Martir Range of Northern Baja California, Mexico.
In each of these areas, Batrachoseps relictus occurs within derivatives of
Arcto-Tertiary or Arcto -Tertiary -Madro -Tertiary ecotonal communities.
Brame and Murray suggested that Batrachoseps stebbinsi of the Piute and
Tehachapi mountains in Kern County, Batrachoseps simatus of the Kern
River Canyon, Kern County, and B. pacificus of the Northern Channel Islands
off Santa Barbara and Ventura counties, probably arose from a relictus-Mke
ancestor at an early time and possibly represent fragments of a once continu-
ous species inhabiting primarily Arcto-Tertiary communities. Batrachoseps
attenuatus, ranging from southwestern Oregon to southwestern California and
the lower elevations of the western slopes of the Sierra Nevada, and B. major
of southwestern California and northwestern Baja California are considered
more recently derived forms from Batrachoseps relictus. Both have been able
^Research Associate, Section of Herpetology, Los Angeles County Museum of
Natural History. (Present address: Eaton Canyon Nature Center, 1750 North
Altadena Drive, Pasadena, California 91107.)
1
2
Contributions in Science
No. 200
to adapt to the drying trends of the Pleistocene and have successfully invaded
the Madro-Tertiary floral communities. Brame and Murray discussed some
dubious records for Batrachoseps from southwestern Alaska, the Nevada de
Colima in Mexico, and the Cape region of southern Baja California, Mexico,
the latter area being isolated by desert to the north and ocean or gulf on the
other three sides. It is hoped that renewed interest in verifying these question-
able records will now be stimulated based upon the unusual desert habitat for
the new Batrachoseps species herein described.
Warden Russel W. Murphey of the California Department of Fish and
Game, hiked down the steep east-facing desert slopes to a tributary of and
above Deep Canyon, Riverside County, during the late summer of 1969, and
found a moist water seepage area below the canyon cliffs. In order to create a
small water hole for the native wildlife, he began pulling off the hard rock-like
encrusted limestone overlying a moist sandy soil and granitic rock beneath
and discovered, to his amazement, clusters of salamanders, the first desert
record and the most eastern known salamander in California. Unfortunately
these specimens died in captivity, but Warden Murphey, in the company of
the author and James E. De Weese, returned to the locality early in 1970
and collected 8 additional specimens. The series of Batrachoseps proved to
represent a new species, and due to the unusual arid environment for a sala-
mander, I propose that it be known as:
Batrachoseps aridus, new species
Figures 1-5; Table 1
Suggested common name. Desert Slender Salamander.
Holotype. LACM 56271, an adult female from Hidden Palm Canyon, a
tributary of Deep Canyon, elevation approximately 2500 feet, (10.5 miles by
road S of the intersection of state Highways 111 and 74, town of Palm Desert),
NW end of Santa Rosa Mountains, from slopes on western side of Coachella
Valley, Riverside County, California; collected by Russel W. Murphey, James
E. De Weese, and the author, February 27, 1970, (see map. Fig. 5, for locality
and Figs. 1 and 2 of holotype).
Paratypes. LACM 56272-78 (7 specimens), collected with the holotype;
and LACM 56570-80 (1 1 specimens), collected March 22, 1970, by Allan L.
Reinholtz, Patricia L. Brame and the author.
Diagnosis. A moderately small species, 14 adults ranging from 30.3 to
48.4 mm (mean 37.4) in standard lengths; with short tails; and distinguished
from all other species of Batrachoseps by its distinctive ventral coloration con-
sisting of blackish maroon on belly and gular area, sharply contrasted with the
flesh colored tail venter (see color pattern, Fig. 1); distinguished from all
other species of Batrachoseps except B. stebbinsi by having an 0-2-3-2 hand
and foot phalangeal formula {B. attenuatus rarely has an 0-2-3-2 formula);
and distinguished from all other species of Batrachoseps, except B. pacificus,
by having a proportionally broader head, standard lengths 5. 7-7. 4 (mean 6.5)
1970
A New Species of Slender Salamander
3
Figure 1. Holotype of Batrachoseps aridus, new species, LACM 56271, an adult
female. Line equals 1 centimeter.
times head widths versus 7.0-8. 1 (mean 7.4) in pacificus and greater than 7.5
for the other Batrachoseps species. See section on comparison for additional
character differences between Batrachoseps aridus and the other seven species
in the genus.
Description of holotype. An adult female, snout moderately short, some-
what pointed at tip; nostril moderately small, nasolabial groove well defined.
Standard length 7.4 times head width; standard length 4.5 times snout-gular
fold length. Eye large and protuberant. Well defined postorbital groove
extends posteriorly from eye as shallow depression to three grooves that pro-
ceed sharply ventrally, for 2.2 mm to 1st groove which becomes the 1st
nuchal groove ventrally, for 3.8 mm to the 2nd groove, which becomes the
2nd nuchal groove ventrally- and 5.7 mm to the 3rd groove which becomes
the gular fold groove ventrally. The 1st nuchal groove is 2.8 mm and the 2nd
nuchal groove is 1.8 mm anterior and parallel to the gular fold groove.
Vomerine teeth 1 1 left, 12 right, extending diagonally in a posterior direction
from the medial border of the internal nares, arranged in patches. Maxillary
teeth 22 left, 23 right, extend about two-thirds through eye. Premaxillary
teeth 10, not piercing lip. Intercostal grooves 18 per side. Tail short, 0.99
times standard length; dorsoventrally compressed at base but more rounded
from middle to tip; not noticeably constricted at base. Postiliac gland not
evident. Limbs very long, appressed limbs leave 6.5 intercostal folds uncov-
ered; hind limb overlaps 5.5 intercostal folds; standard length 5.3 times right
fore limb, 4.8 times right hind limb; coupling value 1.53 (axilla-groin length
divided by combined hind and fore limb lengths). Webbing of hands and feet
moderate, leaving 1 to 1.5 phalanges free of web. Fingers and toes with large
subdigital pads except first digit, which has somewhat reduced pad; fingers
in order of decreasing length: 3, 2, 4, 1; toes in order of decreasing length :
3, 2, 4, 1.
Osteological characters from x-ray of holotype. 20 trunk vertebrae;
2 caudal-sacral vertebrae; 19 ribs; phalangeal formula (both feet) 0-2-3-2,
4
Contributions in Science
No. 200
(both hands) 0-2-3-2, terminal phalanges slightly expanded at tips; single
premaxillary bone.
Measurements (in mm). Head width, 6.5; snout to gular fold (head
length), 10.8; head depth at posterior angle of jaw, 2.7; eyelid length, 2.8;
eyelid width, 2.0; anterior rim of orbit to snout, 2.2; horizontal orbital diam-
eter, 2.0; interorbital distance, 1.2; distance between vomerine teeth and
parasphenoid tooth patch, 1.0; snout to fore limb, 13.0; distance separating
internal nares, 1.8; distance separating external nares, 1.8; snout projection
beyond mandible, 0.8; snout to posterior angle of vent (standard length),
48.4; snout to anterior angle of vent, 45.8; axilla to groin length, 29.4; tail
length, 47.8; tail width at base, 2.8; tail depth at base, 2.4; fore limb length,
9.2; hind limb length, 10.0; width of right hand, 2.6; width of right foot, 3.2.
Color in life. Dorsum blackish maroon, overlaid with an indistinct
lighter dorsal band visible from the tip of the snout, top and sides of head;
trunk dorsum, to four-fifths length of tail, composed of silver to brassy irido-
phores interspersed with some larger patches of metallic golden orange
iridophores. Ventrally, the gular area and trunk are a darker blackish maroon,
darker than dorsally, with numerous silver iridophores on the neck and some
scattering on the gular surface. The tail venter is a pale pinkish brown (flesh
colored) in marked contrast to the rest of the animal. The silver to brassy
iridophores are also abundant on the upper arms and legs dorsally with a
few scattered also on the fore arms. The iris is a deep blackish maroon with
brassy iridophores.
Variation. There are no sexually dimorphic characters, other than the
larger size of females and papillate vents of males. The holotype has the nar-
rowest head (6.5 mm) at 48.4 mm standard length, and another female,
LACM 56574, has the broadest head (6.4 mm), nearly the same head width
at a much smaller size, 36.5 mm standard length. Aside from the somewhat
striking variation in head widths, the series of Batrachoseps aridus is not too
variable in other characters (see Table 1 for comparative measurements). All
are quite similar in color to the holotype, except LACM 56571, a male, which
is more of a uniform chocolate-brown in ground color with no iridophores
present (see Fig. 1 for normal color pattern). The paratypes agree with the
holotype in phalangeal formula, but 10 of the 18 paratypes have only 19
trunk vertebrae and 18 ribs, whereas 7 agree with the holotype in having 20
trunk vertabrae and 19 ribs. In six paratypes, tibial spurs can be seen (prob-
ably present in all Batrachoseps aridus, but difficult to see in x-rays). One
paratype, LACM 56574, has multiple vertebrae fusions with four pairs of
ribs extending from one fusion and three left, and two right from another
fusion, so its vertebral count was not incorporated above.
Comparison. In addition to the characters discussed in the section on
diagnosis, Batrachoseps aridus is distinguished from all other species of
Batrachoseps by having proportionately longer heads, standard lengths
4. 1-4.7 (mean 4.4) times snout-gular fold lengths (= head lengths); by having
1970
A New Species of Slender Salamander
5
proportionately longer limbs, standard lengths 4.0-4.8 (mean 4.4) times right
hind limb lengths; costal folds left uncovered by appressed limbs (= limb
interval) 3. 5-6. 5 (mean 5.0); coupling values (right hind limb length + right
fore limb length divided into axilla-groin length) 1.08-1.53 (mean 1.31); by
having proportionately larger feet, standard lengths 12.5-15.1 (mean 13.7)
times right foot widths; costal grooves 16-19 (mean 17.8) per side (see Brame
and Murray, 1968; 28-29, Table 5, for comparative data for the other seven
species of Batrachoseps); and by its distinctive coloration of black to choco-
late-maroon ground color dorsally with an indistinct band composed of
hundreds of separated silver to brassy iridophores intermixed with orange-
gold patches of iridophores on front part of head, trunk, first four-fifths of
tail, sides of head, neck and shoulders, and silver iridophores scattered on
gular surface and abundant on neck venter region; venter of trunk darker
than dorsum and more of a darker maroon-black, tail flesh colored ventrally
in marked contrast to trunk venter. Distinguished from its apparent closest
relative, Batrachoseps stebbinsi, by its smaller size, 30.3-48.4 (mean 37.4 mm)
standard lengths versus 50.8-60.4 (mean 55.2 mm); by its proportionately
broader head, standard lengths 5. 7-7. 4 (mean 6.5) times head widths in con-
trast to 7. 5-8.4 (mean 8.0) for 10 B. stebbinsi; by its proportionately longer
head, standard lengths 4. 1-4.7 (mean 4.4) times snout-gular fold lengths
versus 4.5-5.4 (mean 5.1) for 10 B. stebbinsi', by its proportionately longer
limbs, standard lengths 4.0-4. 8 (mean 4.4) times right hind limb lengths
compared to 4.5-5. 1 (mean 4.8) for B. stebbinsi’, coupling values for 18
B. aridus = 1.08-1.53 (mean 1.31) versus 1.38-1.62 (mean 1.51) for 10
B. stebbinsi; costal folds uncovered by appressed limbs 3. 5-6. 5 (mean 5.0) in
contrast to 6-7 (mean 6.25) for 10 5. stebbinsi', by having proportionately
broader feet, standard lengths 12.5-15.1 (mean 13.7) times right foot widths
compared to 13.9-16.4 (mean 14.7) for 10 B. stebbinsi; by having usually one-
half to one more costal grooves per side of trunk, costal grooves 16-19 (mean
17.8) per side versus 18-19 (mean 18.5) for 10 B. stebbinsi (trunk vertebrae
19-20 (mean 19.4) for B. aridus and 20 for B. stebbinsi)’, and by marked dif-
ferences in coloration (see color pattern. Figs. 1 and 2, and Brame and Murray,
1968: 20); basic color differences are presence of distinct dorsal band (often
beige), presence of distinct solid or broken dorsolateral black stripes, ground
color of venter and tail black, white iridophores abundant ventrolaterally
though mostly absent midventrally except on gular area, shoulders, and neck;
iris light brown with brassy iridophores for B. stebbinsi in contrast to the
coloration of B. aridus which has an indefinite dorsal band composed of
hundreds of scattered silver to brassy iridophores and some gold-orange
patches of iridophores on a dark black-maroon to chocolate-maroon ground
color; dorsolateral stripes absent, ground color of venter blackish-maroon
except for tail which is flesh color; silver iridophores absent both ventro-
laterally and midventrally except on shoulders, neck and gular area, and iris
very dark black-maroon with brassy iridophores. Batrachoseps aridus and
6
Contributions in Science
No. 200
Figure 2. Upper: dorsal views of 3 Batrachoseps aridus, new species, at left, from
left to right-LACM 56573, LACM 56272, and the holotype, LACM 56571, all
from the type locality; and 3 B. major at right, from left to right-LACM 56283,
LACM 56281, and LACM 56280, all from W. Exposition Blvd. between 5th and
10th Avenues, Los Angeles. Lower: ventral views of the same individuals in the
same order.
1970
A New Species of Slender Salamander
7
B. stebbinsi are the only Batrachoseps to have a phalangeal formula of
0-2-3 -2 for the feet although it occurs rarely in B. attenuatus. The normal
phalangeal formula for the other six species (including attenuatus) is 1 -2-3-2.
Habitat. The type locality is located on the lower eastern slope of Black
Hill, about 500 feet elevation below State Highway 74 (= Pines to Palms
Highway). The exposed slopes, both east and west above Hidden Palm Can-
yon, have a flora composed largely of cholla, agave, barrel cactus, prickly
pear cactus, creosote, manzanita, juniper and mesquite. The canyon floor
flora is dominated by sugar bush, willow, Washington palms, creosote, mes-
quite, and various grasses, and mosses, algae and maiden-hair fern on the per-
petually shaded north and west facing cliff-like walls of the canyon. A continu-
Figure 3. View down the east facing slope with cholla and agave in the lower
right corner of the picture; the canyon floor at the left center of the picture with
the prominent Washington palms; the well shaded west facing slope in the back-
ground, composed of limestone deposits overlying sand and granitic rock beneath;
the type locality of Batrachoseps aridus, new species, at the base of this slope and
several yards up the canyon (hidden from view) to the right (south). This locality
is Hidden Palm Canyon, about 2500 feet elevation. (See text for additional type
locality description and data, and map. Fig. 5.)
8
Contributions in Science
No. 200
ous water seepage occurs at various areas along these cliffs and is apparently
much more widespread in wetter years judging from the extensive sheet-like
calcium carbonate (limestone) deposits. Many of the Batrachoseps aridus
were taken beneath sheets of limestone during the late summer of 1969^, but
only a few were found there during February and March, 1970. Most of
these were taken beneath rocks at the lower level of the cliffs or at the base
of the north and west facing cliffs or in rock talus. The soil underneath the
limestone sheets and talus rocks appears to be composed largely of coarse
sand intermixed with small granitic rocks. Sugar bush is abundant at the base
of the north and west facing cliffs (see habitat views, Figs. 3 and 4).
Range. Known only from the type locality (see map Fig. 5).
Evolutionary relationships. Morphologically, Batrachoseps aridus is closer
to B. stebbinsi than to other species, and it possesses the proportionately
broadest head, longest limbs and biggest feet of all species. It also has fewer
Figure 4. View of the type locality of Batrachoseps aridus, new species, the
majority of the type series being taken from the talus slope at the lower right of
the picture. A few were collected beneath the limestone sheeting at the left center.
Water seepage occurs beneath the limestone and talus.
^On July 25, 1970, 10 additional specimens (LACM 61272-61281) were obtained
from beneath and within moderately soft porous limestone at the type locality,
thus confirming that this species is active in the summer months.
1970
A New Species of Slender Salamander
9
Figure 5. Map of southern California showing the type locality for Batrachoseps
aridus, new species (solid star in open circle). The two localities for Batrachoseps
stebbinsi are indicated by the open stars inside solid circles. The ranges of B. major
in the southwest and B. attenuatus in the northwest are shown. Cross-hatching
indicates the region of geographic overlap. Actual localities of sympatry between
B. major and B. attenuatus are indicated by solid circles.
Table 1 Measurements and data for Batrachoseps aridus
10
Contributions in Science
No. 200
m§u3q pBOH
00
04
o
04
ON
q
00
q
q
q
04
o
00
4>
00
d
ON
00*
00*
ON*
00
d
d
d
00
00*
00
00
d
d
NO
NO
NO*
qiPlM lood
(N
q
q
q
oo
q
q
*^.
q
oo
00
q
q
q
q
q
04
cn
d
d
04*
04*
04
04*
04*
04
04*
04
04*
04
04*
04*
04*
04
04*
ipSuaq iiBX
00
q
04
,
O'
00
q
00
04
q
CO
q
00
q
CO
00
d
00
d
d
d
d
d
^*
04*
NO*
CO
d
04
CO
d
.-h'
m
m
fO
ro
fO
CO
04
04
CO
CO
CO
04
04
04
04
04
o
ON
O
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O
00
ON
ON
04
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00
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00
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00
4''
9UU9UIOA
q
-
(N
6
CTN
-
00
00
o\
On
00
6
6
d
d
NO
d
NO
m
On
O'
O'
On
r-
O'
O'
NO
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00
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cn
(N
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04
f-H
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f-H
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*
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>*
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.B,
P
p
V
0)
»o
04
04
«n
VN
04
VN
NO
NO
NO
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NO
NO
NO
NO
NO
NO
NO
NO
NO
NO
NO
NO
«o
VN
»n
«o
u~)
VN
»n
»o
*Ol
lO)
WO
>OJ
m
s
s
S
s s s s
U
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u
u u u u
<
<
<
<
<
<
<
<
<
<
<
<
<
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< < < <
H-1
d
d
d
d
d
d
d
d
d
d
d
d
d d d
d
II II
a o
Number of intercostal folds left uncovered when the limbs are appressed to sides, b = Holotype.
Aberrant specimen with multiple vertebral fusions, d = Regenerated or broken tails.
1970
A New Species of Slender Salamander
11
trunk vertebrae, 19-20 (mean 19.4), and costal grooves, 16-19 (mean 17.8)
except for some populations of B. relictus and all B. wrighti which have one to
two less. These characters can all be classified as primitive and I believe
Batrachoseps aridus evolved early, probably from a common ancestor of
B. stebbinsi. Now it is apparently restricted to a spring-fed, protected canyon
in an arid desert and possibly to other suitable relict “oases” and similar
habitats in nearby regions, whereas B. stebbinsi is restricted to the two small
area localities in the Piute and Tehachapi mountains of Kern County about
1 60 airline miles to the northeast in a pine-oak and riparian habitat (see map.
Fig. 5). To a lesser extent, Batrachoseps aridus is related also to B. pacificus
of the Northern Channel Islands and more remotely to B. simatus of the
Kern River Canyon of Kern County. The four species, B. aridus, B. stebbinsi,
B. pacificus and B. simatus, may represent fragmentations of a once wide-
spread species. (See Brame and Murray, 1968: 27-34 for discussion of species
relationships and evolution of the genus.)
Remarks
Additional search should be made of well shaded desert canyons, espe-
cially in places where there are limestone deposits and perennial water seep-
age. Because of the limited extent of the habitat (the limestone sheets could
easily be decimated) of Batrachoseps aridus, I strongly urge that it be placed
on the official red book list of rare and endangered species (U.S.A. and Inter-
national); steps should be taken to protect this unusual habitat and its inter-
esting occupants. A similar action has already been taken with regard to
Batrachoseps stebbinsi.
Acknowledgments
My chief debt of gratitude is due Russel W. Murphey who discovered
Batrachoseps aridus and who kindly led me to the locality. Further, I greatly
appreciate the field aid of James E. De Weese of the University of Southern
California, Allan L. Reinholtz and Patricia L. Brame. The latter prepared
the illustration of the holotype. John W. Wright kindly reviewed the manu-
script and made many useful suggestions. I am grateful to William Presch
for x-raying the entire series.
Literature cited
Brame, Arden H., Jr. and Keith F. Murray. 1968. Three new slender sala-
manders {Batrachoseps) with a discussion of relationships and speciation within
the genus. Los Angeles Co. Mus. Nat. Hist., Bull. 4: 1-35.
Accepted for publication July 1, 1970
:i LOS
I
ANGELES
II COUNTY
; MUSEUM C ■2. J-
CONTRIBUTIONS
IN SCIENCE
j'JUMBER 201
■i!
November 13, 1970
A NEW SALAMANDER (GENUS OEDIPINA)
OF THE UNIFORMIS GROUP FROM WESTERN PANAMA
By Arden H. Brame, Jr. and William E. Duellman
Los Angeles County Museum of Natural History • Exposition Park
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be included with the galley proof.
Virginia D. Miller
Editor
A NEW SALAMANDER (GENUS O EDI PIN A)
OF THE UNIFORM IS GROUP FROM WESTERN PANAMA
By Arden H. Brame, Jr/ and William E. Duellman^
Abstract: A new species of Oedipina from western Panama
is described from the montane cloud forest on the northern
slopes of Cerro Pando. In many characters this long-bodied
Oedipina falls in an intermediate position between the uniformis
and collaris subgroups of the uniformis group and its closest
relative appears to be Oedipina cyclocauda. Because it has simi-
lar coloration of a brown trunk dorsum and white to yellow or
silvery lateral stripes as in O. collaris, O. poelzi, and O. altura
(members of the collaris subgroup), it is placed within that
subgroup.
An expedition conducted by Duellman, Charles W. Myers and Linda
Trueb in May, 1966 to the north slopes of Cerro Pando resulted in the
discovery of an extraordinarily rich salamander fauna including three un-
described species of Bolitoglossa plus B. subpalmata and B. marmorea. In
addition, a large species of Oedipina was discovered; it represents the seventh
species of the genus known to occur in Panama, the other six being Oedipina
alfaroi, O. collaris, O. complex, O. cyclocauda, O. parvipes and O. uniformis
(see Brame, 1968 for a recent review). In allusion to its large size (only O.
collaris attain a longer standard length) we propose that it be called :
Oedipina grandis, new species
Figures 1-3; Tables 1 and 2
Holotype. KU 116676, an adult male from the northern slopes of Cerro
Pando, between 1810 and 1930 m elevation (5937-6330 feet), Provincia de
Bocas del Toro, extreme western Panama near the border with Costa Rica;
obtained by Charles W. Myers, on May 14, 1966.
Paratypes. LACM 57055 and 57056, topoparatypes; KU 116673, 1930
m (6330 feet), KU 116674, 1950 m (6396 feet); KU 116678, 1920 m (6298
feet) and KU 116679, 1810 m (5904 feet); all from the northern slopes of
Cerro Pando, Provincia de Bocas del Toro; collected by Charles W. Myers,
William E. Duellman and Linda Trueb, May 11-30, 1966.
^Section of Herpetology (currently Research Associate), Los Angeles County Mus-
eum of Natural History. Present address : Eaton Canyon Nature Center, 1750
North Altadena Drive, Pasadena, California 91107.
^Curator, Division of Herpetology, Museum of Natural History, and Department
of Systematics and Ecology, University of Kansas, Lawrence, Kansas 66044.
1
2
Contributions in Science
No. 201
Figure 1. Dorsal view of Oedipina grandis, holotype, KU 116676, an adult male.
Total length equals 214.9 mm.
Diagnosis. A member of the Oedipina collaris subgroup of the uniformis
group but in some ways intermediate between the collaris and uniformis sub-
groups (see section on evolutionary relationships). The second largest species
of Oedipina; standard lengths of 7 adults range from 55.1 to 71.4 (mean
65.1 mm). Distinguished from all members of the collaris subgroup (O. col-
laris, O. poelzi, O. altura, O. pseudouniformis and O. cyclocauda) by having
proportionally narrower heads, shorter limbs and smaller feet. Oedipina
grandis is similar to O. poelzi, some O. collaris and O. altura in having a
color consisting of dark to medium brown back and head, trunk bordered
by silvery to cream to yellow light dorsolateral stripes boldly demarcated
from the deep black ventral coloration and black dorsum of tail (Figs. 1 and 2).
(See section on comparison and Table 2 for proportional character differences
between Oedipina grandis and the other members of the collaris subgroup
plus Oedipina stuarti of the uniformis subgroup.)
Description of holotype. Adult male, snout short and gently rounded
at tip; mental hedonic gland not evident externally, nostril small, labial pro-
tuberances small, canthus rostralis moderately arched. Standard length 11.1
times head width and 7.1 times snout-gular fold length (head length).
Vomerine teeth 9 left, 9 right, extending to posterior lateral border of the
internal nares. Maxillary teeth 24 left, 25 right, extending posteriorly to a
point two-thirds distance through length of orbit. Two premaxillary teeth,
both protruding through upper lip. Postorbital groove distinct, extending
for 3.3 mm posteriorly from eye as moderate depression, abruptly proceeding
ventrally and extending across gular area (as the nuchal groove) parallel to
and 2.2 mm anterior to gular fold. Tail thick, nearly round at base but slightly
compressed laterally for last half of length with constriction at base barely
1970
A New Salamander from Western Panama
3
evident; 2.21 times standard length. Postiliac glands large, round and promi-
nent. Limbs moderately short, 12 costal folds remaining uncovered when
limbs appressed to sides of trunk; standard length 9.1 times right fore limb,
8.4 times right hind limb, 34.5 times right foot width. Fingers and toes fairly
thickened, inner and outer toes I and V fused to II and IV respectively; rest
of toes extensively webbed with but the terminal one to one and one-half
phalanges free. Fingers in order of decreasing length: 3, 2, 4, 1; toes in order
of decreasing length: 3, 2, 4, 5, 1.
Measurements (in mm). Head width, 5.9; snout-gular fold (head length),
9.3; head depth at posterior angle of jaw, 3.1 ; eyelid length, 2.4; eyelid width,
1.5; anterior rim of orbit to snout, 2.9; anterior rim of orbit to external nares,
1.8; horizontal orbital diameter, 1.4; interorbital distance, 2.5; distance be-
tween vomerine teeth and parasphenoid tooth patches, 0.6; distance between
vomerine teeth and premaxillary teeth, 2.2; internal choanae (nares) to pre-
maxillary teeth, 2.0; distance separating external nares, 2.1; distance separat-
ing internal nares, 1.8; snout to fore limb, 15.6; snout projection beyond
mandible, 0.8; snout to posterior angle of vent (standard length), 65.6; snout to
anterior angle of vent, 61.8; axilla-groin length, 43.9; fore limb length, 7.2;
hind limb length, 7.8; width of right hand, 1.6; width of right foot, 1.9; tail
length, 145.3; tail depth at base, 3.9; tail width at base, 3.8.
Color in life. Dorsum dark brown with minute silver flecks, especially
on head and limbs, and small irregular dark (black) marks on dorsum; chin
pale brown with silver flecks; belly, lower flanks and ventral surfaces of tail
black with or without silver flecks. Iris (under magnification) dark brown with
small light brown flecks.
Color (in 70% ETOH). Color much faded from that of living specimen;
dorsum of trunk brown bordered by silvery whitish dorsolateral stripes of
uneven borders markedly set off from intense black venter and intense black
tail color. Limbs with some brown spots on black background above; gular
area gray and rest of ventral surfaces black.
Variation. Males have proportionally longer legs (standard length/ hind
leg length equals 8. 1-8.7, mean 8.4, for males; 8.9-10. 1, mean 9.3, for
females) and proportionally larger feet (standard length/ right hind foot
equals 34.5-45.9, mean 39.3, for males; 38.3-47.0, mean 42.0, for females).
Males, as usual for bolitoglossines, are smaller than females; males range
from 55.1-67.6 (mean 62.8 mm), compared to 61.2-71.4 (mean 66.9 mm)
for females. The holotype has a somewhat larger foot than the other speci-
mens and KU 116674, another male, has the broadest head and longest legs
proportionally (see Table 1). Other than the above mentioned differences
and the usual sexually dimorphic characters (males with papillate vents and
premaxillary teeth protruding through upper lip), the paratypes and type agree
closely (see Table 1). The color is much the same for all specimens although
the dorsolateral white-silverish to yellow stripes are broader and more dis-
tinct in most paratypes than for the holotype (see Figs. 1 and 2).
4
Contributions in Science
No. 201
Figure 2. Dorsolateral view of Oedipina grandis, paratype, KU 116674, an adult
male measuring 55.1 mm standard length and 154.1 mm total length. Photograph
by Charles W. Myers.
Comparison. Oedipina grandis is the second largest species of the genus,
7 adults ranging from 55.1 to 71.4 (mean 65.1 mm); related to poelzi, altura,
pseudouniformis and cyclocauda (these occurring in various parts of Hon-
duras, Nicaragua, Costa Rica and Panama [see Brame, 1968]). Distinguished
from these four species by its larger size, 55.1-71.4 (mean 65.1 mm) standard
lengths, compared to 45.1-63.6 (mean 55.6 mm) for 29 poelzi, 57.7 mm for
one altura, 39.3-59.9 (mean 45.3 mm) for 23 pseudouniformis, and 36.4-44.1
(mean 41.5 mm) for 20 cyclocauda. Distinguished from Oedipina poelzi of
Costa Rica by having a much narrower head, standard lengths 10.0-1 1.7
(mean 11.2) times head widths versus 5. 1-6.7 (mean 6.1) for 29 poelzi', much
smaller feet, standard lengths 34.5-47.0 (mean 40.9) times right foot widths
in contrast to 25.0-28.9 (mean 26.5) for 5 poelzi', and fewer vomerine teeth,
mean standard length 3.43 times mean vomerine tooth numbers versus 2.42
times in poelzi. Distinguished from altura of Costa Rica by having a short
rounded snout, shorter legs, standard lengths 8.1-10.1 (mean 8.9) times right
hind limb length compared to 8.2 in the single adult altura, smaller feet,
standard lengths 34.5-47.0 (mean 40.9) times right hind foot widths versus
32.1 in altura, and larger numbers of maxillary teeth 38-60 (mean 49) in
contrast to 27 in altura. Distinguished from Oedipina pseudouni for mis of
Nicaragua and Costa Rica by having a considerably narrower head, standard
lengths 10.0-1 1.7 (mean 1 1.2) times head widths compared to 8. 7-9.7 (mean
9.3) for 23 pseudouniformis, shorter limbs, standard lengths 8.1-10.1 (mean
8.9) times right hind limb lengths versus 6.5-8. 1 (mean 7.4) for 23 pseudo-
uniformis, smaller feet, standard lengths 34.5-47.0 (mean 40.9) times right foot
widths in contrast to 30.2-32.1 (mean 31.4) for pseudouni for mis, and fewer
vomerine teeth, 16-23 (mean 19) compared to 17-34 (mean 25) for pseudo-
uniformis, a smaller species, mean standard length times mean vomerine teeth
1970
A New Salamander from Western Panama
5
for grandis 3.43, for pseudouniformis 1.81. Distinguished from the small sized
Oedipina cyclocauda of Honduras, Nicaragua, Costa Rica and Panama by
having proportionally narrower heads, standard lengths 10.0-1 1.7 (mean 1 1.2)
times head width in contrast to 9.1-11.3 (mean 9.9) for 20 cyclocauda, shorter
legs, standard lengths 8.1-10.1 (mean 8.9) times right hind limb versus
7. 2-9.0 (mean 8.2) in 20 cyclocauda, by having smaller feet, standard lengths
34.5-47.0 (mean 40.9) times right hind foot widths in contrast to 29.0-36.7
for cyclocauda, and similar numbers of vomerine teeth 16-23 (mean 19) in
grandis compared to 16-23 (mean 18) in cyclocauda, a much smaller species,
mean standard length times mean vomerine teeth for grandis 3.43 versus 2.31
for cyclocauda.
The similarities in body proportions between grandis and stuarti (from
Honduras) are probably due to convergence; proportions of limb length and
numbers of maxillary teeth and vomerine teeth are similar (Table 2) but
comparisons of head width, [standard lengths 10.7-1 1.7 (mean 1 1.2) times
head widths versus 1 1.1-12.3 (mean 1 1.7)] reveal that grandis has a pro-
portionately broader head, and comparisons of the feet [standard lengths
34.5-47.0 (mean 40.9) times right foot widths in contrast to 30.6-30.8 for
stuarti] show that grandis has smaller feet. In addition, stuarti is uniform
lead-black on all surfaces in sharp contrast to grandis, which has a brown
back, white or silver to yellow sides and deep black venter.
Habitat. All specimens were obtained in undisturbed montane cloud
forest (see Myers, 1969, for detailed description). The area between 1800
and 1950 meters on the northern slope of Cerro Pando is characterized by
a broad-leafed evergreen forest with a canopy about 20 meters above the
ground. The relatively open forest supports an understory of palms and tree
ferns. Thick growths of mosses occur on trees and logs. The leaf litter is
thick and, at least throughout May, 1966, continuously wet (see Fig. 3).
Five specimens were found beneath decaying logs on the forest floor and
one was beneath rotting thatch from a former shelter. One individual was
found as it was crawling on the ground in the camp clearing at night.
Range. Known only from the type locality and vicinity from between
1810 to 1950 meters (5937-6396 feet) elevation, on the northern slopes of
Cerro Pando, Provincia de Bocas del Toro, extreme western Panama, near
the border with Costa Rica.
Evolutionary relationships. In many characters Oedipina grandis is inter-
mediate between the uni for mis subgroup {uniformis, paucidentata, stuarti,
ignea, alfaroi and taylori) and the collar is subgroup (col laris, poelzi, altura,
pseudouniformis and cyclocauda). These two subgroups compose the uni-
formis group. In head width and hind limb length, Oedipina grandis is inter-
mediate between the bulk of the species in the two subgroups. Except for
having tiny feet as in uniformis, grandis seems to be more closely related to
the lowland Oedipina cyclocauda of the Caribbean slopes of Honduras,
Nicaragua, Costa Rica and northwestern Panama than to any other species.
6
Contributions in Science
No. 201
Figure 3. Montane cloud forest habitat of Oedipina grandis, northern slope of
Cerro Pando, 1950 meters (6396 feet) elevation, Provincia de Bocas del Toro,
Panama. Paratype, KU 116674, of Oedipina grandis was found under the log with
hat on it in lower right. Photograph by Charles W. Myers, May 12, 1966.
Oedipina grandis should be placed on the dendogram (Brame, 1968: 58,
Fig. 29) towards cyclocauda in an intermediate position between the two
subgroups. Its dorsal and dorsolateral coloration is remarkably similar to
poelzi, altura and some collaris, which have brown backs bordered by white-
silverish to yellow dorsolateral stripes set off from the deep black ventral
coloration, an additional reason for aligning grandis with the collaris sub-
group. Thus, grandis is the most specialized member of this subgroup; it has
a proportionately larger head and limbs but smaller feet, thereby approaching
members of the uni for mis subgroup. Therefore, it seems that grandis might
be better adapted to a fossorial existence than other members of the collaris
subgroup. The slight tendency in reduction in number of maxillary teeth and
moderate tendency in reduction in number of vomerine teeth are indications
of trends in specializations similar to those in altura, stuarti, paucidentata,
ignea, taylori and alfaroi; possibly reduction in number of teeth is asso-
ciated with a different diet from that of the multidentate species, a factor
probably further influenced by their more fossorial habits. This description
increases the total to 12 species inhabiting the region of suspected origin for
the genus Oedipina in Costa Rica and western Panama (Brame, 1968: 56),
and increases the total number of known species of Oedipina to 16.
Table 1. Meristic data for specimens of Oedipina grandis
1970
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CONTRIBUTIONS
IN SCIENCE
Dumber 205
November 24, 1970
73
(i ^
FURTHER INFORMATION ON THE BREEDING BIOLOGY
OF THE HONEY GUIDES
By Herbert Friedmann
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 sep-
arately, and numbers run consecutively regardless of subject matter. Number 1 was
issued January 23, 1957. The series is available to scientific institutions and scien-
tists on an exchange basis. Copies may also be purchased at a nominal price. Inquiries
should be directed to Virginia D. Miller, Los Angeles County Museum of Natural
History, 900 Exposition Boulevard, Los Angeles, California 90007.
Robert J. Lavenberg
Managing Editor
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Manuscripts for the LOS ANGELES COUNTY MUSEUM, CONTRIBU-
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papers will be determined by the amount and character of new information. Al-
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largely with specimens in the Museum’s collections, other technical papers will be
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torial Committee including review by competent specialists outside the Museum.
Authors proposing new taxa in a CONTRIBUTIONS IN SCIENCE must
indicate that the primary type has become the property of a scientific institution of
their choice and cited by name.
MANUSCRIPT FORM.-(l) The 1964 AIBS Style Manual for Biological
Journals is to be followed in preparation of copy. (2) Double space entire manu-
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are not required but are strongly recommended. Summaries will be published at
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(9) Submit two copies of manuscript.
ILLUSTRATIONS.— All illustrations, including maps and photographs, will
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Editor at the time corrected galley proof is returned. Appropriate order forms will
be included with the galley proof,
Virginia D. Miller
Editor
FURTHER INFORMATION ON THE BREEDING BIOLOGY
OF THE HONEY GUIDES
By Herb^ert Friedmann’
Abstract: In this paper recent additions to our knowledge
of the reproductive biology of four of the African species of
honey guides of the genus Indicator are presented with comments
for their proper evaluation and correlation with earlier infor-
mation. The egg of Indicator maculatus maculatus is described
for the first time; five new host species are given' for I. indicator
(Merops snperciliosus chrysocercus, M. orientalis viridissimus,
M. imbicus, M. pysillus pusillus, and M. bulocki hulocki), and
one each for 1. minor (Lybiiis leucocephaliis senex) and I. coni-
rostris {Gymnobiicco bonapartei cinereiceps).
Our knowledge of the breeding biology of the honey guides is still far
from complete, but the various bits of new information recently made avail-
able help to fill some of the existing gaps. These 'are given below for each
of the four African species of Indicator to which they apply. I hope that
competent observers and students of birds , in various .parts of Africa may
use their opportunities to further ■ opr knowledge of these difficult butdasci-
nating birds.
1. Spotted Honey guide. Indicator maculatus G. R. Gray
The egg of the nominate race of the spotted honey guide has remained
undescribed. The first known example, now in the collection .of the Los
Angeles County Museum of Natural History, was collected at Boron, in the
Itory Coast, toward the end of June, 1968, by Daniel Parelius, who sent it
to me. The egg is pure wffiite with slight gloss, like those of all species of
Indicator whose egg shells are known, and measures 23.4 x 18 mm. It is very
similar to, biit slightly larger than, the single recorded egg of the central
African race 'of this species, I. maculatus stictithorax, which measures 21.6
X 17.7 mm.
These two eggs, one of each of the two subspecies of this honey guide,
were collected under precisely similar conditions. Parelius had set a trap of
fine net very close to the ground, and he caught in it a female spotted honey
guide, which, on dissection, proved to be in active breeding condition. The
maculatus egg was on the ground immediately beneath the net and must
have been laid by the capttired bird, although it was not noticed until the
next morning when another female spotted honey guide was taken in the
trap. The second bird could not have been responsible for the egg as it was
in non-breeding condition.
* Research Associate in Ornithology, Los Angeles County Museum of Natural His-
tory.
1
2
Contributions in Science
1970
In like manner, the egg of 1. maculatus stictithorax, taken in Rio Muni
(now Spanish Continental Guinea) in January, 1952, by Jorge Sabater, was
also found on the bare earth just beneath a net trap close to the ground, in
which a female spotted honey guide was taken at the time.
Lest it be thought that the typical race of this species is new to the
Ivory Coast, it may be recalled that Bigot and Roux (1966: 148) reported
three examples from there in 1962. These were the first actual specimen
records from that country, and were followed by those obtained by Parelius
six years later. Bigot and Roux noted that the stomach contents of their
specimens contained remains of chiton from beetles and spiders. This caused
them to comment that the spotted honey guide was far from being exclusively
a wax eater, but fed equally on small terrestrial arthropods.
I emphasize once more that beeswax is not the whole, or, so far as we
know, even the most important element of, food of this or of any of the
species of Indicator, but is something added to the regular insectivorous diet
of these birds, although with apparent desire on their part. In the honey
guides of the genera Melignomon and Prodotiscus the wax found in their
stomachs is not wax from beecomb, but is from scale insects, Coccidae.
2. Greater Honey guide. Indicator indicator (Sparrman)
Five additional hosts for the greater honey guide have recently been
reported in Nigeria, all of them species of bee eaters. It was known previously
that members of this family are highly favored as hosts by the greater honey
guide, so it is not surprising to find that additional species of the group are
similarly parasitized. Four of the five are species new to the host list; the
other one is a sub-species previously unrecorded in this capacity, but con-
specific with another race already known to be victimized in other parts of
Africa by the same species of honey guide. The new data, given below,
together with the information previously assembled by me (Friedmann, 1955,
1958, 1968), show that nearly 30 per cent of all recorded instances of para-
sitism by the greater honey guide, actually 47 out of 161 cases, have involved
bee eaters as the hosts.
The additions are as follows :
Blue-cheeked bee eater. Merops superciliosus chrysocercus Cabanis and Heine.
Walsh (1966: 74) first recorded this bee eater breeding in a sand bank
on the Niger River about 20 kilometers south of Kainji Dam, Kontagora
Division, Nigeria. This constituted a considerable southern extension of its
known breeding range, as until then the bird was not known to nest south
of 15° 8' N, approximately 60 kilometers south of Ansongo, on the Niger
River, in the French Sudan. From June 29, 1966, Walsh studied the Kainji
Dam colony of some 14 nest holes to which the birds were seen carrying
food, either to incubating mates, or, more probably, to young in the recesses
of the tunnel nests. On July 1 6, Walsh dug out one of these nests and found
it to contain as its sole inmate a young greater honey guide nearly ready to
leave the nest. From data on other nestlings of this honey guide it is known
1970
Breeding Biology of the Honey Guides
3
that the period from hatching to fledging is approximately a month. This
would place the date of egg laying around the beginning of June.
This is the first record for the blue-cheeked bee eater as a host of the
greater honey guide. It is apparently the same instance that was listed later
by Wells and Walsh (1969: 14). Additional reports may be anticipated as
the blue-cheeked bee eater seems to be spreading southward in Nigeria. Walsh
(in litt., September, 1969) informed me that the 1966 colony of 14 nest holes
had increased to 90 nest holes in 1967, and in 1968 he discovered another
colony of about 150 nest holes about eight kilometers farther south on the
Niger River.
Little green bee eater. Merops orientalis viridissimus Swainson
Fry (1967: 12) added the little green bee eater to the list of victims of
the greater honey guide in Nigeria. No details were given, and it is not
possible to say whether but a single such instance or more than one came
to his notice.
Nubian carmine bee eater. Merops imhicus Gmelin
Wells and Walsh (1969: 14) reported this bee eater to be parasitized
by the greater honey guide near the Kainji Dam, Nigeria. In reply to my
inquiry Walsh (in litt., September, 1969) informed me that the report was
based on seeing recently fledged young of Indicator indicator at the colony
of carmine bee eaters, but as the observations were made by Wells he could
not elaborate on them, and Wells had departed for Malaya and could not
be reached readily. It is unfortunate that the observations as recorded did not
specifically mention feeding of the young parasites by the adults of the host
species, but the fact that there were more than one of the young honey guides
in immediate proximity to the nesting colony adds substance to the record,
the exact wording of which is as follows “. . . At the Shugunu nubicus colony
in 1966 young honey guides were on the wing in V (May), before the majority
of young bee eaters had left the next (nest) ...”
Least bee eater. Merops pusillus pusillus Muller
In eastern and southern Africa the race M. p. meridionalis has long
been known to be one of the most frequent, if not the very most frequent,
of the victims of the greater honey guide. It is therefore not surprising to
find that in Nigeria the nominate race of this bee eater is also similarly im-
posed upon by the parasite. The first such instance was reported by Dodds
(1959: 1 1 8) in a paper overlooked by me until recently, and then additional
and wholly independent observations were published by Fry (1967 : 12).
White-fronted bee eater. Merops bulocki bulocki Vieillot
The nominate race of this bee eater has been found to be a frequent
victim of the greater honey guide in Nigeria by Fry (1965: 78, and 1967 : 12).
If we were to follow White (1965: 234) in considering bullockoides conspecific
with bulocki it would be pertinent to state that the former taxon is known
4
Contributions in Science
1970
to be parasitized in Kenya and Rhodesia, where, indeed, it seems to be a
regularly selected host, for which at feast six records have been reported
(Friedmann, 1968: 3). However, Fry (1969: 574) after extensive held as
well as museum studies, considers bullockoides and bulocki to be valid taxa,
although closely related species, and I concur. , '
Boehm^s bee eater. Merops boehmi Reichenow
In my 1968 paper (Friedmann, 1968: 3) I cited Benson,' Brooke, and
Vernon (1964: 67) as reporting a Zambian instance of this bee eater serving
as a honey guide host. In this capacity it was known earlier only from a
single case found in Malawi. Since then Benson has informed me that there
was no Zambian record and that he and his co-authors had combined Malawi
and Zambia data in their paper.
While additional instances of parasitism by the greater honey guide
have come to my notice since my 1968 paper, they have contributed no new
information, but merely added to earlier, similar occurrences. In two other
species of Indicator there are, however, one new host record for each.
3. Lesser Honey guide. Indicator minor Stephens
An additional host for the lesser honey guide (nominate sub-species)
may be reported solely on a set of eggs of the white-headed barbet, Lybius
leucocephalus senex (Reichenow) , with one of the lesser honey guide. These
are currently on display in the exhibition galleries of the National Museum
in Nairobi. The set apparently was acquired by the Museum from Sir Charles
F. Belcher, but I was unable to learn the locality or date of collection when
visiting the Museum in October, 1969.
4. Thick-billed honey guide. Indicator conirostris (Cassin)
Mr. Gorman M. Bond informed me that in the second week of June,
1965, A. Forbes Watson collected a nestling fernale of this honey guide
from a nest of a grey-throated barbet, Gymnobucco bonapartei cinereiceps
Sharpe, in the Kakamega Forest, western Kenya. This is the first record for
the subspecies cinereiceps as a victim of the thick-billed honey guide, although
the nominate race of the barbet had been known earlier to be parasitized
in Cameroon. The Kakamega bird is now in the National Museum of Natural
History, Washington.
, Literature Cited
Bigot, L., and F. Roux. 1966. Premieres donnees sur I’avifaune de la Savane et
de la Foret-Galerie de Lamto (Cote-d’Ivoire). Oiseaux 36 : 145-152.
Dobbs, K. A. 1959. Some birds of Sokoto, Northern Nigeria, with brief notes on
their status. Nigerian Field 24: 102-1 19.
Friedmann, H. 1955. The honey guides. U. S. Nat. Mus. Bull. 208: 1-292.
1958. Advances in our knowledge of the honey guides. Proc. U. S. Nat.
Mus. 108, no. 3404; 309-320. ' '
1968. Additional data on brood parasitism in the honey guides. Proc.
U. S. Nat. Mus. 124, no. 3648: 1-8.
1970
Breeding Biology of the Honey Guides
5
Fry, C. H. 1965. The birds of Zaria III— Residents, Vagrants, and Check-List.
Nigerian Ornithologists’ Soc. Bull. 7: 68-79.
1967. Studies of bee eaters. Nigerian Field 32: 4-17.
1969. The evolution and systematics of bee eaters (Meropidae). Ibis 111:
557-592.
Walsh, F. 1966. Extension of breeding range into Nigeria of the blue-cheeked
bee eater. Nigerian Ornithologists’ Soc. Bull. 3: 74.
Wells, D. R. and F. Walsh. 1969. Birds of northern and central Borgu (Nigeria).
Nigerian Ornithologists’ Soc. Bull. 6: 1-25.
White, C.M.N. 1965. A revised check list of African non-passerine birds. Govern-
ment Printer, Lusaka. 299 p.
Accepted for publication September 18, 1970
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