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ANNALS

OF

CARNEGIE MUSEUM

VOLUME 48

1979

PUBLISHED BY THE AUTHORITY OE THE
BOARD OF TRUSTEES OF THE CARNEGIE INSTITUTE
PITTSBURGH, PENNSYLVANIA
1979

Publications Committee

HARRY K. CLENCH, Chairman
LEONARD KRISHTALKA
C. J= McCOY, JR.
JAMES B. RICHARDSON III

Editorial Staff

HUGH H. GENOWAYS, Editor
DUANE A. SCHLITTER, Associate Editor
STEPHEN L. WILLIAMS, Associate Editor
NANCY J. PARKINSON, Technical Assistant

CONTENTS

Contents .................................................................. v

New Taxa vii

Author Index viii

ARTICLE

1 . A new species of Amphisbaena (Reptilia, Amphisbaenia) from Argentina.

Arthur C. Hulse and C. J. McCoy 1

2. A study of nongeographic variation in Tatera leucogaster (Mammalia:

Rodentia) from Botswana. Pierre Swanepoel, Duane A. Schlitter, and
Hugh H. Genoways 7

3. Floral vascular anatomy of the Himalayan Theropogon pallidus Maxim

(Lilliaceae-Convallarieae). Frederick H. Utech 25

4. Floral vascular anatomy of Scoliopus bigelovii Torrey (Lilliaceae-Pari-

deae = Trilliaceae) and tribal note. Frederick H. Utech 43

5. A systematic review of the olive-backed pocket mouse, Perognathus fas-

ciatus (Rodentia, Heteromyidae). Daniel F. Williams and Hugh H.
Genoways 73

6. Paleontology and geology of the Badwater Creek area, central Wyoming.

Part 17. The late Eocene snakes. J. Alan Holman 103

7. Substrate selection by three species of desmognathine salamanders from

southwestern Pennsylvania: an experimental approach. Anthony J.
Krzysik and Edward B. Miller Ill

8. The Las Avispas burial platform at Chan Chan, Peru. Thomas Pozorski . . 119

9. Late prehistoric llama remains from the Moche Valley, Peru. Shelia

Pozorski 139

10. Karyotype analysis, palynology, and external seed morphology of To-

fieldia tenuifolia (Michx.) Utech (Lilliaceae-Tofieldieae). Frederick
H. Utech 171

11. Edaphosaurus (Reptilia, Pelycosauria) from the Lower Permian of north-

eastern United States, with description of a new species. David S
Berman 185

12. A new Liolaemus (Sauria, Iguanidae) from the high Andes of Argentina,

with ecological comments. Arthur C. Hulse 203

13. Gnathorhiza bothrotreta (Osteichthyes: Dipnoi) from the Lower Permian

Abo Formation of New Mexico. David S Berman 211

14. The yellow-bellied sea snake, Pelamis platurus (Reptilia: Hydrophiidae), in

the Philippines. C. J. McCoy and Donald E. Hahn 231

15. Taphonomy of microvertebrate fossil assemblages. William W. Korth . . . 235

16. Geomyoid rodents from the Valentine Formation of Knox County, Nebras-

ka. William W. Korth 287

17. Notes on bats (Mammalia: Chiroptera) from Bonaire and Curasao, Dutch

West Indies. Hugh H. Genoways and Stephen L. Williams 311

18. Records of bats (Mammalia: Chiroptera) from Suriname. Hugh H. Geno-

ways and Stephen L. Williams 323

19. Alto Salaverry: a Peruvian coastal Preceramic site. Shelia Pozorski and

Thomas Pozorski 337

20. Paleontology and geology of the Badwater Creek area, central Wyoming.

Part 18. Revision of late Eocene Hyopsodus. Leonard Krishtalka ..... 377

21. Paleontology and geology of the Badwater Creek area, central Wyoming.

Part 19. Perissodactyla. Craig C. Black 391

22. Tht Rhyacophila of Pennsylvania, with larval descriptions of^. banksi and

R. carpenteri (Trichoptera: Rhyacophilidae). John S. Weaver III and
Jan L. Sykora 403

23. Checklist of California mammals. Daniel F. Williams 425

24. Eastern North American Pleistocene Ochotona (Lagomorpha: Mammalia).

John E. Guilday 435

NEW TAXA

DESCRIBED IN VOLUME 48

NEW SPECIES

Amphisbaeea minuta, new species, Reptilia Amphisbaenia 2

t Dawsonophis wyomingensis, new species, Reptilia, Squamata 105

t Edaphosaurus colohistion, new species, Reptilia, Pelycosauria 187

t Hyopsodus sholemi, new species. Mammalia, Condylarthra 386

Liolaemus capillitas, new species, Reptilia, Sauria 204

t Perognathus trojectioansrum, new species. Mammalia, Rodentia 290

NEW GENUS

t Dawsonophis, new genus, Reptilia, Squamata 104

t Fossil taxa

AUTHOR INDEX

Berman, D. S 185,

Black, C. C.

Genoways, H. H 7, 73, 311,

Guilday, J. E.
Hahn, D. E.
Holman, J. A.

Hulse, A. C 1,

Korth, W. W 235,

Krishtalka, L

McCoy, C. J 1 ,

Pozorski, S. 139,

Pozorski, T 1 19,

Schlitter, D. A.
Swanepoel, P.
Sykora, J. L. .

Utech, F. H 25, 43,

Weaver, J. S., Ill

Williams, D. F. 73,

Williams, S. L 311,

211

391

323

435

231

103

203

287

377

231

337

337

7

7

403

171

403

425

323

7.7 3
P(7,S-<7:l

ISSN 0097-4463

ANNALS

of CARNEGIE MUSEUM

CARNEGIE MUSEUM OF NATURAL HISTORY
4400 FORBES AVENUE « PITTSBURGH, PENNSYLVANIA 15213
VOLUME 48 6 MARCH 1979 ARTICLE 1

A NEW SPECIES OF AMPHISBAENA (REPTILIA,
AMPHISBAENIA) FROM ARGENTINA

Arthur C. Hulse^

Research Associate, Section of Amphibians and Reptiles

C. J. McCoy

Curator, Section of Amphibians and Reptiles
Abstract

Amphisbaena minuta, new species, is described from Larrea habitat in the arid Bolson
de Pipinaco, Catamarca Province, Argentina, and other amphisbaenian records from the
Province are reviewed.

Introduction

The province of Catamarca, Argentina, has received relatively little
attention from herpetologists. As part of a study of the convergent
evolution of North and South American desert ecosystems, reptiles
were collected (by ACH) in Catamarca during 1973, 1974, and 1975,
principally in the western section of the province. Subandean western
Catamarca is arid, and includes a northward extension of the Monte
Desert (Blair et aL, 1976). The topography consists of a series of iso-
lated valleys, each surrounded by high mountains. The situation sug-
gests great potential for development of local endemism in fossorial
reptiles of low vagility. Among the reptiles collected in one isolated
valley, the Bolson de Pipinaco, is a distinctive new species of Am-
phisbaena. Terminology used follows the system of Gans and Alex-
ander (1962).

‘ Department of Biology, Indiana University of Pennsylvania, Indiana, Pennsylvania
15701.

Submitted 14 August 1978.

1

2

Annals of Carnegie Museum

VOL. 48

Amphisbaena minuta, new species

Holotype.— CM 65531, immature male, from 27 km S Andalgala
(Route 1), Provincia de Catamarca, Argentina, collected by Arthur C.
Hulse on 12 January 1974.

Paratypes .—CM 65532, immature male, from the same locality as
the holotype, collected 1 February 1974 and CM 65533, immature
male, 17 km S Andalgala (Route 1), collected 20 December 1973.

Diagnosis.— A small, slender Amphisbaena lacking major fusions
of head shields; with greatly enlarged prefrontals (Fig. 1); 265 to 271
body annuli along the mid- ventral line; 17 to 19 dorsal and 20 ventral
segments per midbody annulus; 22 caudal annuli; two rows of post-
genial and one row of postmalar chin shields; four distinctly rounded
precloacal pores. Body annuli in the postnuchal region cross the back
at almost right angles, and caudal autotomy is present at the eighth
caudal annulus. The head is not pointed, and the snout is blunt and
rounded.

Description of holotype. — Dorsal body pigmentation is tannish brown. Anteriorly the
pigmentation disappears at the lateral sulci, however, from a third of the length of the
body behind the head to the precloacal annulus the pigmentation extends to the third
annular segment ventral to the lateral sulci. Occasionally the pigmentation fades on the
second, or extends to the fourth segment. Intersegmental raphes are always lighter than
the centers of segments. Ventral coloration on the body is immaculate cream. Dorsal
head coloration is light tan on the rostral and anterior portions of the nasals, but deepens
to a brownish purple on the other dorsal head scales. As with the intersegmental raphes,
the sutures between the dorsal head scales are light. Laterally the dark pigmentation
fades on the upper third of the second supralabial. The remainder of the second, and all
of the first and third supralabials are cream with scattered brown spots. The ventral
surface of the head is immaculate white. Pigmentation of the tail is similar to that of the
body.

The head is short and blunt, and the head scales lack major fusions. The snout is
blunt, somewhat produced, and slightly flattened dorsoventrally. The rostal is barely
visible from above, and is bordered by a pair of large nasals. The paired prefrontals are
the largest head scales, and are about twice the size of the nasals. They are in broad
contact with the nasals, frontals, second supralabials, and oculars. The posterior lateral
margins form 45 degree angles. The frontals are small, about one-third the size of the
prefrontals, and are located in the concavity produced by the posterior angles of the
prefrontals. They are in broad contact with the parietals, and point contact with the
oculars. Four parietals are present. Three and one-half supralabials are present. The
second supralabial is the largest, and is in point contact with the nasal. The oculars are
equal in size to the frontals, and cover large, distinct, darkly-pigmented eyes. The
temporals are paired, in contact with the posterior half-supralabials, and of approxi-
mately that size. The second temporal is in narrow contact with the ocular on the right,
but separated from the left ocular by the first postocular. A single postocular is present
on the left side, but the postocular is divided into two scales on the right.

The mental is relatively small, slightly wider anteriorly than posteriorly. The post-
mental is pentagonal and slightly larger than the mental. Three infralabials are present;
the second is the largest. The first row of postgenials, consisting of a pair of large,
roughly triangular scales, separates the postmental from the malars. The second post-
genial row is composed of five smaller scales. The malars are triangular and in broad
contact with the second infralabials, and in narrow contact with the third infralabials.

1979

Hulse and McCoy— New a MPHiSBAEN a Species

3

Fig. 1, — Dorsal, ventral, and lateral views of the head of Amphishaena minuta (CM
65531, holotype). Scale bar equals 1 mm.

4

Annals of Carnegie Museum

VOL. 48

The first postmalar row is composed of 10 scales approximately the same size as the
segments of the first body annulus. There are two accessory dorsal half-annuli between
the first and second complete body segments.

Ventral annular segments are nearly as wide as long, but not significantly widened.
In the area of the lateral sulcus the annular segments are approximately as long as they
are wide. Middorsally the segments are three times longer than wide. Dorsal annular
segments on the tail are only two times longer than wide. There is a prominent autotomy
constriction at the eighth caudal annulus, where the tail is broken.

The body is cylindrical, and there is no nuchal constriction.

Scale counts are as follows: body annuli, 265; lateral half-annuli, 2; caudal annuli, 22;
preanal pores, 4; annular segments at midbody, 39 (19 dorsal and 20 ventral); suprala-
bials, W2\ infralabials, 3; cloacal segments, 6-4-6. Measurements are snout-vent length,
165 mm; diameter at midbody, 3.5 mm; tail length, 16 mm.

V ariation .—Tht paratypes (CM 65532 and 65533) generally agree
with the holotype in dimensions, coloration, and details of scutellation.
Scale counts for the paratypes are as follows: body annuli, 271, 271;
tail broken at eighth caudal annulus in both; preanal pores, 4, 4; an-
nular segments at midbody, 39 (19 dorsal, 20 ventral), and 40 (20 dor-
sal, 20 ventral); supralabials, 33^, 314; infralabials, 3,3. Measurements
are snout-vent length, 177, 165; diameter at midbody, 3.5 (both).

Distribution and ecology .-—Tht five known specimens of Amphis-
baena minuta, including two collected in 1974 that escaped before they
could be preserved, were taken in bajada habitat in the Bolson de
Pipinaco. All were found alive at night (2100 to 2400 hours), on a paved
section of Route 1, at 17, 20, 22, and 27 km south of Andalgala. Road-
side habitat in this area is typical northern Argentina flatland desert,
with the dominant plant being creosotebush or jarilla {Larrea cunei-
folia), and subdominants Cassia aphylla and Trichocereus sp. (see
Fig. 7, in Williams and Mares, 1978). The soil is relatively loose and
sandy. There was no obvious reason for increased amphisbaenian sur-
face activity on the nights when specimens were collected.

M. A. Mares (personal communication) also collected a small slen-
der species of Amphisbaena, presumably A. minuta, in the Bolson de
Pipinaco, and reported a large, robust species, possibly A. camura,
from near Belen, in the northwestern part of the Bolson. Mares’ spec-
imen, deposited at the Fundacion Miguel Lillo, could not be found for
verification of the tentative identification.

Discussion. —We are unable to reach convincing conclusions about
the affinities of Amphisbaena minuta— by no means an unusual sit-
uation in this difficult genus (Cans and Mather, 1977:36). Although our
specimens are immature, the body diameter/body length relationship
suggests a very slender, relatively small species at maturity. The high
annuli counts, number of preanal pores, moderate number of annular
segments, color pattern, and head shape in combination distinguish A.
minuta amply from all congeners. We at first thought the relationship
of this species might be with A. angustifrons plumbea {sensu Cans

1979

Hulse and McCoy — New Amphisbaena Species

5

and Diefenbach, 1972), a poorly known Patagonian form. Amphis-
baena minuta differs from plumhea in having caudal autotomy, post-
nuchal annuli crossing the middorsal line at right angles, and a higher
number of body segments (maximum 218 in plumbea). Moreover,
plumbea is a robust, heavy-bodied species with a pointed snout. We
continue to regard the group of Amphisbaena angustifrons as the most
likely relatives of A. minuta, despite the observed differences in char-
acter states. This section of the genus is in need of further study,
because the present arrangement of species and subspecies is based
on extremely sparse material from the vast geographic area of western
Argentina and Patagonia.

In the most recent keys to the species of Amphisbaena (Cans and
Diefenbach, 1970; Cans and Mather, 1977) A. minuta runs to A. oc-
cidentalis townsendi. This latter species, in contrast to A. minuta, has
a distinctive color pattern of contrasting dorsal and ventral pigmen-
tation, that appears mottled, and marked elongation of dorsal segments
of the anterior trunk annuli (Cans, 1961).

The herpetofauna of Catamarca Province was first described by Kos-
lowsky (1895), but he recorded no species of Amphisbaena from the
area. Since that time Cans (1965) has reported Amphisbaena angus-
tifrons {angustifrons) from Andalgala and Esquina Grande, and A.
camura from San Antonio and Catamarca (city). Gans (1966) cited A.
darwini heterozonata from the Provincia de Catamarca, without spe-
cific locality. To these published records we can add A. angustifrons
angustifrons from Camino El Alto (FML 431), and A. darwini heter-
ozonata from Rio Balcosna (FML 510a-~510b). Although all three of
these species are potentially sympatric with A. minuta, most of the
records in the province are from the more mesic eastern section.

Acknowledgments

Huise’s field work was supported by NSF grant GB-27125 to Dr. W. Frank Blair of
the University of Texas, Austin for the Origin and Structure of Ecosystems Subprogram
of the International Biological Program. McCoy’s travel to Argentine museums was
supported by the Netting Research Fund, Carnegie Museum of Natural History. For
facilities and assistance we thank Dr. R. F. Laurent, Fundacion Miguel Lillo, Tucuman
(FML), and Dr. J. M. Gallardo and Sr. J. R. Cranwell, Museo Argentino de Ciencias
Naturales, Buenos Aires. For loan of specimens we thank Dr. K. Kramer, Naturhi-
storisches Museum, Basel; Dr. G. R. Zug, National Museum of Natural History, Wash-
ington, D.C.; Dr. Ernest Williams, Museum of Comparative Zoology, Cambridge. We
are particularly indebted to Or. Carl Gans for helpful advice and comments on the
manuscript. The type-specimens are deposited in Carnegie Museum of Natural History
(CM).

Literature Cited

Blair, W. F., A. C. Hulse, and M. A. Mares. 1976. Origins and affinities of ver-
tebrates of the North American Sonoran Desert and the Monte Desert of north-
western Argentina. J. Biogeography, 3:1-18.

6

Annals of Carnegie Museum

VOL. 48

Cans, C. 1961. Notes on amphisbaenids (AmphisbaeniaiReptilia). 2. Amphisbaena
occidentalis Cope from the coastal plain of Northern Peru. Postiila, 56:1-17.

. 1965. Notes on amphisbaenids (Amphisbaenia, Reptilia). 17. A redescription

and discussion of Amphisbaena angustifrons Cope and Amphisbaena camura Cope
of South America. Amer. Mus. Novitates, 2225:1-32.

— . 1966. Studies on amphisbaenids (Amphisbaenia, Reptilia). 3. The small species

from southern South America commonly identified as Amphisbaena darwini. Bull.
Amer. Mus. Nat. Hist., 134:185-260.

Cans, C., and A. A. Alexander. 1962. Studies on amphisbaenids (Amphisbaenia,
Reptilia). 2. On the amphisbaenids of the Antilles. Bull. Mus. Comp. Zool., 128:65-

158.

Cans, C. and C. O. da C. Diefenbach. 1970. Amphisbaena. Pp. 26-38, in Catalogue
of the Neotropical Squamata, Part II, Lizards and Amphisbaenians (by J. A. Peters
and R. Donoso-Barros), Bull. U.S. Nat. Mus., 297:i~viii, 1-293.

. 1972. Description and geographical variation of the South American Amphis-
baena angustifrons: the southernmost amphisbaenian in the world (Reptilia, Am-
phisbaenia). Amer. Mus. Novitates, 2494:1-20.

Cans, C., and S. Mathers. 1977. Amphisbaena medemi, an interesting new species
from Colombia (Amphisbaenia, Reptilia), with a key to the amphisbaenians of the
Americas. Fieldiana Zool., 72:21-46.

Koslowsky, J. 1895. Batracios y reptiles de La Rioja y Catamarca. Rev. Mus. La
Plata, 6:357-370.

Williams, D. F., and M. A. Mares. 1978. A new genus and species of phyllotine
rodent (Mammalia: Muridae) from northwestern Argentina. Ann. Carnegie Mus.,
47:193-221.

^ 73

ISSN 0097-4463

ANNALS

I)/ CARNEGIE MUSEUM

CARNEGIE MUSEUM OF NATURAL HISTORY
4400 FORBES AVENUE ® PITTSBURGH, PENNSYLVANIA 15213
VOLUME 48 6 MARCH 1979 ARTICLE 2

A STUDY OF NONGEOGRAPHIC VARIATION IN
TATERA LEUCOGASTER (MAMMALIA:

RODENTIA) FROM BOTSWANA

Pierre Swanepoee^

Resident Museum Specialist, Section of Mammals

Duane A. Schlitter

Associate Curator, Section of Mammals

Hugh H. Genoways

Curator, Section of Mammals

Abstract

Specimens of Tatera leucogaster from six localities in Botswana were tested for
variation with age, secondary sexual variation, and individual variation. Of the six age
classes recognized, categores I, II, and III each formed their own group, whereas cat-
egories IV, V, and VI were not separable on a morphometric basis and were considered
to be adults. Significant secondary sexual variation was found only in depth of braincase
in which females were larger. The only character exhibiting unusually high individual
variation was length of posterior palatal foramen. All other characters exhibited indi-
vidual variation within acceptable limits.

Introduction

Gerbils of the genus Tatera are members of the rodent subfamily
Gerbillinae, which now generally is placed in the Cricetidae. The geo-
graphic range of the genus includes subsaharan Africa and eastern
Syria and southwestern Turkey across Southwest Asia to the Indian

’ Permanent address: Kaffrarian Museum, King William’s Town 5600, Republic of South
Africa.

Submitted 22 September 1978.

7

8

Annals of Carnegie Museum

VOL. 48

subcontinent, Wroughton (1906), in his early revision of African Ta-
tera, separated the species only on a geographical basis. Later, Hinton
and Kershaw (1920) were able to separate northeastern African Tatera
into two species-groups based on various trenchant characteristics.
Subsequently, Roberts (1929) separated South African species into the
same basic species-groups.

Ellermae (1941) divided the African Tatera into five provisional
groups and admitted an urgent need for a generic revision. Davis (1949)
attempted to analyze the affinities of the southern African species,
although he considered all African species, and concluded that there
were four species in southern Africa and that these were separable
into the same two species-groups of earlier authors. His arrangement
was further refined in a series of papers (Marcus et al., 1960; Davis,
1962, 1965, 1966, 1974, and 1975). Davis (1966) drew attention to the
fact that diagnostic characters did not always hold true in separating
various species of Tatera. He pointed out that it was much easier to
separate coexisting species but that, because of the poorly understood
geographic variation in any given species, the relationships of speci-
mens from more than one area were difficult to comprehend with cer-
tainty.

The current study was undertaken in order to better understand
nongeographic morphometric variation within one species of southern
African Tatera. Specimens identifiable as T. leucogaster, based on
Davis (1966) and Smithers (1971), were studied. Based on the results
of this preliminary study, the senior author plans a detailed revision
of all southern African species of this genus.

Materials and Methods

In the course of this study, 288 specimens of Tatera leucogaster from six localities
in Botswana were examined (see Fig. 1 and Gazetteer). All specimens consisted of
conventional museum skins and skulls. The specimens from these localities in Botswana,
all from west of the Okavango River and Swamp, were grouped for statistical analysis.

Four external and 16 cranial measurements were recorded from most specimens ex-
amined. External measurements were obtained from the labels of specimens prepared
as standard museum skins, whereas cranial measurements were taken by means of dial
calipers.

Cranial measurements are defined below.

Greatest length of skull.— Greatest distance from the anteriormost projection of the
nasal bones to the posteriormost portion of the occipital bone.

Condylobasal length.— Greatest distance from anterior surface of the premaxilla pro-
truding from between the incisors to the posteriormost portion of the occipital condyles.

Zygomatic breadth.— Greatest width across zygomatic arches, measured at right angle
to longitudinal axis of cranium.

Braincase breadth.— Greatest width across braincase measured at right angle to lon-
gitudinal axis of cranium.

Interorbital breadth.- — ^Least width across interorbital constriction at right angle to
longitudinal axis of cranium.

1979

SWANEPOEL ET KI..—TaTERA VARIATION

9

23° 24° 25° 26°

from which specimens were examined are shown in insert of Botswana.

Length of nasals Greatest distance from anteriormost projection of nasal bones to
posteriormost projection of nasals along their medial suture.

Breadth of rostrum.-— Least width across rostrum immediately anterior to zygomatic

plate.

Oblique length of bulla.- — Greatest oblique length of auditory bulla taken from a point
adjacent to paraoccipital process to anteriormost edge of bulla.

Greatest length of bulla. — Greatest oblique length of bulla taken from posterior edge
of mastoidal bulla to anteriormost edge of auditory bulla.

Length of maxillary toothrow. — Least distance from anterior lip of alveolus of M* to
posterior lip of alveolus of M^.

Breadth across upper molars. — ^Least distance, measured at right angle to longitudinal
axis of cranium, from labial side of crowns of one maxillary toothrow to labial side of
other toothrow.

Length of anterior palatal foramen. — Greatest distance from anterior edge to posterior
edge of anterior palatal foramen.

10

Annals of Carnegie Museum

VOL. 48

II III IV V VI

Fig. 2. — Left maxillary toothrow of Tatera leucogaster illustrating wear patterns for
five age categories. See text for description of age categories.

Length of posterior palatal foramen. — Greatest distance from anterior edge to pos-
terior edge of posterior palatal foramen.

Length of diastema. — Greatest distance from posterior edge of incisors at premaxilla
to anterior edge of alveolus of M*.

Depth of braincase. — Skull was placed on a microscope slide and the least distance
measured from dorsalmost portion of skull to ventralmost part of slide, thereafter, the
thickness of the slide was subtracted from this value.

Length of mandibular toothrow. — Least distance from anterior lip of alveolus of M,
to posterior lip of alveolus of M3.

All specimens were assigned to one of six age categories (Fig. 2), defined below.

Age Class I — ^M^ not fully erupted; juvenile pelage present; bullae not ossified; oc-
cipital-sphenoid suture not fused; cranium domed dorsally.

Age Class II — M^ fully erupted; no noticeable wear on occlusal surfaces of upper
molars; bullae not completely ossified; occipital-sphenoid suture not
completely fused.

Age Class III — Occlusal surfaces of upper molars worn; paracone-protocone connec-
tion in M^ complete; bullae nearly to completely ossified.

Age Class IV — Occlusal surfaces of upper molars worn; anterocone approaching worn
level of paracone-protocone on M^; well worn; adult pelage present;
supraorbital ridge present; bullae completely ossified.

Age Class V — ^Three alternative wear patterns on upper molars exist: a) mure of M^
lacking, cusps forming a single enamel lake, b) paracone-protocone and
metacone-hypocone connected on M^ forming single enamel lake, or
c) a combination of a and b; occipital-sphenoid suture completely os-
sified; supraorbital ridge developed but skull not rugose.

Age Class VI — Mure of M‘ and M^ lacking, cusps of both molars each forming a single
enamel lake, anterocone connected to protocone; skull rugose; supra-
orbital ridge well-developed.

1979

SWANEPOEL ET AL. — TaTERA VARIATION

11

Davis (1966) proposed six age categories for Tatera brantsi based on toothwear. His
aging criteria differ in the following ways from the one we propose: his category I (young
and subadults) encompass both our I and II; his II (young adults) corresponds to our
III; his III (adults) to our IV; his IV (mature adults) to our V; his V and VI (old adults)
to our VI.

Statistical analysis was performed on an IBM 360 computer at Carnegie-Mellon Uni-
versity, Pittsburgh, Pennsylvania. Univariate analyses of variation with age, secondary
sexual variation, and individual variation were performed using the UNIVAR program,
developed and introduced by Power (1970). Standard statistics (mean, range, standard
deviation, standard error, variance, and coefficient of variation) are generated by this
program. In the event two or more groups are compared, a single-classification analysis
of variance (ANOVA) is employed to test for significant differences between or among
means. When means were found to be significantly different, the Sums of Squares
Simultaneous Test Procedure (SS-STP) (Gabriel, 1964) was used to determine maximally
non-significant subsets.

For earlier uses of this procedure in mammalian systematics, see, among others,
Choate, 1970; Genoways and Jones, 1971, 1972; Birney, 1973; Smith, 1972; and Geno-
ways, 1973.

Results and Discussion

Three kinds of nongeographic variation are discussed — -variation
with age, secondary sexual variation, and individual variation.

Variation with Age

It is necessary in the study of geographic variation of a taxon to
identify those age categories that represent “adult” animals— that is,
individuals that have stopped growing, or nearly so. Each of the four
external and 16 cranial characters were tested separately for males and
females with single classification ANOVA to determine if any of the
means of age categories differed significantly at the .05 level. If the
means were found to be significantly different, SS-STP was used to
find maximally nonsignificant subsets (Table 1),

Because of small sample sizes, age category VI in males and age
categories I and VI in females were not included in SS-STP analyses.
Relevant standard statistics of these age categories, however, are in-
cluded in Table 1 for comparative purposes.

The only two characters for which no significant differences among
means of all age categories were found were length of mandibular
toothrow in both males and females, and length of posterior palatal
foramen in females. In males, the latter showed two broadly overlap-
ping subsets. Nonoverlapping subsets of ages II, III, and IV and V
were found in three characters for both sexes (condylobasal length,
breadth across upper molars, and length of diastema) and in nine ad-
ditional characters for females only (total length, length of tail, greatest
length of skull, zygomatic breadth, length of nasals, oblique length of
bulla, greatest length of bulla, length of anterior palatal foramen, depth
of braincase). Age categories I and II formed a subset that differed

12

Annals of Carnegie Museum

VOL. 48

Table 1. — Variation with age in external and cranial measurements of Tatera
leucogaster from six localities in Botswana. Statistics given are sample size, mean,
two standard errors of the mean, range, coefficient of variation, F-value, critical
F-value, and results of SSSTP analysis showing nonsignificant subsets. Age categories,
studied within each sex with analysis of variance, are listed in decreasing order with
the largest mean first. Age categories VI for males and I and VI for females were
not analyzed but their values are listed for comparative purposes. Groups of means

that were found to be not significantly different at the 5% level are marked

ns.

Sex and
age classes

N

Mean

± 2 SE

Range

cv

Fs

F

Results

SS-STP

Total length

Male

V

8

295.5

± 6.72

280-310

3.2

35.42

I

IV

11

283.5

± 7.66

266-306

4.5

2.49

I I

III

28

264.1

± 7.18

235-303

7.2

I

II

37

245.7

± 7.89

197-289

9.8

I

I

8

196.5

± 8.48

189-220

6.1

I

VI

2

294.5

± 1.00

294-295

0.2

Female

V

10

298.7

± 7.68

282-321

4.1

30.12

I

IV

22

291.9

± 4.06

277-316

3.3

2.71

I

III

39

267.3

± 6.67

225-308

7.8

I

II

30

247.9

± 8.71

193-286

9.6

I

VI

4

305.5

± 13.70

29^326

4.5

I

3

208.7

± 19.23

190-222

8.0

Length of tail

Male

V

8

158.0

± 4.17

145-164

3.7

33.03

I

IV

11

150.2

± 4.73

140-162

5.2

2.49

I I

III

28

137.8

± 4.53

113-159

8.7

I

II

37

128.2

± 4.44

101-155

10.5

I

I

8

101.6

± 4.05

94-111

5.6

I

VI

2

151.0

± 2.00

150-152

0.9

Female

V

10

158.3

± 6.01

143-173

6.0

23.41

I

IV

22

154.3

± 3.16

140-169

4.8

2.71

I

III

39

141.6

± 4.02

116-165

8.9

I

II

30

130.1

± 5.30

94-155

11.2

I

VI

4

161.5

± 9.88

151-174

6.1

I

3

111.3

± 15.68

97-124

12.2

Length of hindfoot

Male

IV

13

35.5

± 0.84

33-38

4.2

13.01

I

III

28

35.1

± 0.41

33-37

3.1

2.49

I I

V

10

35.0

± 0.67

34-37

3.0

I I

II

37

34.2

± 0.52

31-37

4.7

I

I

8

31.6

± 0.65

30-33

2.9

I

VI

2

35.5

± 1.00

35-36

2.0

1979

SWANEPOEL ET AL. — TATERA VARIATION

13

Table 1. — {Continued)

Sex and
age classes

N

Mean ± 2 SE

Range

cv

Fs

F

Results

SS-STP

Female

III

39

35.2 ± 0.34

32-37

3.0

3.10

I

IV

26

35.0 ± 0.49

33-38

3.6

2.70

I I

V

11

34.8 ± 0.75

33-36

3.6

I I

II

31

34.2 ± 0.61

30-37

4.9

I

VI

4

35.3 ± 0.96

34-36

2.7

I

3

32.7 ± 1.33

32-34

3.5

Length of ear

Male

V

10

21.6 ± 0.68

20-23

5.0

14.92

I

IV

13

21.2 ± 0.46

20-23

3.9

2.49

I I

III

28

21.1 ± 0.37

20-23

4.6

I I

II

36

20.5 ± 0.35

19-24

5.2

I

I

8

18.5 ± 0.53

17-19

4.1

I

VI

2

22.0 ± 4.00

20-24

12.9

Female

V

11

21.8 ± 0.70

20-23

5.4

7.55

I

IV

25

21.7 ± 0.49

10-24

5.6

2.70

I

III

39

21.1 ± 0.28

19^23

4.2

I I

II

31

20.5 ± 0.40

17-22

5.5

I

VI

4

2"^2.5 ± 1.90

21-25

8.5

I

3

18.7 ± 0.67

18-19

3.1

Greatest length of skull

Male

V

6

38.6 ± 0.90

37.2-39.6

2.9

36.55

I

IV

9

37.7 ± 0.74

35.0-39.3

2.9

2.54

I I

III

24

36.3 ± 0.59

33.2-39.1

3.9

I

II

20

33.5 ± 0.68

30.6-36.2

4.6

I

I

3

30.4 ± 1.01

29.4-31.1

2.9

I

VI

1

39.6

Female

V

5

38.3 ± 0.91

37.0-39.5

2.6

27.96

I

IV

18

38.1 ± 0.56

36.0-39.7

3.1

2.76

I

III

24

36.0 ± 0.64

32.2-39.7

4.3

I

II

19

33.7 ± 0.89

30.0-36.5

5.8

I

VI

3

39.7 ± 2.16

37.6-41.2

4.7

I

3

30.8 ± 0.46

30.4-31.2

1.3

Condylobasal length

Male

V

9

34.5 ± 0.40

33.6-35.1

1.7

48.36

I

IV

12

33.5 ± 0.52

31.9^34.8

2.7

2.52

I

III

25

32.1 ± 0.51

29.3-34.2

4.0

I

II

26

29.9 ± 0.61

26.3-32.3

5.2

I

I

4

26.3 ± 0.74

25.6-27.1

2.8

I

VI

1

35.2

N

8

16

31

23

4

3

8

10

21

12

2

2

11

20

23

15

3

2

9

11

30

27

5

1

10

20

34

28

4

3

16

11

31

35

7

2

Annals of Carnegie Museum

VOL. 48

Table 1. — {Continued)

Mean ± 2 SE

Range

cv

Fs

F

Results

SS-STP

34.4 ± 0.60

33.1-36.0

2.5

34.02

I

33.9 ± 0.54

32.3-35.5

3.2

3.74

I

32.2 ± 0.52

28.6-35.9

4.5

I

29.8 ± 0.78

26.5-32.3

6.3

I

35.2 ± 0.90

34.0-35.9

2.6

26.4 ± 0.24

26.2-26.8

0.8

Zygomatic breadth

20.0 ± 0.56

18.9-20.8

4.0

18.32

I

19.3 ± 0.56

17.5-20.5

4.6

2.58

I I

18.8 ± 0.37

16.9-19.9

4.5

I

I

17.9 ± 0.43

16.9-19.0

4.2

I

15.2 ± 1.20

14.6-15.8

5.6

20.1

20.1

19.9 ± 0.27

19.1-20.6

2.3

22.69

I

19.7 ± 0.27

18.8-20.6

3.1

2.75

I

18.7 ± 0.38

16.7-20.1

4.8

I

17.8 ± 0.46

16.6-19.7

5.0

I

19.8 ± 0.72

19.1-20.3

3.2

16.3 ± 0.40

16.1-16.5

1.8

Breadth of braincase

16.2 ± 0.24

15.7-16.6

2.2

19.52

I

16.1 ± 0.24

15.4-16.8

2.5

2.51

I

16.0 ± 0.14

15.0-17.0

2.4

I

15.4 ± 0.18

14.5-16.0

3.0

I

14.7 ± 0.46

14.2-15.3

3.5

I

16.2

16.3 ± 0.23

15.7-16.9

2.2

16.82

I

16.2 ± 0.18

15.3-17.0

2.5

2.72

I I

15.9 ± 0.10

15.2-16.5

1.9

I

15.4 ± 0.25

16.2-16.6

4.3

I

16.3 ± 0.24

16.1-16.6

1.5

14.6 ± 0.41

14.2-14.9

2.4

Interorbital breadth

6.1 ± 0.13

5.6-6.6

4.3

17.83

I

6.1 ± 0.12

5. 7-6.4

3.2

2.46

I

6.0 ±0.11

5. 3-6.5

5.0

I

5.7 ± 0.09

5. 1-6.2

4.7

I

5.3 ± 0.11

5. 1-5.5

2.8

I

6.5 ± 0.30

6.3-6.6

3.3

1979

SWANEPOEL ET AL. — TaTERA VARIATION

15

Table 1. — {Continued)

Sex and
age classes

N

Mean ± 2 SE

Range

cv

Fs

F

Results

SS-STP

Female

V

12

6.2 ± 0.13

5. 8-6. 7

3.8

11.16

I

IV

28

6.1 ± 0.10

5.4-6.9

4.4

2.70

I

III

38

5.9 ± O.IO

5. 2-6.4

5.3

I

II

34

5.7 ± 0.11

5. 2-6. 5

5.4

I

VI

4

6.3 ± 0.30

5.9-6.6

4.8

I

3

5.3 ± 0.24

5. 1-5.5

3.9

Length of nasals

Male

V

8

16.4 ± 0.44

15.4-17.1

3.8

44.90

I

IV

13

15.6 ± 0.43

14.3-16.7

5.0

2.51

I

I

III

27

14.8 ± 0.39

13.3-16.7

6.9

I

II

25

13.4 ± 0.40

11.1-15.5

7.4

I

I

7

11.3 ± 0.26

10.9^11.8

3.1

I

VI

2

15.8 ± 0.10

15.7-15.8

0.4

Female

V

10

16.0 ± 0.30

15.1-16.7

2.9

34.63

I

IV

26

16.0 ± 0.24

14.8-17.2

3.8

2.73

I

III

30

14.7 ± 0.43

12.0-16.5

8.0

I

II

24

13.4 ± 0.49

11.2-15.9

8.9

I

VI

3

16.2 ± 1.16

15.1-17.0

6.2

I

3

11.6 ± 0.13

11.5-11.7

1.0

Breadth of rostrum

Male

V

10

5.4 ± 0.14

5. 1-5.7

4.0

31.90

I

IV

16

5.3 ± 0.13

4. 8-5. 7

5.0

2.48

I

I

III

31

5.0 ± 0.08

4. 3-5.4

4.6

I

II

37

4.8 ± 0.08

4.4-5. 3

4.9

I

I

8

4.4 ± 0.09

4. 3-4. 7

2.9

I

VI

2

5.6 ± 0.40

5.4-5.8

5.1

Female

V

12

5.5 ± 0.14

5. 1-6.0

4.5

43.53

I

IV

27

5.3 ± 0.05

5. 1-5.6

2.6

2.69

I

III

40

5.1 ± 0.06

4. ^5. 5

3.4

I

II

34

4.9 ± 0.09

4.3-5. 5

5.3

I

VI

3

5.5 ± 0.07

5.4-5.5

1.1

I

3

4.5 ± 0.16

4.4-4. 6

2.2

Obliqui

? length of bulla

Male

V

10

10.9 ± 0.15

10.5-11.2

2;1

38.80

I

IV

12

10.6 ± 0.20

10.2-11.0

3.3

2.50

I

I

III

27

10.3 ± 0.16

9.5-11.2

4.0

I

II

32

9.7 ± 0.15

8.7-10.3

4.3

I

I

6

9.0 ± 0.16

8.7-9.2

2.2

I

VI

2

10.8 ± 0.20

10.7-10.9

1.3

16

Annals of Carnegie Museum

VOL. 48

Table 1. — (Continued)

Sex and
age classes

N

Mean ± 2 SE

Range

cv

Fs

F

Results

SS-STP

Female

IV

23

10.7 ± 0.12

10.0-11.1

2.6

35.08

I

V

12

10.7 ± 0.23

10.1-11.3

3.7

2.71

I

III

33

10.3 ± 0.12

9.4-11.0

3.4

I

II

30

9.8 ± 0.17

8.7-10.5

4.8

I

VI

4

10.8 ± 0.22

10.6-11.1

2.1

I

3

9.1 ± 0.47

9.0-9.3

1.9

Greatest length of bulla

Male

V

8

13.7 ± 0.34

12.7-14.2

3.5

22.37

I

IV

8

13.3 ± 0.33

12.7-14.1

3.5

2.52

I

III

25

13.2 ± 0.16

12.0-14.1

3.0

I

II

26

12.6 ± 0.17

11.5-13.2

3.5

I

I

4

11.6 ± 0.35

11.2-12.0

3.0

I

VI

1

13.5

Female

V

7

13.7 ± 0.25

13.2-14.1

2.4

18.39

I

IV

18

13.5 ± 0.16

12.8-14.0

2.6

2.74

I

III

31

13.0 ± 0.14

11.7-13.7

3.1

I

II

24

12.6 ± 0.25

11.1-13.4

4.9

I

VI

4

13.8 ± 0.26

13.6-14.2

1.9

I

3

12.0 ± 0.18

11.8-12.1

1.3

Length of maxillary toothrow

Male

V

9

6.3 ± 0.09

6.2-6.6

2.2

9.09

I

IV

11

6.1 ± 0.13

5.7-6.4

3.6

2.50

I

I

III

30

6.0 ± 0.08

5.7-6.5

3.6

I

I

II

31

5.9 ± 0.07

5.5-6.3

3.4

I

I

I

7

5.8 ± 0.18

5.5-6.2

4.1

I

VI

2

6.4 ± 0.10

6.3-6.4

1.1

Female

V

12

6.3 ± 0.20

5. 7-6. 8

5.5

12.46

I

IV

26

6.3 ± 0.07

5.S-6.6

2.9

2.71

I

III

36

6.1 ± 0.06

5.7-6.6

2.7

I

II

30

6.0 ± 0.08

5. 5-6.4

3.8

I

VI

2

6.5 ± 0.30

6.3-6.6

3.3

I

3

5.9 ± 0.12

5. 8-6.0

1.7

Breadth across upper molars

Male

V

9

8.1 ± 0.19

7.4-8. 3

3.5

32.18

I

IV

12

7.7 ± 0.24

6.8-8.3

5.3

2.50

I

III

30

7.3 ± 0.11

6.7-7.8

4.0

I

II

33

7.0 ± 0.09

6.4-7.6

3.8

I

I

7

6.8 ± 0.16

6.6-7. 1

3.1

I

VI

2

8.2 ± 0.30

8. 0-8. 3

2.6

1979

SWANEPOEL ET AL. — TaTERA VARIATION

17

Table 1. — (Continued)

Sex and
age classes

N

Mean ± 2 SE

Range

cv

Fs

F

Results

SS-STP

Female

V

12

8.0 ± 0.22

7. 2-8. 6

4.8

56.65

I

IV

26

8.0 ± 0.10

7. 5-8. 6

3.3

2.71

I

III

36

7.4 ± 0.11

6.6-8. 1

4.6

I

II

29

7.1 ± 0.09

6.7-7. 5

3.3

I

VI

2

8.5 ± 0.40

8. 3-8. 7

3.3

I

3

6.7 ± 0.07

6.6-6.7

0.9

Length of anterior palatal foramen

Male

V

10

7.3 ± 0.25

6.8-8. 1

5.5

36.60

I

IV

16

7.0 ± 0.14

6.3-7.4

4.1

2.48

I

I

III

29

6.7 ± 0.19

5.7-7.5

7.7

I

II

36

6.2 ± 0.16

5. 3-7. 3

7.9

I

I

8

5.1 ± 0.20

4.8-5.5

5.6

I

VI

2

7.0 ± 0.60

6. 7-7. 3

6.1

Female

V

12

7.2 ± 0.18

6.9-7.8

4.4

32.23

I

IV

28

7.1 ± 0.11

6. 7-7. 7

4.0

2.69

I

III

40

6.7 ± 0.12

5. 8-7. 5

5.5

I

II

33

6.3 ± 0.17

5. 2-7. 3

7.8

I

VI

3

7.4 ± 0.31

7. 1-7.6

3.6

I

3

5.4 ± 0.29

5. 1-5.6

4.7

Length of posterior palatal foramen

Male

V

9

1.7 ± 0.19

1.3-2. 2

16.9

2.82

I

III

27

1.6 ± 0.12

1.0-2. 5

18.7

2.50

I

I

IV

14

1.6 ± 0.17

1. 2-2.4

19.5

I

1

II

31

1.5 ± 0.10

1. 1-2.1

18.7

I

I

I

6

1.3 ± 0.13

1.0-1.4 .

11.9

I

VI

1

1.4

Female

V

12

1.7 ± 0.14

1. 5-2.2

14.0

2.53

ns

III

37

1.6 ± 0.10

0.9^2. 1

18.1

2.71

IV

23

1.6 ± 0.11

1. 1-2.2

16.5

II

28

1.5 ± 0.11

1.0-2. 5

20.0

VI

3

1.5 ± 0.18

1. 3-1.6

10.4

I

2

1.5 ± 0.30

1.3-1. 6

14.6

Length of diastema

Male

V

10

9.6 ± 0.23

9.1-10.1

3.7

48.80

I

IV

15

9.4 ± 0.18

8.7-10.0

3.8

2.48

I

III

30

8.7 ± 0.22

7. 5-9. 7

7.0

I

II

38

8.0 ± 0.19

6.7-9.0

7.4

I

I

8

6.9 ± 0.21

6. 5-7. 3

4.2

I

VI

2

10.2 ± 0.30

10.0-10.3

2.1

18

Annals of Carnegie Museum

VOL. 48

Table 1. — (Continued)

Sex and
age classes

n

Mean ± 2 SE

Range

cv

Fs

F

Results

SS-STP

Female

V

12

9.7 ± 0.21

9.2-10.4

3.8

45.00

I

IV

28

9.4 ± 0.12

8.8-10.0

3.4

2.70

I

III

38

8.8 ± 0.18

7.6-9.7

6.3

I

II

33

8.1 ± 0.21

7.0-9. 1

7.5

I

VI

4

10.1 ± 0.64

9.5-10.5

4.5

I

3

6.8 ± 0.42

6.7-7.0

2.2

Depth of braincase

Male

V

9

15.3 ± 0.22

14.9^15.8

2.2

18.37

I

IV

10

15.3 ± 0.39

14.4-16.2

4.1

2.52

I

III

25

15.0 ± 0.24

13.7-16.0

3.9

I

II

26

14.3 ± 0.21

13.2-15.3

3.8

I

I

4

13.3 ± 0.37

12.9^-13.7

2.8

VI

1

15.7

Female

IV

17

15.7 ± 0.19

14.7-16.4

2.5

25.76

I

V

9

15.6 ± 0.23

14.8-16.0

2.2

2.73

I

III

32

15.0 ± 0.18

13.5-16.2

3.4

I

II

26

14.3 ± 0.28

12.8-15.3

4.9

I

VI

4

16.0 ± 0.19

15.9-16.3

1.2

I

3

13.2 ± 0.29

12.9^13.4

1.9

Length of mandibular toothrow

Male

V

9

5.9 ± 0.10

5. 6-6.2

2.6

0.40

ns

IV

11

5.8 ± 0.15

5. 2-6.3

5.0

2.76

III

30

5.8 ± 0.05

5.5-6. 1

2.6

II

31

5.8 ± 0.08

5. 3-6.4

4.0

I

7

5.8 ± 0.32

5.4-6. 1

6.2

VI

2

6.1 ± 0.20

6.0-6. 2

2.3

Female

V

12

6.0 ± 0.14

5.6-6.6

4.2

2.04

ns

IV

24

5.9 ± 0.11

5. 5-6.6

4.4

2.71

III

36

5.9 ± 0.07

5.5-6.3

3.5

II

32

5.8 ± 0.08

5.4-6.3

3.7

VI

4

6.0 ± 0.14

5. 9^6.2

2.4

I

3

5.7 ± 0.47

5. 6-5.9

3.0

significantly from a subset formed by ages III to V in breadth of brain-
case for males. Nonoverlapping subsets of I and II, III, and IV and V
were exhibited by breadth across upper molars for males. Nonover-
lapping subsets of I, II, and III to V were found in three characters
for males (interorbital breadth, length of bulla, and depth of braincase).

1979

SWANEPOEL ET AL. — TaTERA VARIATION

19

Nonoverlapping subsets of II and III, and IV and V were exhibited for
females by interorbital breadth and length of maxillary toothrow. In
only breadth of rostrum for females, of all characters tested, was a sig-
nificant difference found between age categories IV and V. The means
of age categories IV, V, and VI were found not to be largest in only
two cases (length of hindfoot and length of posterior palatal foramen,
for both males and females). Age category I was found to be the small-
est in all characters tested. Only in length of posterior palatal foramen
for males was age category II not the second smallest. Here the mea-
surement of the one age category VI specimen available fell between
I and IT

In Tatera leucogaster, age categories IV and V cannot be separated
on a statistically significant basis. Additionally, means of measure-
ments taken of age VI individuals fall within the range of those for IV
and V. Therefore, specimens from these age categories were consid-
ered “adults” and were pooled for subsequent analysis of secondary
sexual variation. The remaining three categories appear each to fall
into its own group. Age category III, however, shows a closer asso-
ciation with IV than with II, although it is clearly distinct from the
latter.

From the results of a mark-release study, where age was estimated
by comparing body weight at first capture with a known growth curve,
Davis (1966) was able to apply chronological age to his toothwear
classes in Tatera brantsi from a southwestern area of Johannesburg
(Republic of South Africa). If these same ages are comparable to our
proposed toothwear categories in Tatera leucogaster, individuals in
our age category I and II would be up to four months of age; III, five
to eight months of age; IV, nine to 18 months of age; V, 18 to 24
months of age; and VI, older than two years. Davis (1966) further
found that individuals assignable to his category III (corresponding to
our IV) were mature adults and in full breeding activity. This agrees
with our findings where individuals in our age category IV can at least
be considered “adult” in overall size and undoubtedly were repro-
ductively active.

Secondary Sexual Variation

Adult (age categories IV, V, and VI) males of Tatera leucogaster
were tested against adult females using a single classification ANOVA
to learn if the sexes were significantly different in the characters stud-
ied. The results of the analysis are shown in Table 2.

Females were significantly larger than males in one (depth of brain-
case) of 20 measurements tested. The means for females were larger
than those of males in 14 of the remaining measurements. The means
of four (zygomatic breadth, interorbital breadth, oblique length of bul-
la, length of posterior palatal foramen) were the same for the two

20

Annals of Carnegie Museum

VOL. 48

Table 2. — Secondary sexual variation in external and cranial measurements of Tatera
leucogaster from six localities in Botswana. Statistics given are sample size, mean,
two standard errors of the mean, range, coefficient of variation, F-value, and critical
F-value. Means for males and females that are significantly different at the 5% level
are marked with an asterisk, and those that are not significantly different are marked ns.

Sex

N

Mean ± 2 SE

Range

cv

Fs

F

Total length

Male

21

289.1 ± 5.32

266^310

4.2

3.63

ns

Female

36

295.3 ± 3.81

277-326

3.9

4.02

Length of tail

Male

21

153.2 ± 3.32

140-164

5.0

1.75

ns

Female

36

156.2 ± 2.81

140-174

5.4

4.02

Length of hindfoot

Male

25

35.3 ± 0.51

33-38

3.6

1.40

ns

Female

41

35.0 ± 0.38

33-38

3.4

3.99

Length of ear

Male

25

21.4 ± 0.43

20-24

5.1

1.39

ns

Female

40

21.8 ± 0.40

19-25

5.8

4.00

Greatest length of skull

Male

16

38.2 ± 0.59

35.9-39.6

3.1

0.17

ns

Female

26

38.3 ± 0.51

36.0-41.2

3.4

4.08

Condylobasal length

Male

22

34.0 ± 0.40

31.9-35.2

2.8

1.06

ns

Female

28

34.3 ± 0.40

32.3-36.0

3.1

4.05

Zygomatic breadth

Male

20

19.7 ± 0.39

17.5-20.8

4.4

0.28

ns

Female

34

19.7 ± 0.19

18.^20.6

2.8

4.03

Breadth of braincase

Male

21

16.1 ± 0.16

15.4-16.8

2.3

1.45

ns

Female

34

16.3 ± 0.12

15.3-17.0

2.2

4.03

Interorbital breadth

Male

29

6.1 ± 0.09

5.6-6.6

4.0

0.01

ns

Female

44

6.1 ± 0.08

5.4-6.9

4.2

3.99

Length of nasals

Male

23

15.9 ± 0.33

14.3-17.1

4.9

0.72

ns

Female

38

16.0 ± 0.19

14.8-17.2

3.6

4.00

1979

SWANEPOEL ET AL.—TaTERA VARIATION

21

Table 2,— {Continued)

Sex

N

Mean ± 2 SE

Range

cv

F,

F

Male

28

Breadth of rostrum

53 ± OJO

4.8

0.79

ns

Female

42

5.4 ± 0.06

5. 1-6.0

3.7

3.99

Male

24

Oblique length of bulla

10.7 ± 0.13 10.2~=-11.2

2.9

0.04

ns

Female

39

10.7 ± 0.10

10.(V11.3

2.9

4.00

Male

17

Greatest length of bulla

13.5 ± 0.23 12.7-14.2

3.6

0.74

ns

Female

29

13.6 ± 0.13

12.&-14.2

2.6

4.06

Male

22

Length of maxillary toothrow

6.2 ± 0.10 5. 7-6. 6

3.7

3.48

ns

Female

40

6.3 ± 0.08

5.7-6.8

3.8

4.00

Male

23

Breadth across upper molars

7.9 ±0.17 6.8-83

5.2

2.81

ns

Female

40

8.0 ± 0.10

7.2-8.7

3.9

4.00

Male

28

Length of anterior palatal foramen

7.1 ±0.13 6.3-8. 1

4.9

2.09

ns

Female

43

7.2 ± 0.09

6.7-7.8

4.1

3.99

Male

25

^Length of posterior palatal foramen
1.6 ±0.14 0.8-2.4

21.1

0.09

ns

Female

38

1.6 ± 0.09

1. 1-2.2

16.0

4.00

Male

27

Length of diastema

9.5 ± 0.15 8.7-10.3

4.2

0.15

ns

Female

44

9.6 ± 0.12

8.8-10.5

4.2

3.99

Male

20

Depth of braincase

15.3 ± 0.22 14.4-16.2

3.2

9.75

Female

30

15.7 ± 0.13

14.7-16.4

2.3

4.05

Male

27

Length of mandibular toothrow

5.9 ±0.10 5.2-6.3

4.2

2.94

ns

Female

40

6.0 ± 0.08

5.5-6.6

4.1

3.99

22

Annals of Carnegie Museum

VOL. 48

sexes. In only one character (length of hindfoot) did the mean for
males exceed that of the females.

Although females average larger than males, the difference is sig-
nificant in only one character. Based on these results, specimens of
Tatera leucogaster from the study area were considered not to exhibit
secondary sexually dimorphism and morphometric data for males and
females can be combined in analyses of geographic variation.

Individual Variation

Young animals (age categories I, II, and III) generally were more
variable than were adults (age categories IV, V, and VI) (Table 1).

In adults, relatively low individual variation was exhibited. With the
exception of length of posterior palatal foramen (CV: males, 21.1; fe-
males, 16.0), all characters studied had coefficients of variation of 5.8
or less (Table 2), well within the limits of those found for other small
rodents (Long, 1968, 1969). Although external measurements generally
have relatively high individual variation, only length of tail and length
of ear had high coefficients of variation in the present study. The higher
coefficient of variation exhibited by external measurements usually are
explained by the fact that different collectors are involved in measuring
the specimens (Sumner, 1927). However, material examined for the
present study was collected, and external measurements were taken,
by a relatively few workers from the African Mammal Project of the
U.S. National Museum of Natural History.

With the exception of length of posterior palatal foramen, all char-
acters (both external and cranial) studied exhibited individual variation
within acceptable limits and can be used in future studies.

Specimens examined (288). — Botswana: 15 mi N Nokaneng, 21 (USNM), Nokaneng,
3 1 (USNM); 7 mi N Shakawe, 2 (USNM); Shakawe, 100 (USNM); Sepopa, 122 (USNM);
Tsau, 12 (USNM).

Gazetteer

Nokaneng, 15 mi N

Nokaneng

Shakawe, 7 mi N

Shakawe

Sepopa

Tsau

19°25'S, 22°16'E
19°40'S, 22°16'E
18°14'S, 21°52'E
18°21'S, 2r52'E
18°45'S, 22°15'E
20°09'S, 22°27'E

Acknowledgments

We are indebted to Henry W. Setzer, Smithsonian Institution, Washington, D.C., for I
allowing us to examine the large samples of Tatera leucogaster from northern Botswana
housed at that institution. Numerous people helped make this study possible. Among I

them, we would like to thank Margaret Popovich for assisting the senior author in j

processing specimens and recording data, Judy Schlitter for keypunching computer
cards, John F. Sutton for assistance with computer analyses of data, Nancy Perkins for

1979

SWANEPOEL ET AL. — TaTERA VARIATION

23

drawing the figures, Teresa Bona for typing various drafts of the manuscript, and
Wanda Maritz for secretarial assistance.

This study was completed while the senior author visited the Section of Mammals,
Carnegie Museum of Natural History, as a Resident Museum Specialist in the Museum’s
International Visitor Program. The Department of Nature and Environmental Conser-
vation of the Cape Provincial Administration, Republic of South Africa, and the admin-
istration of the Kaffrarian Museum are greatefully acknowledged for allowing the senior
author to participate in the program.

Literature Cited

Birney, E. C. 1973. Systematics of three species of woodrats (genus Neotoma) in

central North America. Misc. Publ. Mus. Nat. Hist., Univ. Kansas, 58:1-173.

Choate, J. R. 1970. Systematics and zoogeography of Middle American shrews of the
genus Cryptotis. Univ. Kansas Publ., Mus. Nat. Hist., 19:195-317.

Davis, D. H. S. 1949. The affinities of the South African gerbils of the genus Tatera.
Proc. Zool. Soc. London, 118:1002-1018.

. 1962. Distribution patterns of southern African Muridae, with notes on some

of their fossil antecedents. Ann. Cape Prov. Mus., 2:56-76.

. 1965. Affinities of the South African gerbils of the genus Tatera: corrections

and notes. Proc. Zool. Soc. London, 144:323-326.

. 1966. Contribution to the revision of the genus Tatera in Africa. Ann. Mus.

Roy. Afr. Centr., 8vo., Zool., 144:49-65.

■ — . 1974. The distribution of some southern African mammals (Mammalia: Insec-

tivora, Rodentia). Ann. Transvaal Mus., 29:135-184.

— — — . 1975. Part 6.4. Genera Tatera and Gerbillurus. Pp. 1-7, in The Mammals of
Africa: an identification manual (J. Meester and H. W. Setzer, eds.), Smithsonian
Inst. Press, Washington, D.C.

Ellerman, j. R. 1941. The families and genera of living rodents. With a list of named
forms (1758-1936) by R. W. Hayman and G. W. C. Holt. Volume IT Eamily Mur-
idae. Trustees British Mus. (Nat. Hist.), London, xii + 690 pp.

Gabriel, K. R. 1964. A procedure for testing the homogeneity of all sets of means in
analysis of variance. Biometrics, 20:459-477.

Genoways, H. H. 1973. Systematics and evolutionary relationships of spiny pocket
mice, genus Liomys. Spec. Publ. Mus., Texas Tech Univ., 5:1-368.

Genoways, H. H., and J. K. Jones, Jr. 1971. Systematics of southern banner-tailed
kangaroo rats of the Dipodomys phillipsii group. J. Mamm., 52:265-287.

. 1972. Variation and ecology in a local population of the vesper mouse {Nyc-

tomys sumichrasti). Occas. Papers Mus., Texas Tech Univ., 3:1-22.

Hinton, M. A. C., and P. S. Kershaw. 1920. On a collection of mammals from the
Dinka Country, Bahr-el-Djebel. Ann. Mag. Nat. Hist., ser. 9, 6:94-101.

Long, C. A. 1968. An analysis of patterns of variation in some representative Mam-
malia. Part 1. A review of estimates of variability in selected measurements. Trans.
Kansas Acad. Sci., 71:201-227.

. 1969. An analysis of patterns of variation in some representative Mammalia.

Part IT Studies on the nature and correlation of measures of variation. Pp. 289-
302, in Contributions in mammalogy (J. K. Jones, Jr., ed.), Misc. Publ. Mus. Nat.
Hist., Univ. Kansas, 51:1-428.

Marcus, T., B. DeMeillon, and D. H. S. Davis. 1960. New fleas (Siphonaptera)
from Central African gerbils, with notes on the taxonomic status of their hosts,
Tatera leucogaster and T. taborae. Novos Taxa Ent., 20:3-8.

Power, D. M. 1970. Geographic variation of red-winged blackbirds in central North
America. Univ. Kansas Publ., Mus. Nat. Hist., 19:1-83.

Roberts, A. 1929. New forms of African mammals. Ann. Transvaal Mus., 13:82-121.

24

Annals of Carnegie Museum

VOL. 48

Smith, J. D. 1972. Systematics of the chiropteran family Mormoopidae. Misc. Publ.
Mus. Nat. Hist., Univ. Kansas, 56:1-132.

Smithers, R. H. N. 1971. The mammals of Botswana. Mus. Mem., Nat. Mus. Rho-
desia, 4: 1-340.

Sumner, F. B. 1927. Linear and colorimetric measurements of small mammals. J.
Mamm., 8:177-206.

Wroughton, R, C. 1906. Notes on the genus Tatera, with descriptions of new species.
Ann. Mag. Nat. Hist., ser. 7, 17:474-^99.

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ANNALS

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CARNEGIE MUSEUM OF NATURAL HISTORY

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j 4400 FORBES AVENUE ® PITTSBURGH, PENNSYLVANIA 15213

i

'' VOLUME 48 6 MARCH 1979 ARTICLE 3

FLORAL VASCULAR ANATOMY OF THE HIMALAYAN
THEROPOGON PALLIDUS MAXIM
(LILIACEAE^CONVALLARIEAE)

Frederick H, Utech^

Associate Curator, Section of Plants

Abstract

The floral vascular anatomy and carpel morphology of the berry-fruited, monotypic,
Himalayan endemic Theropogon pallidus are presented and compared to that of its
tribal cohort Convallaria. The down turned flowering pedicels of Theropogon have three
large compound bundles from which the total floral vasculature is derived. Through a
series of radial divisions these three pedicel bundles establish branch bundles, which
fuse between the gaps of the divisions and also fuse between the original three bundles.
The three outer tepals, the three outer stamen bundles, and the three dorsals are fusion
products along gap radii which bisect the large pedicel bundles. Similarily the three
inner tepa! medians, the three inner stamen bundles, and the three septal axiais are
fusion products along radii mid- way betm'een the three pedicel bundles. The fleshy
imbricated tepals have only a single bundle, that is a median-— there are no tepal laterals
in either whorl. There is no terminal interconnection between the dorsal and ventral
supplies. The six simple ascending ventrais have direct horizontal funicular branches.
Due to the large number of fusion bundles and radial divisions which can be traced to
the three compound pedicel bundles, the overall floral vasculature must be viewed as
reduced and advanced. The vascularization of the berry fruit in Theropogon is different
from that in Convallaria.

The trilobed gynoecium in Theropogon is tricarpellate, syncarpous, non-stipitate, and
unilocular terminally. Septal grooves, which persist, are present, but there are no septal
glands. Rhaphides were not observed. The average ovule number is 24. The gynoecium
morphology of Theropogon is different from that in Convallaria.

‘ Research and publication supported by the M. Graham Netting Research Fund through
a grant from the Cordelia Scaife May Charitable Trust.

Submitted 12 September 1978.

25

26

Annals of Carnegie Museum

VOL. 48

Introduction

Theropogon palUdus Maximowicz is an endemic species of the Hi- !'

malayan region (Baker, 1875; Hooker, 1894; Engler, 1888; Krause,
1930). This perennial species has a stocky rhizome with numerous, [
thick fibrous roots (Fig. lA, C). The radial, distichous leaves are am |
nual, glaucous above and below, and have prominent median costa
(Fig. IB). The scape is acutely angled to narrowly winged longitudi-
nally and is shorter than the ascending leaves. Fach terminal raceme i
has between five to eighteen flowers. Fach flower is subtended by both
a linear-subulate bract (B), and a linear but shorter braceteole (Brt)
(Fig. ID). Both the bract and the bracteole are shorter than the asso- |
ciated down-curved pedicel (Fig. ID). The pale, rose-red perianth of
the campanulate-shaped flowers are inverted during anthesis. ;

Though this species has been known since the latter part of the last i
century (Maximowicz, 1871) and figured in Curtis' Botanical Maga- |
line (Hooker, 1875; t. 6154), floral anatomical information is scarce. |
Nair and Sharma (1965) using light microscope observations briefly :
described the pollen grains of this species, and Stenar (1953) reported !
a "very probably” Polygonum-iypQ of embryogenesis.

Fngler (1888) and Krause (1930) placed Theropogon Maxim, in the |
tribe Convallarieae subtribe Convallarine with Convallaria L. (Eu-
rope, eastern North America, and eastern Asia), Speiranthe { = Speir-
antha) Baker (eastern Asia), and Reineckea Kunth (Japan), whereas |
Hutchinson (1934, 1959) circumscribed the same genera in his tribe |
Convallarieae without the use of a subtribe. The berry fruit of Ther- |
opogon is the chief character used for this tribal association. Because
the vascular floral anatomy of the berry-fruited genus Convallaria has I
been previously presented (Utech and Kawano, 1976), this presenta-
tion of the vascular floral anatomy of Theropogon and subsequent i
comparison to Convallaria are desirable for classical tribal evaluation.

Materials and Methods s

For observation a series of buds through mature flowers of Theropogon paUidus were
selected from excess material on unmounted herbarium sheets (Nepal: Thare-Dunche, j
200 m, 25 June 1960, Kanai and Shakya 672,012, TI). The flowers were rehydrated in |
4% KOH, dehydrated in an ethanol series to 70% ethanol, and passed through the usual
TBA-wax series (Johansen, 1940; Sass, 1958). Fifteen flowers of varying age (buds to
fruits) were sectioned between 12 to 16 p. and stained in safranin and methylene blue.
Several of the rehydrated flowers that were cleared in the basic solution were stained
in 1% fuchsin (Fuchs, 1963) to show the total vasculature.

This report presents the pattern of floral vascularization from the pedicel to the tips
of the stamens and stigma, and does not imply either ontogenetic nor teleological de-
velopment in the manner of presentation. Cross-sectional, projected cross-sectional, and
roll out longitudinal diagrams are presented for this species (Figs. 2-4). This illustrative
style of vascular presentation prallels that of our previous floral vascular studies (Utech,
1978«, 1978^, 1978c, 1978<i, 1978c). Table 1 compares the vasculature Theropogon and
Convallaria.

1979

Utech — Theropogon Floral Vascular Anatomy

27

Observations
Pedicel Vascularization

The inflorescence of Theropogon pallidas is a simple, erect raceme
that usually attains a height of 2.0 to 4.0 dm (average 3.1 dm). The
flowering raceme portion usually spans the upper one-third of the in-
florescence. Each of the five to eighteen flowers is subtended by a
linear to subulate bract, which averages 1.7 cm in length (range 1.0™
2.4 cm), and by a smaller linear bracteole, which average 0.6 cm (range
0.4-0.9 cm) in length. Both the bract and the bracteole are persistent
(Fig. ID). Each bract has a single, unbranched midvein and two lateral
veins, whereas the bracteole has a single midvein.

There is a single flower per node (Fig. ID). The departing pedicels
clearly exhibit a spiral (1/3) phyllotaxy around the raceme axis. At
each point of pedicel attachment, the pedicels are triangular in cross-
section, and recurved downwards. In cross-section the flowering ped-
icel lacks a sclerenchymatous sheath around the pedicel vasculature,
however, a sclerenchymatous sheath is associated with the pedicel
vasculature in the fruiting pedicel. The pedicel is gibbose below the
receptacle.

For most of the flowering pedicel’s length, there are three large
vascular bundles. Near the base of the flower, that is, the inverted
receptacle, the pedicel is circular in cross-section (Figs. 2A, 3A),
which represents a shape change from the lower triangular cross-sec-
tion. Near the receptacle base each of the three pedicel bundles under-
goes a radial division that results in three pairs of pedicel bundles
(Figs. 2B, 3B).

Tepal and Stamen Vascularization

The six reddish-rose tepals of the inverted campanulate flower are
free to their bases. There is no connate floral tube as in Conv allaria
(Utech and Kawano, 1976). In Theropogon a slight basal size differ-
ence occurs between the three outer tepals and three inner tepals. The
outer tepals are thicker basally (Figs. 2C-D, 3C-D). The ovate to
subacute tepals are basally fleshy and enlarged throughout their midrib
regions. From the middle of the tepals to their bases, the outer and
inner tepals are broadly imbricated. During anthesis the tepal tips
which have adaxial papillae are outwardly recurved. Both tepal cycles
have a single midrib vein, that is, an outer tepal bundle (OT) in each
of the three outer tepals and an inner tepal bundle (IT) in each of three
inner tepals. Weakly differentiated nectiferous cells occur on the lower
inner bases of all six tepals.

Morphological elaboration occurs in the inverted receptacle as the
tepals and stamens are freed. The transition from the circular pedicel
to a broadly six-lobed receptacle is sudden (Figs. 2B-D, 3B-D). The

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VOL. 48

Fig. 1, — Herbarium specimens of Theropogon pallidus Maxim, showing gross vegetative
and floral morphology. A) Specimen from Nepal showing the root and shoot systems
{Kanai 610,100 TI). B) Size gradient in basal leaves, prominent median costa evident,
leaves longer than inflorescence. C) Thick fibrous root from stout rhizome. D) Upper
flowering raceme showing nodding, campanulate flowers each subtended by both a bract
(B) and a bracteole (Brt), (Scale indicated) (Photographs courtesy of the University
Museum, University of Tokyo, Japan, TI).

1979

Utech — Theropogon Floral Vascular Anatomy

29

lateral, inner margins of the outer are freed first, then the adaxial zones
along the midribs. At this level, the midribs of the outer tepals are
fleshy (Figs. 2D, 3D). Following this outer tepal initiation, the gynoe-
cium is partially freed from the adnate outer stamen and outer tepals
along the outer (dorsal) carpellary wall (Figs. 2E, 3E). Shortly there-
after all six tepals are freed from the gynoecium. A shallow filament
tube results from the lateral connation of the filament bases which
surround the gynoecium (Figs. 2F, 3F). The depth of the filament tube
corresponds to the distributional zone of the nectiferous cells on the
lower, adaxial tepal bases. Though there is limited epitepaly between
the tepal and the stamen cycles, the lateral connation between the
dilated filament bases in conjunction with the nectiferous tepal bases
probably has a functional significance in the pollination of this species.
In Conv allaria, the filaments are adnated to the connate floral tube,
and there are no nectiferous cells.

The six stamens, the three outer and the three inner, are equal, much
shorter than the tepals, incurved and laterally dilated (fleshy) through-
out. The filaments are attached to the lower abaxial anther surface
between the divergent outer anther lobes (introrse). Each of the yel-
lowish brown anthers has four cells, and dehiseces via lateral slits.
The anthers are tilted inwards and ring the upper gynoecium. Connec-
tive tips extend beyond the anther sacs.

Vascularization of the tepal and stamen cycles is simple and direct,
and involves sets of fusion bundles (Figs. 2B-~E, 3B-E, 4). At the base
of the receptacle there are three pairs of bundles. The three outer tepal
medians (OT) are formed first along radii between the three bundle
pairs via fusing branches derived from each pair member. The inner
tepal medians (IT) have a similar pattern of origin from the lower
receptacle bundle pairs. The inner median (IT) formation involves ra-
dially derived branches followed by fusion between different members
of the lower receptacle bundle pairs (Fig, 4). Both the OT and IT
medians are fusion products. A given OT is 120° from another OT, and
60° from an adjacent IT bundle. The formation and departure of the
tepal supply is extremely horizontal within the expanded receptacle.
Each free tepal has a single trace. There is no further radial division
basally or within the laminal surface to form tepal laterals. The tepal
traces are large in cross-section with the xylem portion broadly two-
armed.

The stamen traces are derived in a manner which parallels that of
the tepal medians (Fig, 4). The three outer stamen traces (OS) are
formed via radially derived branches from the continuing lower recep-
tacle bundle pairs which fuse along the OT radii. The branches, which
formed the OS traces, were derived from the same side of the lower
bundle pairs as the OT traces. Radial divisions on the opposite sides

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VOL. 48

1979

Utech — Theropogon Floral Vascular Anatomy

31

of these pair bundles and fusions along the IT radii result in the for-
mation of the three inner stamen traces (IS) (Fig. 4). All six stamen
traces depart horizontally through the receptacle soon after their for-
mation. Both the tepal and stamen vascularization originates within a
limited vertical distance. The various sets of tepal and stamen bundles
are derived from the three, lower, axial, receptacle pairs. These same
axial bundle pairs continue and establish the total gynoecial vascular
supply.

Gynoecial Morphology and Vascularization

The ovoid to subglobose gynoecium in Theropogon pallidus is non-
stipitate, tricarpellate, and unilocular terminally (Figs. 2D-G, 3D-~G).
In the various floral stages examined, rhaphides were not observed,
though they were common in Conv allaria (Utech and Kawano, 1976).
Though dorsal grooves are lacking, there are longitudinal septal
grooves in the upper gynoecium which continue up the complete length
of the narrow elongated style (Figs. 2H~~I, 3H-I). However, these
upper septal grooves showed no evidence of forming septal nectaries.
Though the fruit is a black, fleshy berry, there are persistent gynoecial
indications of weakened zones along the septa. The anther tips attain
approximately the same height as the upper ovary. The style length
including the minute stigma is approximately twice that of the flow-
ering ovary.

As the outer tepals are cut off (Figs. 2D~E, 3D-E), three fusion
dorsals (D) are formed like the OT and OS traces via radial branches

Fig. 2. — ^Serial cross-sections of Theropogon pallidus. A) Circular midpedicel section
with the three-bundled configuration. B) Trilobed upper pedicel with the six-bundled
configuration which is due to radial divisions. C) Six-lobed lower receptacle showing
the formation of the fusion outer (OT) and inner (IT) tepal median bundles from the six,
continuous, axial bundles; the OT bundles are formed at a lower level than the IT
bundles. D) Upper receptacle showing the formation of the fusion outer (OS) and inner
(IS) stamen traces from the six continuing axial bundles, both the OT and OS bundles
and the IT and IS bundles share a common radii. E) Receptacle above D showing the
tepal and stamen formation, fusion dorsals (D) are present along the OT-OS radii. F)
Lower gynoecium surrounded by a laterally connate staminal tube, both tepal cycles
freed; each fleshy tepal has a single unbranched fusion bundle; dorsals in peripheral
position; fusion septal axials (SA) formed between the inner paired sets of ventrals (V)
along the IT-IS radii. G) Midgynoecial section showing the fused central placentae; the
locules have opened transverse to the OT-OS-D radii; the septal axials (SA) have ter-
minated; the free filaments are laterally dilated; the tepal cycles have been omitted. H)
Upper gynoecial section showing the centrally free placentae; ventrals (V) present on
the inner septal wing margins; six large anthers surround the upper gynoecium; the tepal
cycles have been omitted. I) Mid-sty lar section showing the tripartite, unilocular cavity;
septal margins grooved; the dorsals (D) evident and unbranched; both the tepal stamen
cycles have been omitted.

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VOL. 48

Theropoflon pellidus Maxim.

Fig. 3. — Theropogon pallidus, cross-sections of Fig. 2 after transformational projections
with selected vascular bundles connected. The letter sections correspond to those in
Fig. 2.

1979

Utech — Theropogon Floral Vascular Anatomy

33

ODD

Fig. A.- — Roll-out longitudinal summary diagram for the floral vasculature of Theropogon
pallidus. The various text discussed bundles have been coded: OT = outer tepal, IT =
inner tepal, OS = outer stamen, IS = inner stamen, D = dorsal, SA = septal axial, and
V = ventral.

from the inner axial bundle pairs, which fuse (Figs. 2D-E, 3D-E, 4).
There is a direct, horizontal movement to the carpellary periphery of
the newly formed dorsals under the unopened locules. The three dor-
sals (D) remain in this peripheral position to the tip of the style without
branching or fusion.

Following the formation of the three dorsals (D), the freed gynoe-
cium in cross-section is smoothly six-lobed. Along the IT-IS radii
another set of three fusion bundles is formed. These three fusion bun-
dles are the septal axials (SA). Their location is slightly more periph-
eral than that of the six remaining central axial bundles. Though the
unbranched vertical course of the three septal axials (SA) is limited,
the septal axial formation is similar to that of the dorsal (D) (Fig. 4).
A given dorsal (D) is 60° from an adjacent septal axial (SA) (Figs. 2,
3F).

The simultaneous opening of the three locules is transverse to the

dorsal-center radii (Figs. 2G, 3G). The central placental region is large

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VOL. 48

and remains undivided up to the midgynoecium level. The gynoecium
base has 12 bundles, that is, the three septal axials (SA), the three
dorsals (D), and the six continuing axial bundles. The latter six bundles
function directly as the ventrals (V). The two bundles, which serve as
the ventrals (V) of a given carpel, were originally a pair and were
derived from a single pedicel bundle at a lower level (Fig. 4). Ovule
supply is directly via horizontal funicular (F) branches from the ver-
tical ventrals. Funicular branching occurs in the midcarpellary zone as
the central, massive placenta is subdivided along the dorsal-center
radii (Fig. 2H, 3H). The number of lateral funicular branches, which
supply the horizontal ovules, varies between three and five per ventral.
Therefore, there are on the average between 6 to 10 ovules per locule
or 18 to 30 ovules per gynoecium. The total seed production per gy-
noecium is, however, reduced to approximately half the ovule number.
Each fruiting pedicel has a subglobose to trilobed, pulpy, black berry
which is approximately 1.0 cm in diameter.

Distribution and Specimens Examined

Theropogon pallidus is a common, temperate zone, Himalayan re-
gion plant. Several general range descriptions have been given— -tem-
perate Himalayas, from Kumaon, alt. 60,000 ft to Sikkim, alt. 6,000 to
10,000 ft, Khasia Hills, alt. 5,000 to 6,000 ft (Hooker, 1875, 1894), and
Himalaya: Kumaon, Nepal, Sikkim, Khasia Hills, Tibet (Kihara,
1953). Nair and Sharma (1965) observed pollen from plants collected
in Cherrapunji (Assam).

The following cited specimens were observed at the University of
Tokyo (TI) in 1975. Besides detailed itineraries and maps, floristic
compilations of the University of Tokyo’s botanical expeditions to the

eastern Himalayas are available (Hara, 1966, 1971; Ohashi, 1975).

Theropogon pallidus (Wall, ex Kunth) Maxim., in Bull. Acad. Sci. St.

Petersb., i5:89. 1871.

Ohiopogon pallidus Wallich (Cat. no. 2138 (1829)) ex Kunth, Enum. 5:300. 1850.

Specimens examined — Bhutan: Dugye Dzong to Gunitsaura, elev. 2900 m, 21 June
1966, Nishioka s.n. (TI). Gunitsura to Cheka, elev. 2900 m, 24 June 1966, Nishioka s.n.
(TI). Nepal: Tare to Dunche, elev. 200 m, 25 June 1970, Kanai and Shakya 672,012
(TI). Gosinkund to Dhunche, 2100 m, 3 June 1969, Hara, Kanai, Kurosawa and Ohashi
69,614 (TI). Gosinkund, Ramche (1800 m) to Thare (2200 m), 2 June 1969, Hara, Kanai
and Kurosawa 69,615 (TI). Thare, elev. 6000 ft, on shady place on mossy rock, flowers
pink, 10 June 1969, Samen and Bista 13,032 (TI). Thare (2000 m) to Dunche (200 m),
3 June 1969, Kanai 670,100 (TI). Kalingchok, Thala (2050 m) to Tale Bisauna (2750 m),
10 September 1970, Kanai, Chuma and Nagano 674,258 (TI).

Summary and Concluding Remarks

The floral vascular anatomy of Theropogon pallidus has been pre-
sented, and the most salient features of this monotypic species include

1979

Utech — Theropogon Floral Vascular Anatomy

35

the following. The raceme has a spiral (1/3) phyllotaxy with each in-
verted, nodding flower subtended by both a bract and bracteole. This
six, rose-red tepals are free to their fleshy bases and are broadly im-
bricated laterally. A zone of nectiferous cells adaxially lines the outer
tepals. The tepal tips are outwardly recurved. Though different from
Conv allaria, which has a connate floral tube with adnate filaments,
the overall tepal configuration in Theropogon shows a campanulate
similarity with Conv allaria.

Basally the filament bases in Theropogon are laterally connate into
a shallow staminal tube which surround the inverted gynoecial base.
Beyond this level the freed filaments are broadly dialated. This stamen
modification does not occur in Conv allaria. In Theropogon the anthers
dehisce via lateral slits and are abaxially attached to the short filaments
along the lower one-third of the anther length (dorsifixed). The anther
tips are tilted inwards over the upper gynoecium. The six anthers con-
sequently present an appressed conical configuration. Apical connec-
tive extensions are present in all six anthers.

Pedicel vascularization consists of three large bundles each 120°
apart. Near the base of the inverted receptacle, the three pedicel bun-
dles divide radially to form a ring that contains three pairs of bundles.
It is via gap-closing fusion between and among the pairs of radially
derived branches that the tepal, stamen, and gynoecial vasculature is
established. The three outer tepal bundles (OT), the three outer stamen
traces (OS) and the three dorsal bundles (D) are gap-closing fusion
products formed along the radii, which bisect the three lower pedicel
bundles. The three inner tepal bundles (IT) and the three inner stamen
(IS) bundles similarily are gap-closing fusion products, but their radii
of formation are 60° from those of the outer whorl bundles. The six
ventrals (V), that is, the three continuing, unfused bundle pairs, supply
the ovules via simple, horizontal funicular (F) branches.

Table 1 presents a morphological and anatomical comparison of
Theropogon and Conv allaria. Though comparative emphasis is on the
floral vascular anatomy of both genera, other vegetative characters are
compared. Additional data on Convallaria for Table 1 have been taken
from Utech and Kawano (1976), Streveler (1966), Stenar (1941, 1953),
and Bjornstad (1970). The gross morphological differences between
Theropogon and Convallaria include the following. Mature flowering
specimens of Theropogon have numerous elongate grass-like leaves
with prominent costa, massive fibrous roots from a stocky rhizome,
and flowers, which are subtended by both a bract and a bracteole. The
three similar, but disjunct species of Convallaria (C. majalis L., C.
keiskei Miq., and C. montana Green), on the other hand, have flow-
ering individuals with long petiolate, paired leaves, an elongate rhi-
zome with scattered roots, and a single, simple bract, which subtends
each inverted flower.

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VOL. 48

Table 1. — Floral comparison between Theropogon and Convallaria.

Character

Theropogon pallidus

Convallaria spp.

(Utech and Kawano, 1976;

and as indicated)

Number of species

Monotypic, endemic to the

Three temperate species;

and range

Himalayan region of

Nepal and India

Europe, eastern U.S. and
eastern Asia (Strevelar,
1966)

Root system

Stout rhizome with numer-
ous, thick, and clustered
roots

Elongate rhizome with

numerous, thin and
scattered roots

Sheath leaves

Basal, two-six,
membranaceous

Basal, two-six,
membranaceous

Main leaves

Annual; 12-16, distichous,
grass-like, narrowly
elongate leaves with
prominent median costa;
parallel-veined (Fig. 1)

Larger leaves

Length: 15.0-45.0 cm;
Mean: 36.0 cm

Width: 0.6-2.0 cm;

Mean: 1.4 cm

L/W Ratio: 25.70

Spring ephemeral; typically
leaves paired; petiolate;
strongly net-veined
(Streveler, 1966 —
composite)

Larger leaves

Length: 25.0-45.0 cm;
Mean: 34.0 cm

Width: 6.0-11.5 cm;

Mean: 7.5 cm

L/W Ratio: 4.05

Inflorescence

Terminal raceme with 5-
18~(20) inverted flowers;
raceme central and axial
to imbricated leaves;
raceme shorter than
leaves

Terminal raceme with (3)-5-
12-(20) inverted flowers;
raceme outside and lateral
to paired leaves; raceme
longer than leaves

Flowers

Trimerous, hypogynous, bi-
sexual, actinomorphic

Trimerous, hypogynous, bi-
sexual, actinomorphic

Bract

Present; linear-subulate,
3-nerved, persistent, one
per node (Fig. 1)

Length: 1.0-2. 4 cm;

Mean: 1.7 cm

Present; lanceolate, 1-
nerved, persistent, one
per node (Streveler,

1966 — composite)

Length: 0.6-1. 5 cm;

Mean: 0.9 cm

Bracteole

Present; linear, 1-nerved,
persistent, one per node

Length: 0.4-0.9 cm;

Mean: 0.6 cm

Absent

Pedicel

Down turned during anthe-
sis; triangular in cross-
section at node, circular
at receptaclar base;
sclerenchyma sheath
present in fruit

Down turned during anthe-
sis; triangular in cross-
section at node, circular
at receptaclar base;
sclerenchyma sheath
present in fruit

1979

Utech — Theropogon Floral Vascular Anatomy

37

Table 1. — {Continued)

Character

Perianth

(OT = outer tepal)
(IT = inner tepal)

Stamens

(OS = outer stamen)
(IS = inner stamen)

Theropogon pallidus

Vascularization:
three large compound
bundles each with broadly
V-shaped xylem; 120° apart

Pale pink to reddish rose;
non-persistent; fragrant
(?); campanuloid, tepals
free to base and broadly
imbricated; tepal tips with
adaxial papillae and re-
curved outwards; lower
adaxial base of outer
tepals with a zone of
nectiferous (secretory)
cells

OT: 1-nerved (fusion)

IT: 1-nerved (fusion)

No tepal laterals

OT and IT are fusion
bundles of receptacular
origin formed among and
between the three pedicel
bundles (Figs. 2-4)

Tepal and stamen traces free
from each other

Six filaments free from the
tepals and gynoecium to
their bases; free filaments
broadly dilated; filament
bases laterally connate
into a shallow staminal
tube; filament cycles equal

Anthers six, yellowish
brown; abaxially attached
to filaments within lower
Vi length (dorsifixed);
anthers tilted forward
to surround the upper
gynoecium; lateral de-
hiscence (introrse); 2-sacs,
4-celled, fibrillar thicken-
ings on endothecial cells;
connectives apically
extended

Convallaria spp.

(Utech and Kawano, 1976;
and as indicated)

Vascularization:
six compound bundles (3
large -I- 3 small) each
60° apart

Creamy to yellowish white
with light green on the
tepal tips; non-persistent;
sweet scented; campanu-
late, tepals fused into floral
tube; tepal tips free and
recurved outwards; necti-
ferous cells absent from
the inner surface of the
floral tube

OT; 1-nerved (fusion)

IT: 1-nerved (fusion)

No tepal laterals

OT and IT are fusion
bundles of receptacular
origin formed from the six
compound, axial pedicel
bundles

Tepal and stamen traces free
from each other

Six filaments adnate to a
floral tube for ca. Vi the
tube length; filaments

"" ovoid to circular in cross-
section; short, but equal
cycle lengths

Anthers six, yellow;
abaxially attached to
filaments within lower Vi
length (dorsifixed);
anthers tilted forward
to surround the upper
gynoecium; lateral de-
hiscence (introrse); 2-sacs,
4-celled, fibrillar thicken-
ings on endothecial cells;
no apical connective
extensions

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VOL. 48

Table \.— {Continued)

Character

Gynoecium
(D = dorsal)

(V = ventral)

(P = placental)

(SA = septal axial)
(F = funicular)

Theropogon palUdus

Both OS and IS bundles
are fusion products
formed within the
receptacle among and
between the three pedicel
bundles which divide

Tricarpellate, syncarpous,
non-stipitate (sessile),
unilocular terminally
subglobose; septal groove
(indentation) present, but
no septal gland, no dorsal
groove

Rhapides not observed

Locules open perpendicular
to the dorsal-center radii;
basal septa fused centrally
and freed in lower levels
prior to ovule attachment

Gynoecium trilobed in cross-
section (Figs. 2-3)

Placental epidermis weakly
stigmatoidal, but con-
tinuous into style

Style columnar; stigma

minute; trilobed in
cross-section

Embryogensis:

Polygonum-iypo.

(Stenar, 1953)

Ovule number — average of
eight per locule

Fruit: berry; black,
subglobose to shallowly
trilobed due to persistent
septal grooves; inde-
hiscent, pulpy; style per-
sistent

Dorsals (D) fusion products,
unbranched, no terminal
fusion, ends in upper style

Convallaria spp.
(Utech and Kawano, 1976;
and as indicated)

Both OS and IS bundles
are fusion products
formed within the
receptacle from the six
axial pedicel bundles

Tricarpellate, syncarpous,
non-stipitate (sessile),
unilocular terminally,
globose; no dorsal or
septal grooves or glands
present

Rhapides present

Locules open perpendicular
to the dorsal-center radii;
basal septa fused centrally
and freed in mid-ovary
above lower ovule tier
attachment

Gynoecium circular in cross-
section

Placental epidermis strongly
stigmatoidal and con-
tinuous into style

Style columnar; stigma
capitate; circular in
cross-section

Embryogensis:

A///Mw-type
(Stenar, 1941;

Bjdrnstad, 1970)

Ovule number — average of
four per locule

Fruit: berry; red, globose,
indehiscent; no dorsal or
septal fruit markings,
pulpy, style deciduous
with scar evident

Dorsals (D) fusion products,
unbranched, no terminal
fusion, ends in upper style

1979

Utech—Theropogon Floral Vascular Anatomy

39

Table 1. — {Continued)

Character Theropogon pallidus

Convallaria spp.

(Utech and Kawano, 1976;
and as indicated)

Ventral plexus: simple

Ventral plexus: simple

ascending ventral (V)

ascending ventral (V)

supply with direct, hori-

supply with parallel,

zontal funicular (F)

ascending placental

supply; two ventrals

bundles, funicular (F)

(V) per septum

traces associated (1:1)

with placental (P) bundles;

two ventrals (V) per

septum, four or more

placentals per septum

Septal axial (SA): one per

Septal axial (SA): two simple

septum; present in lower

bundles which may fuse

levels only, absent as

per septum; terminates in

locules open; no inter-

upper gynoecium without

connection to ventral

vascular inter-connection

supply

to ventral supply

Chromosome number Unknown

In = 38 (Strevelar, 1966;

Fedorov, 1969; Moore,

1973).

In both Theropogon and Convallaria, the flowers are nodding, but
there are significant tepal and stamen differences between the two
genera. In Theropogon the broadly imbricated tepals are free to their
bases and have a basal adaxial nectiferous zone, whereas in Conval-
laria, a fused floral tube is present and there is no nectiferous tissue
associated with the perianth. The tepal tips in both genera are out-
wardly recurved. The filaments in Theropogon are free to their bases
and there are laterally connate for a short vertical distance into a shal-
low filament tube. In Convallaria, on the other hand, the filaments are
adnate to the surrounding floral tube for most of their filament length.
In neither genus is there any evidence of tepal or staminal adnation to
the outer gynoecial wall. Whereas the “lily-of-the-valley” scent of
Convallaria is well known, it was impossible in this study, because of
the material used, to determine whether or not Theropogon had a
scent. None has been reported.

Significant differences also occur between the gynoecia of Thero-
pogon and Convallaria. In the former, septal grooves, which persist
into the fruit, are present, whereas these carpellary indentations are
lacking in Convallaria. The species of Convallaria have abundant rha-
phides distributed throughout the mature pericarps, septal wings, and
funicular stalks. No rhaphides were observed in Theropogon. Though

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VOL. 48

both genera have simple fusion dorsals, there are differences in their
ventral supplies. A fusion septal axial occurs only basally in each sep-
tum of Theropogon, whereas two simple ascending septal axials,
which may or may not fuse, occur in each septum of Conv allaria.
Funicular branches depart directly and horizontally in Theropogon
from the simple ascending ventral traces. In Conv allaria, on the other
hand, the ventral supply is more complicated. Besides the simple as-
cending ventral bundles, two per septum, placental bundles branch
from these ventral bundles and ascend parallel to them. It is from these
parallel ascending placental bundles that the funicular traces are de-
rived in Convallaria. The average ovule in Convallaria is 12, whereas
the average number in Theropogon is 24. Embryogenesis also differs
between the two gmem— Theropogon has a reported Polygonum -type
(Stenar, 1953), whereas Convallaria has a reported Allium-typt (Sten-
ar, 1941; Bjornstad, 1970). Though the chromosome number of Ther-
opogon is still unknown, comparison to the repeatedly verified count
of 2n = 38 for Convallaria (Strevelar, 1966; Fedorov, 1969; Moore,
1973) should prove interesting.

Though there are superficial similarities in the inflorescence and
floral vasculature, the overall dissimilarities far outweigh the similari-
ties (Table 1). Furthermore, whereas many of the similarities (Table 1)
are basic and widespread throughout the Liliaceae Alliance (Anderson,
1940), other similarities, such as the floral shape and berry fruit, which
has a different morphology and pattern of vascularization in Thero-
pogon and Convallaria, are undoubtedly the result of convergent evo-
lution. Parallel observations on the floral vascularization and carpel
morphology in Speiranthe and Reineckea, which are Englerian tribal
cohorts, are needed to fully assess the berry as an evolutionary unit
and as a taxonomic unit in the Convallarieae.

Acknowledgment

The author would like to thank the herbarium staff of the University Museum, Uni-
versity of Tokyo (TI), for their specimen assistance, and the following agencies for their
grant support during which this project was in part completed^ — the United States-Japan
Cooperative Science Program (GF-41367), the Japanese Society for the Promotion of
Science (Grant-in-Aid 934053), and the M. Graham Netting Research Fund (Carnegie
Museum of Natural History) through a grant from the Cordelia Scaife May Charitable
Trust. Special thanks are due Ms. Valerie Barankovich for her artistic assistance in
manuscript production.

Literature Cited

Anderson, C. E. 1940. Some studies on the floral anatomy of the Liliales. Unpublished
Ph.D. dissertation, Cornell Univ., Ithaca, New York, 142 pp.

Baker, J. G. 1875. Revision of the genera and species of Asparagaceae. J. Linn. Soc.,
Bot., 14:508-632.

1979

Utech — Theropogon Floral Vascular Anatomy

41

Bjornstad, I. N. 1970. Comparative embryology of Asparagoideae-Polygonateae, Lil-
iaceae. Nytt Magasin Bot., 17:169-207.

Engler, a. 1888. Liliaceae. Pp. 10-22, in Die oatiirlichen Pflanzenfamilien (A. Engler,
and K. Prantl, eds.), Engelmann Verlag, Leipzig, 2:10-91.

Fedorov, A. A. 1969. Chromosome numbers of flowering plants. Nauk, Acad. Sci.
U.S.S.R., Leningrad, 926 pp. (in Russian).

Fuchs, C. 1963. Fuchsin staining with NaOH clearing for lignified elements of whole
plants or plant organs. Stain Tech., 38:141-144.

Kara, H. 1966. The flora of eastern Himalaya. Univ. Tokyo Press, Tokyo, 744 pp.

. 1971. The flora of eastern Himalaya. Second Report. Univ. Tokyo Press, To-
kyo, 840 pp.

Hooker, J. D. 1875. Theropogon pallidus. Curtis' Bot. Mag., No. 6154.

- — . 1894. Flora of British India. VoL VI. Orchideae to Cyperaceae. L. Reeve and

Company, London, pp. 299-365.

Hutchinson, J. 1934. The families of flowering plants. Vol. IT Monocotyledons.
MacMillan and Company, London, ed. 1, 234 pp.

— . 1959. The families of flowering plants. Vol. 11. Monocotyledons. Clarendon

Press, Oxford, ed. 2, 260 pp.

Johansen, D. A. 1940. Plant microtechnique. McGraw-Hill Book Company, New
York, 523 pp.

Kihara, H. 1953. Fauna and flora of Nepal Himalaya. Vol. 1. Flowering plants and
ferns. Fauna and Flora Research Society, Kyoto University, Kyoto, 310 pp.

Krause, K. 1930. Liliaceae. Pp. 354-355, in Die natiirlichen Pflanzenfamilien (A.
Engler, and K. Prantl, eds.), Engelmann Verlag, Leipzig, 2 (15a):227-390.

Kunth, C. S. 1843. Enumeratio Plantarum. 5:300-301.

Maximowicz, C. 1871. Theropogon nov. gen. Bull. Acad. Imp. St. Petersb., 15:89.

Moore, R. J. 1973. Index to plant chromosome numbers 1967-1971. Regnum Vege-
tabile, 90:114.

Nair, P. K. K., and M. Sharma. 1965. Pollen morphology of Liliaceae. J. Palyn.,
1:38-60.

Ohashi, H. 1975. Flora of eastern Himalaya. Third Report. Bull. Univ. Mus. Univ.
Tokyo, 8:130-136.

Sass, j. E. 1958. Botanical microtechnique. Iowa State Univ. Press, Ames, 228 pp.

Stenar, H. 1941. Uber die Entwicklung des Embryosackes bei Convallaria majalis
L. Bot. Not., 1941:123-128.

. 1953. The embryo sac type in Smilacina, Polygonatum and Theropogon (Lil-

iaceae). Phytomorph., 3:326-338.

Streveler, B. E. 1966. A taxonomic study of the genus Convallaria (Liliaceae). Un-
published M.S. thesis, Univ. Wisconsin, Madison, 89 pp.

Utech, F. H. 1978a. Floral vascular anatomy of Medeola virginiana L. (Liliaceae-
Parideae = Trilliaceae) and tribal note. Ann. Carnegie Mus., 47:13-28.

— - — . 1978^. Comparison of the vascular floral anatomy of Xerophyllum asphode-
hides (L.) Nutt, and X. tenax (Pursh) Nutt. (Liliaceae-Melanthioideae). Ann. Car-
negie Mus., 47:147-167.

— . 1978c. Vascular floral anatomy of Helonias bullata (Liliaceae-Helonieae), with

a comparison to the Asian Heloniopsis orientalis. Ann. Carnegie Mus., 47:169-191.

— . \91M. Floral vascular anatomy of Pleea tenuifolia Michx. (Liliaceae-Tofiel-

dieae) and its reassignment to Tofieldia. Ann. Carnegie Mus., 47:423-454.

— . 1978c. Floral vascular anatomy of the monotypic Japanese Metanarthecium

luteoviride Maxim. (Liliaceae-Melanthioideae). Ann, Carnegie Mus., 47:455-477.

Utech, F. H., and S. Kawano. 1976. Floral vascular anatomy of Convallaria majalis
L. and C, keiskei Miq. (Liliaceae-Convallarinae). Bot. Mag. (Tokyo), 89:173-182.

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0/ CARNEGIE MLISELIM

CARNEGIE MUSEUM OF NATURAL HISTORY
4400 FORBES AVENUE ® PITTSBURGH, PENNSYLVANIA 15213
VOLUME 48 6 MARCH 1979 ARTICLE 4

FLORAL VASCULAR ANATOMY OF SCOLIOPUS BIGELOVII
TORREY (LILIACEAE-PARIDEAE - TRILLIACEAE)

AND TRIBAL NOTE

Frederick H. Utech^

Associate Curator, Section of Plants

Abstract

The floral vascular anatomy and carpel morphology of the coastal Californian endemic
Scoliopus bigelovii Torrey are presented and this is followed by a comparison to its
assumed Paridean relative of eastern North America Medeola virginiana L. The floral
vasculature of S. bigelovii is established from a 15-bundled, axial pedicel configuration,
which consists of an outer zone of 12 bundles and an inner zone of three large bundles.
Six of the outerzone bundles depart directly as the three outer and the three inner tepal
medians. Each of the six remaining outer zone bundles undergo two successive radial
divisions whereupon 12 tepal laterals and six ventrals are formed. Basally each of the
six tepals receives three bundles, that is, a median and two laterals. Whereas several
radial divisions occur laminally within the outer tepal laterals to create a maximum of
13 bundles, that is, 12 laterals and a median, no such radial division occurs within the
inner tepal laterals. Between each of the three pairs of ventrals a fusion septal axial of
short vertical duration is formed. Two-ranked, horizontal funicular traces depart directly
from each of the paired ventrals. Whereas the ventral vasculature has an outer zone
origin, the three dorsals and the three outer stamens (there are no inner stamens) are
division products of the three large inner zone bundles. The dorsals are unbranched and
terminate in the tips of the three recurved stylar arms. There is no interconnection
between the dorsal and ventral supplies.

Several aspects of the floral morphology are derived or advanced and relate to the
pollination and seed dispersal (myrmecochory) of this species. Basally the broad, re-
curved outer tepals are imbricated into grooves on the abaxial surface of the narrow
erect inner tepals. The adaxial surfaces of the outer tepals are lined with nectiferous

* Research and publication supported by the M. Graham Netting Research Fund through
a grant from the Cordelia Scaife May Charitable Trust.

Submitted 12 September 1978.

43

44

Annals of Carnegie Museum

VOL. 48

tissue which in conjunction with the basal tepal imbrication forms a floral tube. Opposite
and above these tepal nectaries are the extrorse anthers, which are also versatile. The
tricarpellate, syncarpous gynoecium is unilocular from the level of opening into the
common style. The triquetrous ovary is flattened along the dorsal area and ridged along
the ventral area. The bitegmic ovules are attached in two-ranked rows in the ventral
corners. Stigmatoidal tissue is continuous from the three recurved stigma tips down
through the common style and along the funicular-ventral margins. Whereas rhaphides
are not present, subepidermal cells with colored material are.

Floral vascular evidence has been added to the continuing discussion on the relation-
ships of the four genera of the Englerian Parideae (or the Hutchinsonian Trilliaceae).
The vascularization of S. bigelovii differs markedly from that of Medeola virginiana.
Besides these differences, previously reported differences in vegetative and floral mor-
phology, embryology, and cytology are summarized as a step towards an overall tribal
evaluation.

Introduction

Though distinctive and showy, the genus Scoliopus Torrey is small
with only two species-^, bigelovii Torrey and S. hallii Watson. These
two species are respectively confined to the Coastal and the Cascade
Mountain Ranges of California and Oregon. In California, the coastal
distribution of S. bigelovii (Fig. 1) approximates the generalized, pa-
leoendemic range of the “coastal redwood,” Sequoia sempervirens
(D. Don) Endl. Such similarities in present-day, relictual ranges are
usually indicative of an isolated Arcto-Tertiary element (Munz and
Keck, 1959; Stebbins and Major, 1965; Axelrod, 1976; Raven and Ax-
elrod, 1978). The Oregonian endemic, 5. hallii, on the other hand, is
only known from the western slopes of the Cascades and the coastal
mountains from Tillamook County south to the Californian line (Hitch-
cock and Cronquist, 1973). Therefore, within the mountains of the far
west Scoliopus hallii is replaced southward by S. bigelovii.

Since 1857 when Torrey first described and illustrated S. bigelovii
(t. 22) as a new species, it has repeatedly been illustrated (Regel, 1875,
t. 834; Hooker, 1897, t. 7566; Parsons, 1907:263; Thomas, 1961: Fig.
49). A photograph of the type specimen, which is presented (Fig. 2),
shows enough similarities to the illustration presented by Torrey (1857:
t. 22) to know with some certainty that this material was used for the
artwork. From this material (Fig. 2) a lectotype is designated.

Four common names have been used for S. bigelovii— "" slink pod,”
“slink lily,” “fetid adder’s tongue,” and “brownies.” Each name and
its usage denote some particular aspect of this species’ biology. Both
“slink pod” and “slink lily” are in reference to the looped twisting or
coiling of the elongated fruiting pedicels, which locates each terminal
“pod” on the ground near or under the parental leaves whereupon the
released seeds are dispersed by ants (myrmecochory) (Berg, 1959).
Scoliopus, Torrey (1857), means crooked foot and is also an allusion
to the tortuous course of the fruiting pedicels (Eig. 2). “Fetid adder’s

1979

Utech—Scoliopus Floral Vascular Anatomy

45

Fig. 1.— Coastal Californian distribution of Scoliopus bigelovii Torrey based on speci-
mens in the University of California-Berkeley (UCB) herbarium.

tongue,” on the other hand, combines a reference to the reddish-brown
mottling on the two, expanded, basal leaves and an unpleasant floral
odor, which is not unlike that of beached star-fishes (Parsons, 1907) or
decaying seaweed (Hooker, 1897). “Brownies” also refers to the gen-

46

Annals of Carnegie Museum

VOL. 48

Fig. 2 — The above sheet (NY) represents a mixed collection from two distinct gather-
ings. Both fortunately were the only two gatherings cited by Torrey (1857:145) in the
type description. One gathering is limited to flowering material, whereas the other is
chiefly of fruiting material. Both materials were used for the composite type description
and the accompanying illustration (t. 22) judging from the materials and the sketches on

1979

Utech — ScoLiopus Floral Vascular Anatomy

47

eralized reddish-brown mottling on the leaves as well as the floral
parts. Jepson (1922) pointed out that this common name was confined
to Humboldt County, California. The coastal populations of S. bige~
lovii in this county are clearly disjunct from those further south along
the coast (Fig. 1). Flowering occurs between January and February
within the total range and seed dispersal between May and June.

Taxonomically Scoliopus has been associated with Trillium, Med-
eola and Paris. This association began with Watson’s (1879) Liliaceae
monograph and has continued in the present Englerian (Engler, 1888;
Krause, 1930) tribe Parideae and the Hutchinsonian (Hutchinson,
1934, 1959) family Trilliaceae. Berg (1959, 1962<3, 1962i?) strongly ques-
tioned this association for both Scoliopus and Medeola. Eor the for-
mer, Berg (1959) presented a thorough investigation of the gross veg-
etative and floral morphology of S. bigelovii, which also including brief
observations on the floral vascular anatomy. Additional and critical
observations on the embryogenesis, pollination ecology, and seed dis-
persal provided abundant data (Berg, 1959) to seriously question this
association for Scoliopus (Berg, 1962^).

The present research report on the floral vascular anatomy of S.
bigelovii as a representative of the genus Scoliopus, which only has
two species, is part of a long range study on the evolution of the
liliaceous berry (Utech and Kawano, 1975, \916a, \91bb, 1976c; Utech
1978a, 1978/?, 1978c, 1978J, 1978c). Whereas the floral vascular anat-
omy of the monotypic Medeola virginiana has previously been pre-
sented (Utech, 1978a) and is here compared to that of S. bigelovii, a
final appraisal of the tribal relationships of Scoliopus must await the

the upper portion of the sheet. Consequently, the specimens from both gatherings are
potential types. In the lower left one-quarter, there are two flowering elements each
with elongated pedicels. These are the second of the gatherings cited by Torrey, that is
“specimens in full flower, collected by Mr. Samuels, but in what part of California we
have not been informed.” A note on the sheet under these two elements further indicates
“rec’d from Dr. Gray, Dec. 1856.” The other gathering is of a complete, mature spec-
imen with numerous, looped pedicels and is centered directly on the sheet. Most of the
flowers are past anthesis which corresponds to Torrey’s note, “past flowering early in
April.” This element is clearly the Bigelow collection made during the 1853-1854 Whip-
ple Railway Route Expedition (Torrey, 1857), and was cited first by Torrey and should
be designated the lectotype for Scoliopus bigelovii. The Samuels’ collection therefore
becomes a paralectotype. In the lower left corner is a standard Whipple Expedition label
with Dr. J. M. Bigelow printed as the collector, California as the locality, and Scoliopus
hand written, presumably by Torrey, on it. The lower right corner has this hand written
information — “Tamul Pass, California, Whipple’s Exped., Dr. Bigelow.” Torrey’s Ta-
mul Pass is presently Mt. Tamalpais, California, which is just north of San Francisco.
(Photograph courtesy of the New York Botanical Garden.)

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voL= 48

Fig. S.-^-Pedicel and receptacle vascularization in ScoUopus bigelovii. A) Upper pedicel
crosS“Section showing a large and small lobe and the two zones of vascular bundles,
which are in the 15-bundled configuration (25 x). B) Enlargement of A with emphasis on
the 15-buedled configuration, that is, the three large, inner zone bundles and the 12,
outer zone bundles which can be further subdivided into the three OTM establishing
bundles (arrows), the three ITM establishing bundles located 60° from each of the for-
mer, and six additional bundles each located between an OTM and an ITM establishing
bundle (40 x). C) Upper pedical showing two of the three large inner bundles, a marked
ITM, but no ITL at this level and the radial origin of the OTL bundles (white dotted
lines); the two OTM (top and bottom) are further from the center than the marked ITM
(40 X). D) Lower receptacle showing the origin of both the ITL and the OS bundles; the
two ITL which accompany each ITM are radially derived; each inner zone bundle which

1979

Utech — ScoLiopus Floral Vascular Anatomy

49

completion of our work on Trillium and Paris (manuscripts in prepa-
ration).

Materials and Methods

Floral material of ScoUopus bigeiovii used in this study consisted of buds through

fully opened flowers, as well as isolated gynoecia of varying ages. These flowers had
been fixed, processed through a TBA series and embedded in wax by Marion S. Cave
(Botanical Garden, University of California-Berkeley), and given as gift research ma-
terial for which the author is indebted. The isolated gynoecia had been fixed in Craf
fixative, whereas the other materials had been fixed in FAA. Both fixatives were noted
on tags within the wax. The collection locality was tag noted as “Muir Woods” (Marin
Co., California). Though no collection dates were given for these wax materials, a
reference to slide material made in 1948 by Cave (1966) indicated Muir Woods. Presum-
ably the materials used for these slides and that in the wax were from the same gathering
or at least from the same general area and collected at a different time.

Standarized paraffin sectioning (12-16/4,) and staining (safranin-methylene blue) tech-
niques (Johansen, 1940; Sass, 1958) were used on the above materials. A total sample
of 35 complete, serial cross-sectional series were prepared on flowers of varying ages,
as well as several serial longitudinal sections. Fig. 1 presents a distribution map for S.
bigeiovii based on specimens in the herbarium at the University of California-Berkeley
(UCB). Fig. 2 is a photograph of the lectotype of S. bigeiovii, which was collected north
of San Francisco (Mt. Tamalpais). Figs. 3-5 are photomicrograph composites, which
show selected aspects of the floral morphology and the associated vascular anatomy.
Figs. 6-8 are summary diagrams for the pedicel to stigma vascularization. The meth-
odology of vasculature diagramming and presentation, as well as the letter coding of
bundles, parallels that in our previous liliaceous studies (Utech, 1978u, \91W, 1978c,
\91M, 1978c). Although the letters for the various vascular bundles used here are similar
to those of our previous papers, a correspondence should be given between them and
those used by Berg (1959) for S. bigeiovii: D (dorsal) = m.s. (median carpellary strand);
V (ventral) = p.s. (placental strand). Table 1 compares the vasculature of S. bigeiovii
as presented in this paper with that of Medeola virginiana (Utech, 1978cf).

Observations

Pedicel Vascularization

The sympodial umbel of ScoUopus bigeiovii commonly contains
three to eight, rarely up to 12, flowers. Each flower is attached to a
slender, bractless, erect pedicel, which is shorter than or equal to the
leaves at anthesis. The flowering pedicels are green and often spotted
with reddish-brown dots, which according to Berg (1959) are “most
numerous on pedicels or pedicel parts exposed to the sun.” The length

divides forms an outer stamen (OS) trace abaxially and a dorsal (D) adaxially (35 x). E)
Upper receptacle showing the departure of the inner tepal vasculature (an ITM and two
ITL) and the OS traces; the dorsal and ventral establishing bundles are located centrally
(30 x). F) Lower gynoecium showing the formation of the septal axials (SA) between
each ventral (V) pair; subepidermal cell with stained (colored, cf. text) material indicated
(lower arrow); no rhaphides are present (35 x).

50

Annals of Carnegie Museum

VOL. 48

of the flowering pedicel usually averages 15.0 cm, although the fruiting
pedicel is longer and recurved to the ground. In cross-section the ped-
icel is triangular for most of its length. However, near the base of each
flower, the number of pedicel angles is six, that is, the three continuing
ones, each 120° apart and three smaller, alternating ones. The three
larger pedicel angles are continuous with the lower midribs of the three
outer tepals (Figs. 3 A, 6A-D, 7A-D).

Throughout the triangular portion of the flowering pedicel, a 15-bun-
died configuration occurs in cross-section (Figs. 3A-B, 6A, 7A). All
of these bundles have a normal phloem (abaxial) and xylem (adaxial)
arrangement. The 15-bundles can be grouped into two broad, concen-
tric zones. The inner-most zone contains three large bundles each of
which lies along a radius that passes through one of the pedicel’s tri-
angular edges (Fig. 3B, arrow). The outer zone contains 12 bundles.
Six of these are closer to the pedicel’s periphery than the remaining
six. The outer six are spaced 60° apart and establish at a higher level
the tepal median supply of both the outer and inner tepal whorls, that
is, the three outer tepal medians (OTM) and the three inner tepal me-
dians (ITM) (Fig. 8). The three bundles, which establish the three outer
tepal medians (OTM)f like the three larger inner bundles, are also on
the radii of the pedicel’s triangular edges. Besides the six tepal median
establishing bundles in the outer zone, there are six additional bundles
that alternate with them. This overall vascular pattern characterizes
the elongated pedicel for most of its length. Furthermore, Berg (1959)
presented a cross-sectional figure (Fig. 24), which clearly shows this
pedicel vascular configuration and organization in the short stem below
the level of leaf attachment.

Approximately 5 mm below the base of the flower or receptacle,
changes occur in both the pedicel’s cross-sectional shape and its vas-
cular configuration. The pedicel changes from three-angled to six-an-
gled. The three new lobes are smaller and alternate with the three
larger lobes. Radii which bisect the smaller lobes also pass through the
inner tepal medians (ITM). With an increase in the pedicel’s cross-
sectional area, the six tepal medians depart outward radially along the
radii of the six lobes (Figs. 3C-D, 6A-C, 7A-C). Also at this level
there is a radial division within each of the six, outer, nontepal median
establishing bundles (Fig. 8). Consequently, the upper pedicel has 21
bundles, that is, the six tepal medians, the 12 outer zone bundles (six
pairs via the radial divisions), which alternate with the medians, and
the three larger inner bundles.

The erect flowering pedicel lacks a perivascular sclerenchymous
sheath. However, such a fibrous sheath is present in the twisted,
looped fruiting pedicel. The postanthesis pedicel recurvation is not a
geotropic response, but rather is due to a differential rate of elongation

1979

Utech — ScoLiopus Floral Vascular Anatomy

51

within the pedicel and the delayed appearance of the sclerenchymous
sheath. Before the fruit dehisces on the ground, the looped pedicel is
extremely turgid. A double pedicel loop with the fruit directed upwards
is not uncommon. The pedicel then, however, loses its turgidity and
collapses downward along with its terminal fruit. Subsequently the
dehiscing fruit with its oibappendaged seeds is grounded closer to the
parental plant than would have occurred had the pedicel remained
erect and had fallen laterally.

Tepal Morphology and Vascularization

The flowers of Scoliopus bigelovii are of the general liliaceous type,
viz. bisexual, actinomorphic, hypogynous, cyclic, and trimerous. They
deviate from the typical liliaceous flowers in only having three, outer
stamens and no inner stamens opposite the inner tepals. The flowers
are consequently tetracyclic.

Although the segments of the two tepal cycles are different, both
the outer and inner tepals are distinct and free. The broadly lanceolate
to ovate outer tepals are spreading and recurved at their midlengths,
and average 16.2 mm (range: 14.5-17.8 mm) in length and 6.9 mm
(range: 5. 8-7. 4 mm) in width. The outer tepals are showy. Their basic
color is greenish-yellow with prominent reddish-brown longitudinal
stripes. Berg (1959) notes that these stripes are due to “densely placed
individual color dots, which results from individual cells containing
coloring matter.”

The three, linear to subulate, inner tepals, on the other hand, are
erect with their upper tips curved inwards over the top of the gynoe-
cium. The lengths of the ascending inner tepals equals that of the
recurved outer tepals. However, a significant size difference occurs in
their width. The inner tepals only average 1.3 mm (range: 0.8-1. 7 mm)
at their widest point. The inner tepals also show a color difference
from the outer tepals. Both the upper adaxial and abaxial surfaces of
the inner tepals are a deep maroon purple, whereas the basal portions
of the inner tepals are a palish yellow adaxially and reddish-brown
striped abaxially. This reddish-brown basal striping is “again caused
by colored, papillate epidermal cells, arranged more or less distinctly
in three longitudinal stripes, one median and two marginal” (Berg,
1959). These colored stripes correspond to the locations of the inner
tepal’ s vasculature. Whereas the stripes on the inner tepals are re-
stricted to the lower abaxial surface, the numerous, colored stripes on
the outer tepals are found on the adaxial surface.

During anthesis, the open flower has a cup-like appearance, which
is due to the lower imbrication of the two tepal cycles (Figs. 4A-B,
6C-E, 7C-E). The outer tepal margins fit laterally into longitudinal
grooves on the abaxial surface of the inner tepals. This imbrication

52

Annals of Carnegie Museum

VOL. 48

Fig. 4,— Tepal and stamen vascularization in S. bigelovii. A) Basal tepal cross-section
showing the lateral edges of two outer tepals (OT) imbricated into abaxial grooves on
an erect inner tepal; the ITM and the two ITL are indicated; each inner tepal has a
maximum number of only three veins (40 x), B) Basal outer tepal (OT) and outer stamen
(OS) cross-section showing basally divergent anther sacs, the free filament (Fil) and the
large, multibundled outer tepal with nectiferous tissue adaxially; each outer tepal has
a maximum number of 13 veins, that is, six OTL plus an OTM plus six OTL (25 x). C)
Lower level stamen cross-section showing the free filament (Fil) with normally arranged
phloem and xylem in the OS bundle and the free anther in which the phloem and xylem
of the OS are reversed due to its downward course; another sac dehiscence is via
vertical, abaxial slits (extrorse); the outer tepal nectiferous tissue is still present (25 x).
D) Midlevel stamen cross-section showing the fusion of the free filament and the free
anther as well as stigmatic tip of the recurved stylar (St) arm (cf. Figs. 6I-J; 7I-J) (25 x),
Scoliopus bigelovii only has three outer stamens.

(Fig. 4A--B) persists up to the level where the outer tepals recurve.
The lower inner surfaces of the outer tepals have a broad, centrally
located band of nectiferous cells (Fig. 4B), which extends from the
level where the outer tepals are freed up into the recurved region. The
tepals are cut-off from the lobed or angled regions of the upper pedicel.
The three outer tepals are freed first with the three outer stamens
(filaments) adnated to them for a short vertical distance. The imbri-

1979

Utech — ScoLiopus Floral Vascular Anatomy

53

cated tepal arrangement and basal outer tepal nectaries are directly
related to the outbreeding pollination mechanism of this species.

As the morphology of the two tepal whorls differs significantly, so
too does the vascularization of these two whorls. Within the expanded,
six-lobed upper pedicel, the three outer tepal medians (OTM) and the
three inner tepal medians (ITM) depart outward along the six lobe
radii. There is no division within or fusion among the medians of either
whorl from their axial course in the pedicel to their termination in the
upper tepal tips. With the departure of the six medians, the outer zone
of the upper pedicel has 12 remaining bundles, that is, the six pairs
noted earlier. The two bundles, which are adjacent to the departing
outer tepal median (OTM), are pair halves of two different radial di-
visions and they depart outward with the centered outer tepal median
(Figs. 6A-D, 7A-D). Three such two-bundle sets, that is, a total of six
of the 12 outer ring bundles, exhibit this departure pattern, and directly
establish the outer tepal lateral (OTL) vascular network.

Although each outer tepal receives three bundles basally, that is, the
outer tepal median (OTM) and two outer tepal laterals (OTL) (Figs.
6A-D, 7A-D, 8), the expanded recurved outer tepals usually have 13
vein bundles, that is, the outer tepal median (OTM) and 12 laterals. Six
tepal laterals (OTL) are on each side of the OTM. To form this large
number of laterals, the two basally derived laterals (OTL) each
undergo a series of successive radial divisions (Figs. 6D-H, 7D~H, 8).
There is no cross-connection between the laterals within the laminal
tepal surface and each laterals ends marginally within the upper tepal
surface.

Another radial division occurs among the six remaining outer zone
bundles. The level of this second radial division is within the recep-
tacle. The division product halves which are adjacent to the departing
inner tepal medians (ITM) depart with the centered median (Figs. 4A,
6C-H, 7C-H, 8). These bundles are the inner tepal laterals (ITL).
Basally, therefore, each inner tepal receives a median (ITM) and two
laterals (ITL) (Fig. 8). But unlike the outer tepals, there is no further
radial subdivision among the laterals within the narrow, inner tepals.
The six remaining bundles after the departure of the inner tepal laterals
(ITL) directly establish the gynoecium’s ventral supply (Fig. 8). The
bundles in the outer pedicel ring, therefore, establish the six, indepen-
dent tepal medians and via successive radial divisions the outer and
inner tepal laterals as well as the ventral supply.

Stamen Morphology and Vascularization

There are only three stamens in Scoliopus bigelovii and these three
are consistently in the outer stamineal position. No vestigial traces of
the inner stamens or of staminodes were observed in our material, nor

54

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VOL. 48

Fig. 5. — Gynoecial vascularization in S. bigelovii. A) Midlevel cross-section showing
the tricarpellate, unilocular gynoecium; the dorsal (D) bundles are along the flattened
sides, whereas the ventral network is in the ridged comers (20x). B) Enlargement of A
showing the paired ventrals in the comer; dorsals (D) indicated (40x). C) Upper gynoe-
cium cross-section with nine bundles, that is, the three dorsals (S) and the three pairs of
ventrals; stigmatoidal tissue covers the funicular attachment area and is continuous from
the basal locular region up into the common style (25 x). D) Cross-section showing the
transition from the upper ovary to the common style; the unilocular cavity closes as it
opened, that is, along the dorsal (D) radii; ventral bundle pairs still present; colored cells
present in the subepidermis (25 x). E) Cross-section showing the erect common style;

1979

VTEcnS coLiopus Floral Vascular Anatomy

55

have any ever been reported. The three stamens are basally adnated
to the outer tepals for a short distance, ca. 0.5 mm (Figs. 6E-H, 7E-
H). The free filaments are short (ca. 5 mm), slightly dilated in cross^
section and curved outwards at the level of anther attachment. The
greenish-yellow, oblong anthers are adaxially attached slightly below
their midlengths (Fig. 4B~D). Between the anther sacs, the connective
tissue is flat. The anther sacs are, however, free and divergent basally
(Fig. 4B). Due to the location and level of filament attachment, the
anthers are versatile. Dehiscence is extrorse, that is, via two vertical
abaxial slits (Fig. 4C-D), The endothecial cells, which line the anther
sacs, have banded thickenings.

Whereas the origins of both the tepal and ventral vasculatures can
be traced to the outer ring of pedicel bundles (Fig. 8), the origin of the
three outer stamens (OS) traces is from the three, large, inner pedicel
bundles (Figs. 3D-E, 6A-D, 7A-D). The three inner ring bundles share
common radii with the outer tepal medians (OTM). Each inner bundle
undergoes a division that leaves two bundles along each centered OTM
radius. This division is unlike the radial divisions associated with the
formation of the tepal laterals and the ventrals. It is an internal division
or splitting, whereby the derived outer stamen bundles appear to be
derived from within the large parental bundles. The outer most bundle
of each division is the outer stamen (OS) trace, whereas the inner most
becomes the dorsal (D) bundle (Figs. 3D-E, 6C-F, 7C“F, 8). The OS
traces may in all likelihood be fusion products due to the rapid opening-
closing division of the large (compound) inner bundles. The OS traces
depart directly and horizontally following their formation.

The OS traces exhibit a peculiar downturned course as they enter
the connective regions of the anthers. A cross-section through the
lower anthers shows a given OS trace in two places, that is, in the
filament where a normal arrangement of phloem and xylem occurs and
in the lower connective tissue where the arrangement of the conducting
cells is reversed (Fig. 4D). One could infer that the anther had been
inverted such that the existing extrorse condition had been derived
(evolutionarily) from a pre-existing introrse condition. The abaxial zone
of pollen dispersal is a definite adaptation for outbreeding, and con-
forms to the tepal geometry with its nectary.

the dorsals (D) are present, the ventrals are not (30x). F) Cross-section of the upper
style which is subdivided into three recurved stylar arms; the axis of common style
cavity is indicated by a white arrow; the dorsals in the recurved stylar arms have

reversed xylem and phloem (30x).

56

Annals of Carnegie Museum

VOL. 48

Gynoecium Morphology and Vascularization

Unlike most tricarpellate, syncarpous, liliaceous gynoecia in which
the dorsal regions correspond to the corner or ribbed portions of the
pistil, the dorsal regions in Scoliopus bigelovii are laterally flattened or
compressed and the ventral regions occupy the corner or ribbed por-
tions of the pistil (Figs. 5A™B, 6G-I, 7G-I). The compound gynoecium
is formed by limited fusion along the outer septal margins which sub-
sequently forms the three ribs. The gynoecium is triangular in cross-
section with the ventral regions occupying the points.

Furthermore, most liliaceous gynoecia have septal wings, which are
laterally fused and protrude into the common locular spaces. Septal
glands may often occur in areas where the lateral septal fusion is in-
complete. This septal inrolling and fusion usually results in the mar-

Fig. 6. — Serial cross-section ofS. bigelovii. A) Three-lobed mid-pedicel with 15-bundled
configuration, which consists of three bundles in an inner zone and 12 bundles in an
outer zone. B) Six-lobed upper pedicel with 21 -bundled configuration, which occurs
following the origin of the six outer tepal laterals (OTL); the six tepal medians (3 OTM
plus 3 ITM) are peripheral in the six lobes; and three inner zone bundles remain centrally
located. C) Upper pedicel transition to the receptacle with 21 bundles; the outer and
inner tepals defined. D) Receptacle base showing the origin of the inner tepal laterals
(ITL) via radial division of outer zone bundles, and the three outer stamens (OS) and
the three dorsals (D) via divisions along the OTM radii; radial division has occurred in
the outer tepals to increase the number of laterals present; the three inner zone bundles
divided to form the dorsals adaxially and the outer stamen bundles abaxially. E) Ad-
ditional radial division within the outer tepals to increase the number of laterals; the
inner tepal laterals (ITL) depart with their centered median (ITM); a nine-bundled con-
figuration remains centrally, that is, the three dorsals (D) and the three pairs of ventrals
(V). F) Gynoecium freed from the tepals and stamens; the three outer stamens, there are
no inner stamens, are adnated to the outer tepals for a short vertical duration; the inner
tepals only have three bundles (an ITM and two laterals), whereas continued radial
division occurs within the outer tepal lateral supply; the three dorsals remain along the
OTM-OS radii, whereas the paired ventrals (V) close the gaps along the three ITM radii.
G) The three outer stamens (OS) are freed from the outer tepals; additional radial division
has occurred within the laterals of the outer tepals; the locule centrally and along the
dorsal radii; three septal axials (SA) are formed, one for each ventral pair; the outer
tepal lateral margins are imbricated into grooves on the abaxial surface of the small,
erect, inner tepals. H) Triquetrous gynoecium with flattened dorsal (D) sides and ridged
ventral (V) corners; the three septal axials have ended; ovule supply is via ranked,
horizontal funicular (F) traces from the corner ventral pair; stigmatoidal tissue present
in the ventral corners; each outer tepal has 13 bundles, that is, six OTL plus an OTM
plus six OTL. I) Upper gynoecium located centrally with three, recurved stylar arms
located peripherally; the phloem and xylem of the dorsals (D) in the stylar arms are
reversed in relation to the arrangement in the carpellary walls; the stylar arms of the
tripartite style pass over the top of the three extrorse anthers. J) Upper gynoecium with
only the three dorsal (D) bundles remaining in the common stylar portion and in the
three recurved arms.

1979

Utech — ScoLiopus Floral Vascular Anatomy

57

58

Annals of Carnegie Museum

VOL. 48

ginal placentae being centrally located with an associated reversal of
the phloem and xylem elements of the ventral bundles. Neither pro-
truding septal wings nor central placentae with reversed conducting
elements occurs in the gynoecium of S. bigelovii, and in comparison
to other liliaceous gynoecia, the gynoecium of the former must be
considered as secondarily primitive or highly reduced.

The gynoecium of S. bigelovii is also unusual in that it is unilocular
from its basal level up through the common hollow style. The locule
opens from the center outwards along the dorsal radii. Since the var-
ious gynoecial bundles remain in a peripheral position as the locule
opens to its maximum size, there is no inward movement of the ven-
trals (V) (Figs. 3F, 5A-B, 6F-I, 7F~I). The placentation is parietal,
due to the peripheral and corner position of the ventrals (Fig. 5A-B).
At anthesis the greenish color of the gynoecium is marked with red-
dish-brown dots that are located in the subepidermal cells of the peri-
carp. However, no rhaphides were observed in the gynoecium or in
the other floral elements of the various flowers of the different ages ex-
amined.

The upper gynoecium is also strongly three-angled (Fig. 5C) and
unilocular. As the common gynoecial cavity (locule) closes along the
three dorsal radii in a pattern similar to its opening, a hollow stylar
canal is formed. This canal is lined with papillate, stigmatoidal cells
(Fig. 5D~E), which are continuous from this stylar region down along
the three placental edges of the inner gynoecium (Fig. 5A~B) to the
basal level of the gynoecium. The common ascending style is short
and is subdivided into three, free, recurved, stylar branches (Fig. 5F).
This stylar recurving is along the dorsal radii. A cross-section in the
stylar region shows a given dorsal in two places, that is, in the style
with a reversed orientation of its xylem and phloem and in the car-
pellary wall or upper common style where the arrangement of the
conducting elements is normal. The recurved stylar branches pass over
the tops of the anthers (Fig. 4D). Each stylar branch has an upwardly
and outwardly open groove, which is directed away from the same
flower’s anther and pollen. The stigmas, which terminate each of the
well-developed stylar arms, are minute. The tripartite style with lo-
calized stigmas is a further adaptation for outbreeding.

The ovule position in S. bigelovii is peripheral, because the ventrals
(V) are along the angled periphery of the gynoecium and the placentae

Fig. 7. — S. bigelovii, cross-sections of Fig. 6 after transformational projections with
selected vascular bundles connected. Lettered cross-sections correspond to those in
Fig. 6.

1979

Utech-=tS'col/o/’c/5 Floral Vascular Anatomy

59

0 0 0

60

Annals of Carnegie Museum

VOL. 48

do not protrude into the common locule. The anatropous, bitegmic
ovules are arranged in loose, two-rowed ascending ranks. Each angled
periphery has two rows of ovules (Fig. 5A). The epidermis of the
distinct funiculus is stigmatoidal like the placentae. The funicular trace
(F) is a simple division product of the ventrals (Fig. 8), An average
ovule number of 30 is not uncommon for the gynoecium of S. bigelovu.

The oblong to lanceolate, mature fruit is strongly three-angled and
is terminated by the persistent, tripartite style. The pericarp is thin
and membranous with no fleshy or pulpy tissue evident. The reddish-
brown dots of the flowering gynoecium persist into the fruiting stage.
The dehiscence is irregular and causes by parenchymatic cell degen-
eration of pericarp tissue between the dorsals and ventrals. Each fruit
has six such zones. This splitting does not follow the normal zone of
weakness, that is, the dorsals (loculicidal dehiscence) or the ventrals
(septicidal dehiscence). The fruit is a capsule for the following reasons:
it dehisces; it remains attached to the pedicel and the plant; it pos-
sesses no pulpy tissue. Berg (1959) used the term “untypically locu-
licidal” for this type of capsule dehiscence and added further, that
“in comparison with other liliaceous capsules, the ScoUopus capsule
must be considered highly reduced, because of its lack of sclerenchyma
and normal dehiscence.”

The gyeoecial vascularization is relatively simple. As the tepals of
both cycles and the three outer stamens are freed, the gynoecial base
has nine distinct bundles (Figs. 3F, 6E-G, 7E-G). This vascularization
is all peripheral for three reasons. The gynoecium is widely unilocular
throughout, the placental margins are unprotrudieg and the septal wing
fusion is minimal. Three of the nine bundles are the dorsals (D). They
do not branch to form dorsal laterals or fuse throughout their vertical
gynoecial course (Fig. 8). However, the styles and their associated
dorsals are recurved. The dorsals (D) and the outer stamens (OS) share
a common axial vascular origin (Fig. 8).

The six remaining gynoecial bundles of the basal nine establish a
simple ventral supply. These six bundles, a pair between each dorsal,
are continuous below with those outer ring bundles of the pedicel that
underwent repeated radial divisions to form the lateral tepal network
(Fig. 8). A fusion product with a short vertical duration is formed
between each ventral pair located in the gynoecial angles. These three
fusion products are the septal axials (SA) (Figs. 3 A, 6F™G, 7F-G, 8).
The paired ventrals have a normal phloem-xylem arrangement due to
their peripheral position and the lack of inrolling of the lateral septal
wings (Figs. 5A-B). A pair of ventrals (V) supplies via ranked, hori-
zontal, funicular (F) traces two ascending rows of ovules (Figs. 5B, 8).
Following ovule vascularization, the ventrals continue upwards into
the common stylar region (Fig. 5C-D), but terminate as the tripartite,

1979

Utech — ScoLiopus Floral Vascular Anatomy

61

« c
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outer tepal median. These bundles merely share a common rdius.

62

Annals of Carnegie Museum

VOL. 48

recurved style is formed (Fig. 5E). There is no terminal interconnec-
tion between the ventrals or with the dorsal network (Fig. 8).

Summary of the Floral Vascular Anatomy of

SCOLIOPUS BIGELOVn

The floral vascularization of Scoliopus bigelovii is determined by a
multiaxial configuration of bundles, which are established in the stem
below the level where the pedicels are freed (Figs. 6-8; Table 1). The
pedicel configuration is constant throughout its complete length. All
of the pedicel bundles have a normal phloem (abaxial) and xylem
(adaxial) arrangement. In cross-section, there are two concentric rings
or zones of pedicel bundles. Within the inner ring or zone, there are
three large bundles, which establish via internal divisions the three
outer stamen (OS) traces and the three dorsals (D). The outer pedicel
ring or zone has 12 bundles, which can be subdivided into two groups
of six. One group of six directly establishes the tepal median supply,
that is, the three outer tepal medians (OTM) and the three inner tepal
medians (ITM), whereas the six remaining bundles undergo several
radial divisions each and thereby establish both the tepal lateral supply
(OTL and ITL) and the ventral network. Therefore, although the dor-
sals (D) have their origins from within the inner pedicel area, the ven-
tral supply is derived from the peripheral area.

All of the tepal medians and laterals of both whorls are derived from
bundles in the peripheral pedicel area, that is, from among the 12
bundles in the outer zone. The six median establishing bundles are 60°
apart, for example a given OTM is 60° from ITM on both sides and
120° from OTM on both sides. Between these six median bundles,
there are six additional bundles, which divide radially to form an outer
tepal lateral (OTL). A pair of OTL depart with each OTM. The con-
tinuing, axial portion undergoes another radial division to form an
inner tepal lateral (ITL). Similarly, a pair of ITL depart with each
ITM. Basally each outer and inner tepal receives three bundles, that
is, a median and two laterals. However, in the outer whorl each basal
tepal lateral undergoes several successive radial divisions to form a
maximum number of six laterals. This division does not occur within
the inner tepals. Consequently, there is a significant difference in the
maximum number of tepal bundles within the two whorls. Each outer
tepal has six OTL plus an OTM plus six OTL for a total of 13 veins,
whereas each inner tepal has only three veins, that is, an ITL plus an
ITM plus an ITL.

The gynoecial base has a nine-bundled configuration, which is com-
posed of the three dorsals (D) derived from the inner pedicel (recep-
tacle) zone and the six ventrals (V) derived from the peripheral zone.
The dorsals (D) continue directly into the recurved tripartite stylar

1979

Utech — ScoLiopus Floral Vascular Anatomy

63

Table 1. — Comparison o/ Medeola virginiana and Scoliopus bigelovii.

Character

Medeola virginiana L.

Scoliopus bigelovii Torr.

Underground parts
(Bell, 1974; Berg,

1959, mia, mib)

Stem tuber; sympodial
growth; duration 1 year;
dispersal by stolon;
vegetative reproduction
immediate and consider-
able; short-lived, fragile
roots, many roots per year

Rhizome; long-lived; con-
tractile roots

Stem

(Bell, 1974; Berg,

1959, 1962«, mib)

Terminates underground

stem; unbranched; erect;
leafy, not scapose; dies
down each season; scle-
renchyma present; hairy

Subterraneous; unbranched;
erect; leafy, not scapose;
dies down each season;
sclerenchyma cylinder
absent

Leaves

(Bell, 1974; Berg,

1959, 1962a, mib)

Several; clustered on stem,
in two groups; verticillate;
convergent-reticulate
venation

Two; near stem apex; al-
ternate; parallel-veined

Inflorescence
(Berg, 1959, 1962fl,
mib; Utech, 1978a)

Subterminal, sympodial
umbel; fruiting pedicel
with sclerenchyma; short
pedicel recurved upwards
in fruit

Terminal, bractless,

sympodial umbel; fruiting
pedicel with scleren-
chyma; long pedicel
recurved downwards in
fruit

Flowers

(Bell, 1974; Berg,

1959, 1962a, mih;
Utech, 1978a)

Perianth segments distinct
and inconspicuous; tepal
whorls alike; perianth
deciduous; six stamens in
two whorls; filaments free,
hypogynous; anthers
dorsifixed; extrorse de-
hiscence via vertical
(abaxial) slits; single
pollination unit; no
nectaries present;
hercogamous

Perianth segments distinct
and showy; outer tepals
petaloid, inner linear;
perianth deciduous; three
outer stamens; filaments
free, hypogynous;
anthers versatile;
extrorse dehiscence
via vertical (abaxial)
slits; three pollination
units; tepal nectaries
present; not hercogamous

Gynoecium (Pistil)

(Berg, 1959, 1962a,
1962^; Utech, 1978a)

Tricarpellous, syncarpous;
ovary globose; parietal
placentation; protruding
placentae; placentae be-
tween angles; placental
epidermis stigmatoidal;
seeds central and ± hori-
zontal; style nearly
absent; stigmata large;
apical part of carpels
deciduous

Tricarpellous, syncarpous;
ovary triquetrous;
parietal placentation;
unprotruding placentae;
placentae in angles of
iocules; placental epi-
dermis stigmatoidal;
seeds peripheral and
ascending; style well-
developed, tripartite;
stigma minute; apical
part of carpels
persistent

64

Annals of Carnegie Museum

VOL. 48

Table 1. — {Continued)

Character

Medeola virginiana L.

Scoliopus bigelovii Torr.

Fruit

(Berg, 1959, 1962fl,
1962^;Utech, 1978fl)

Berry; pulp from pericarp
and placentae

Capsule (cf. text); fruit not
berry descendant; de-
hiscense irregularly
loculicidal

Seed

(Berg, 1959, 1962«,
1962^; Utech, 1978a)

More or less rounded;
brownish, smooth and
unstriated; no appendage;
embryo small and straight

More or less rounded;
brownish, coloring un-
even; minutely pubescent,
striated; appendage small
via cell enlargement
only; oil in appendage;
embryo small and straight

Dispersal

(Berg, 1959, 1962a,
1962^)

Endozoochorous

Myrmecochorous

Ovule

(Berg, 1959, 1962a,
mib)

Anatropus shape; central
pleurotropous orientation;
indistinct funiculus;
stigmatoidal tissue
present; raphides absent

Anatropus shape;

hypotropous orientation;
distinct funiculus;
stigmatoidal tissue
present; raphides absent,
but colored cells present

Nucellus

(Berg, 1959, 1962a,
1962/?)

Embryo sac
(Berg, 1959, 1962a,
1962/?)

Medium size; periclinal
walls in epidermis absent;
basal part short
Fritillaria-iypQ length/
width ratio 2.5; haustorium
absent; synergid filiform
apparatus absent; polar
nuclei do not fuse before
fertilization; 3-2 anti-
podals; nuclear type of
endosperm

Large size; periclinal

walls in epidermis present;
basal part medium
Normal-type (monosporic);
length/width ratio 2.5;
haustorium absent;
synergid filiform apparatus
present; polar nuclei
fuse before fertilization;

3 antipodals; nuclear

3 antipodals; nuclear
type of endosperm

Pedicel vascularization
(Anderson, 1940;

Berg, 1959, 1962a;
Utech, 1978a)

12-bundled configuration;
broad zone with six outer
bundles alternating with
six inner bundles; all
bundles with normally
arranged phloem and
xylem

15-bundled configuration;
peripheral zone with six
outer bundles alternating
with six inner bundles
plus three large inner
zone bundles; all bundles
with normally arranged
phloem and xylem

Tepal vascularization

OTM = outer median

ITM = inner median

OTL = outer lateral

Outer tepal:

OTL -h OTM + OTL

Inner tepal:

ITL + ITM + ITL

Outer tepal:

6 OTL -h OTM + 6 OTL

Inner tepal:

ITL + ITM -h ITL

1979

Utech^-tS COL/OF f/s Floral Vascular Anatomy

65

Table 1.— {Continued)

Character

Medeola virginiana L.

Scoliopus bigelovii Torr.

ITL = inner lateral

OTM and ITM directly
derived from axial pedicel

OTM and ITM directly
derived from axial pedicel

(Anderson, 1940;

• supply; OTL and ITL

supply; OTL and ITL

Berg, 1959, 1962^;

derived via radial

derived via radial divi-

Utech, 1978fl)

divisions

sions; multiple laterals
in outer tepal

Stamen vascularization

Three outer and three inner

Three outer stamens only;

OS = outer stamen

stamens; OS and IS are

each derived via a planar

IS = inner stamen

fusion products via
radially derived branches;

division within the three
large inner zone bundles;

(Anderson, 1940;

OS and IS traces

shared origins with the

Berg, 1959, 1962^;

extending upwards in the

dorsals; OS recurved

Utech, 1978a)

connective tissue

downwards in connective
tissue

Gynoedal vasculari-

Base of gynoecium:

Base of gynoecium:

zation

D = dorsal

3D + 6V + 3SA

3 D + 6 V

V = ventral

Dl = dorsal lateral

followed by:

followed by:

Vl = ventral lateral

3D + 6Dl + 6V +

6 Vl + 3 SA

3 D + 6 V + 3 SA

SA = septal axial

F funicular

vib = ventral lateral

The D, Vl and SA bundles

The SA bundles are

branch

are fusion products, the

Dl and V bundles are not;

fusion products of short
vertical duration, whereas

(Berg, 1959, 1962a;

vertical duration of SA is

the dorsals (D) may be

Utech, 1978fl)

complete; neither lateral
nor terminal inter- or
intraconnection between
the dorsal and ventral
supplies; funicular (F)
supply horizontal and
direct from the ventrals
(V); average ovule num-
ber—18

Upper gynoecium:
both the D and vlb which
are division half products
of the Vl continue through
the common style into the
recurved stylar arms

compound, there is no
fusion involved in their
origin; neither lateral nor
terminal inter- or intra-
connection between the
dorsal and ventral
supplies; funicular (F)
supply horizontal and
direct from the ventrals
(V); average ovule num-
ber^30

Upper gynoecium:
only the D bundles are
in the upper common
style and continue into
the freed tripartite stylar
arms

Chromosome number

Basic chromosome number

Basic chromosome number

(Cave, 1966, 1970;

X = 1

X = 7, 8

Johansen, 1932;

Stewart and

Bamford, 1942;

Woodward, 1948)

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Annals of Carnegie Museum

VOL. 48

arms unbranched and without any fusion. A pair of ventrals (V) occurs

between each of the three dorsals (D) from the base to the upper
gynoecium (Figs. 6-8). A fusion septal axial (SA) of short vertical
duration is formed between each ventral pair. Ovule supply is via
direct, horizontal funicular (F) branches from the ascending ventrals
(V). Within each ventral pair the funicular branching is ranked. Each
of the three locular angles has an average of 10 ovules, that is, an
average of 30 ovules per gynoecium. There is no vascular inter- or
intraconnection between any bundles of the dorsal and ventral sup-
plies.

Though there are some shared similarities in the floral vascular anat-
omy of S. bigelovii and Medeola virginiana (Utech, 197&3), the dif-
ferences are significant (Table 1). The pedicels of both have multibun-
died axial configurations — S. bigelovii has 15 bundles, whereas M.
virginiana has 12 bundles. Basally both the outer and inner tepals in
both species receive three bundles, that is, a median and two laterals,
which is not unusual in the Liliaceae (Anderson, 1940). The six tepal
medians in both species are established directly and simply from axial
pedicel bundles, whereas the 12 tepal laterals in both species are the
result of radial divisions. In M. virginiana, both freed tepal whorls
have a maximum number of three bundles, which contrasts markedly
with the 13 maximum bundles in the outer whorl and three in the inner
whorl of S. bigelovii. The six stamen bundles and the three dorsals in
M. virginiana are fusion products from radial divisions within the re-
ceptacle’s periphery, whereas the three stamens and the three dorsals
in S. bigelovii are not fusion products and their origins are from more
central receptacle areas. The branched dorsal laterals (DJ that occur
in M. virginiana are not present in S. bigelovii. Though three fusion
septal axials (SA) occur in the ventral networks of both species, the
overall ventral network in M. virginiana is more complex than that in
S. bigelovii. In the former, in addition to the ascending ventrals and
the septal axials, there are three additional fusion bundles in the ventral
network. These fusion bundles, the ventral laterals (VJ, divide in the
upper gynoecium with the resulting ventral lateral branches (vlb) con-
tinuing along with the dorsals (D) into the terminal areas of the style.
There is no interconnection between the dorsal and ventral supplies
in either species above their level of origin within the receptacle.

Discussion and Concluding Remarks

The history and debate concerning the correct tribal position of Sco-
liopus is as remarkable as the plant. Torrey (1857:145) placed his new
genus Scoliopus at the end of the Melanthaceae “chiefly on account
of it extrorse anthers, notwithstanding its one-celled fruit and parietal
placentation. The somewhat dichlamydeous flowers are suggestive of

1979

Utech — ScoLiopus Floral Vascular Anatomy

67

Trilliaceae, but the extrorse anthers, as well as other characters, would
seem to forbid its being placed in that group.” The Melanthaceae used
by Torrey (1857) was based on an older, stricter concept of the Lili-
aceae Alliance (Gray, 1856), and essentially corresponds to the current
Englerian subfamily Melanthioideae (Engler, 1888; Krause, 1930).

Baker (1879:405), when revising the Liliaceae Alliance, placed Sco~
liopus in his Colchiaceae, a group that somewhat corresponded to the
older Melanthaceae in part. Yet within this rank, Baker (1879) did not
assign Scoliopus to a definite tribe. Rather he placed it in an appen-
daged group, “genera anomala Colchicacearum.” Earlier doubts of
Baker (1875:509) on the tribal status of Scoliopus resulted in his ref-
erence of this genus to a group containing aberrant liliaceous tribes.
One of these was the monotypic Scoliopeae.

In Watson’s (1879:222, 227) revision of the North American Lili-
aceae, Scoliopus is transferred from the older Melanthaceae to the
Trillieae, which then included Trillium and Medeola. {Paris is an Old
World genus.) The association of Scoliopus with Trillium and Medeola
dates from Watson’s monograph (1879). Watson’s tribe, Trillieae, was
redefined and enlarged with Scoliopus placed in its own subtribe, Sco-
liopeae, to show that Scoliopus was only remotely related to the other
genera.

Watson’s inclusion of Scoliopus in the Trillieae was accepted by
Bentham and Hooker (1883:762; cf. Medeoleae), by Engler (1888:83;
cf. Asparagoideae-Parideae) and others, and soon became the estab-
lished view on the genus’ classification. Doubts were expressed by
Hooker (1897), who noted in Curtis' Botanical Magazine that Scolio-
pus “clearly belonged to the tribe Medeoleae of Liliaceae, as deter-
mined by Bentham (Bentham and Hooker, 1883), but is not very
closely allied to any congener in that group.” However, modern taxo-
nomic treatments, for example those of Krause (1930:373; cf. Aspar-
agoideae-Parideae which follows Engler, 1888) and Hutchinson (1934:
8, 104; cf. Trilliaceae, which follows and elevates the association of
Bentham and Hooker, 1883), give the impression of a close natural,
and unquestionable relationship between Scoliopus and the other
members of the association (Berg, 1959).

Berg’s initial studies on the vegetative and floral morphology and
anatomy of both Scoliopus (Berg, 1959) and Medeola (Berg, 1962^)
were followed (Berg, 1962i?) by detailed embryological observations
for these two genera, as well as for Trillium and Paris, and concluded
with a comprehensive comparison and summary of the four genera in
the Englerian Parideae. Neither Scoliopus nor Medeola are at all sim-
ilar to each other (Table 1), and both are vastly different from Trillium
and Paris, which, on the other hand, do show a very high degree of
similarity. Furthermore, Berg (1962/?) concluded that only Trillium and

68

Annals of Carnegie Museum

VOL. 48

Paris should be kept together within the liliaceous tribe Parideae, that
Medeola should be transferred as a monotypic tribe (Medeoleae) to
the Englerian subfamily Lilioideae, and that Scoliopus should be re^
turned as a monotypic tribe to the subfamily Melanthioideae. How-
ever, considerable work still needs to be done within these two
subfamilies to demonstrate the affinities and relationships, if any, of
Medeola and Scoliopus to these proposed recipient subfamilies.

On all levels of biological organization differences occur between
Scoliopus bigelovii and Medeola virginiana. These differences have
been summarized in Table 1. Because both genera are small, Scoliopus
has two species and Medeola is monotypic, the summarized differ-
ences provide an excellent generic circumscription of each. Berg’s
reported (1959, 19626r, 1962/?) morphological and embryological differ-
ences are included in Table 1, as well as our reported (Utech, 1978a)
differences on their floral vascular anatomy.

The most remarkable features of S. bigelovii, in addition to its flow-
ers, are its reduced, subterranean stem with two alternate (not opposite)
distichous leaves, which are mottled reddish-brown, its contracted
sympodial umbel with elongate fruiting pedicels that are twisted and
recurved, and its "irregular loculicidal” capsules with seeds that are
ant dispersed due to oil-containing appendages (raphe derived), or
elaiosomes. The adaptive significance of the appendaged seeds, the
twisted to the ground pedicels, and the short stems are inter- related
into an advanced pattern of dispersal.

The structural adaptive significance of the showy flowers correlates
with its mode of pollination. Berg (1959) demonstrated that a single
Scoliopus flower actually functions as three "secondary flowers” or
independent pollination subunits, a feature usually associated with Iris.
Each of the three petaloid outer tepals is lined adaxially with nectarifer-
ous tissue and is laterally imbricated to form a broad cup or shallow
floral tube via the interdigitating of the lateral edges of the outer tepals
into grooves on the narrow, abaxial surface of the inner tepals. Be-
cause the triquetrous gynoecium is ridged along the ventral position,
which is opposite the narrow, appressed inner tepals and is flattened
along the dorsal surface, which is opposite the outwardly expanded
outer tepals and extrorse anthers, a configuration of three independent
pollination subunits is established. The three versatile stamens, and
there are consistently three, are in the outer position. Each extrorse
anther dehisces along two vertical abaxial slits such that the pollen
dispersal zone is within the space between the flatten dorsal region of
the gynoecium and the outwardly expanded outer tepal. The style is
tripartite terminally with each free stylar arm recurved over the top of
the anther. The minute stigmas are located terminally and along the
abaxial surface, which is removed from the pollen dispersal zone. The

1979

Utech — ScoLiopus Floral Vascular Anatomy

69

recurved style, the extrorse anthers, the unusual triquetrous gynoecium
and the large, nectiferous outer tepals are all inter- related for outbreed-
ing.

The embryology of S. bigelovii also differs from Medeola virginiana
(Table 1), as well as from Trillium and Paris (Berg, 1962^). Scoliopus
has a Normal-type (monosporic) of embryo sac, whereas Medeola has
a Fritillaria -iypQ (tetrasporic). Furthermore, Scoliopus possesses a
distinct funiculus with stigmatoidal tissue, a series of subepidermal
coloring cells and a fusion of its polar nuclei prior to fertilization.

Chromosome counts for both species of Scoliopus are known. Cave
(1966) reported from a single Oregonian population of S. hallii a count
of 2n = 14. However, Johansen’s report (1932) of 2n = 14 for S. bi-
gelovii was not confirmed by Cave (1966, 1970) who observed counts
of 2n = 16 from several different populations. One population from
Humboldt County, California, for example, had a somatic (root tip)
number of 2n = 16 but also had several extra small meiotic chromo-
somes, which indicates a certain degree of structural hybridity within
the northern part of the range. Consequently the chromosomal base
number of Scoliopus should be associated with both x = 7 and x =
8 until further research is done. Although Medeola virginiana also has
a base number of x == 7, there are significant morphological differences
between its karyotype and that of Scoliopus (Cave, 1970; Stewart and
Bamford, 1942; Woodward, 1948). Both Paris and Trillium are based
on a chromosome number of x = 5.

Scoliopus bigelovii Torrey, Pac. Rail. Report 4:145, pL 22. 1857.

Lectotype .—Tsimul Pass (Mt. Tamalpais), California, Whipple’s Rail-
way Expedition, April 1854, J. M. Bigelow s.n. (NY!)™-center
element; Fara/ecfo/ype.— California, Samuels s.n., s.d. (NY!)-—
lower left element, same sheet as above, Torrey ’s note: ‘Tec’d
from Dr. Gray, Dec. 1856,” cf. Fig. 2.

Specimens examined. — California: Humboldt Co.: Hubbard’s Station, 10 June
1899, J. B. Davy & W. C. Blasdale 5399 (UCB); Near Glendale on Mad River, elev. 0-
500 ft, 2 April 1905, J. P. Tracy 2156 (UCB); Near Hydesville, elev. 200 ft, 9 March
1913, J. P. Tracy 4017 (UCB); Dinsmore’s Ranch, Van Duzen River, opposite Buck
Mt., elev. 2500 ft, 20 June 1913, J. P. Tracy 4223 (UCB); Lawrence Creek, near Knee-
land Prairie, elev. 2000 ft, 9 March 1924, J. P. Tracy 6615 (UCB): Big Lagoon, elev.
300 ft, 26 March 1933, J. P. Tracy 10,941 (UCB), Kneeland Prairie, elev. 2500 ft, 26
April 1936, J. P. Tracy 14,808 (UCB). Marin Co. : West end of Alpine Lake, 13 February
1955, R. Y. Berg s.n. (UCB); Lagunitas Creek, 21 March 1891, Chesnut & Drew s.n.
(UCB); Mill Valley, 26 January 1913, A. Eastwood 2450 (UCB); Mt. Tamapais, 22
February and 30 March 1889, E. L. Greene s.n. (UCB); Mt. Tamalpais, Lagunitas Road,
1 February 1925, H. L. Mason 1231 (UCB); Sequoia Canyon, 31 January 1892, Michener
& Bioletti 2111 a (UCB); West side of Bolinas Ridge, east of Bolinas Lagoon, elev. ca.
750 ft, 23 February 1963, H. K. Sharsmith 5174 (UCB); Pipeline trail up Mt. Tamalpais,
30 March 1932, B. O. Schreiber 112 (UCB). San Mateo Co.: King’s Mt., 18 March

70

Annals of Carnegie Museum

VOL. 48

1902, C. F. Baker 322 (UCB); Lake Pelareitos, 28 February 1902, A. Eastwood s.n.
(UCB); San Andreas, s.d., A. E. Ehlers 278 (UCB); Above Portola on La Honda Road,
3 March 1939, R. F. Hoover 3897 (UCB); Pilarcitos Lake, 13 April 1946, F. A. Pitelka
s.n. (UCB); San Mateo Creek at SE end of Buriburi bridge, 12 March 1955, G. B.
Rossbach 214 (UCB). Santa Cruz Co.'. Big Tree Grove, April 1887, C. F. Sonne s.n.
(UCB); San Lorenzo River, ca. 3.5 miles from mouth of river near Southern Pacific
Railroad trestle, elev. ca. 300 ft, 24 March 1954, J. H. Thomas 2897 (UCB). Sonoma
Co.: Near Guerneville, 3 April 1898, M. S. Baker s.n. (UCB); Redwood Canyon ca, 2.5
mi S of Ft. Ross, elev. 100 ft, 25 February 1939, F. W. Gould 562 (UCB); Occidental,
18 March 1906, E. Lobenstein s.n. (UCB).

Acknowledgments

The author would like to thank the M. Graham Netting Research Fund of the Carnegie
Museum of Natural History (Pittsburgh) for supporting this research through a grant,
Ms. Marion S. Cave (University of California-Berkeley) for the gift of the embedded
floral material of Scoliopus, the director of the herbarium at the University of California-
Berkeley (UCB) for the use of their specimens, and the New York Botanical Garden
(NY) (Bronx) for the type photograph of S. bigelovii and its use. Special thinks are also
due Ms. Linda Plowman, Ms. Sally Garber, Ms. Siobhan Tierney, and Mr. Steve Harvey
for their technical assistance in the preparation of the slide material, and to Ms. Nancy
Perkins for artistic aid in the figure preparation.

Literature Cited

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Ph. D. dissertation, Cornell Univ., Ithaca, New York, 142 pp.

Axelrod, D. I. 1976. History of the coniferous forest, California and Nevada, Univ.
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Baker, J. G. 1875. Revision of the genera and species of Asparagaceae. J. Linn. Soc.,
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. 1879, A synopsis of Colchicaceae and the aberrant tribes of Liliaceae. J. Linn.

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Bell, A. D, 1974. Rhizome organization in relation to vegetative spread in Medeola
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Bentham, G., and j. D. Hooker. 1883. Genera Plantarum. London, vol. Ill, 1258
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Berg, R. Y. 1959. Seed dispersal, morphology and taxonomic position of Scoliopus,
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. 1962a. Morphology and taxonomic position of Medeola, Liliaceae. Skrifter det

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Engler, a. 1888. Liliaceae. Pp. 10-22, in Die natiirlichen Pflanzenfamilien (A. Engler,
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Gray, A. 1856. Manual of the botany of the northern United States. American Book
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Hitchcock, C. L., and A. Cronquist. 1973. Flora of the Pacific Northwest. Univ.
of Washington Press, Seattle, 730 pp.

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Hooker, J. D. 1897. Scoliopus bigelovii, native of California. Curtis Botanical Maga-
zine, t. 7566.

Hutchinson, J. 1934. The families of flowering plants. Vol. IT Monocotyledons.
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— . 1959. The families of flowering plants. Vol. II. Monocotyledons. Clarendon

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Krause, K. 1930. Liliaceae. Pp, 227-260, in Die natiirlichen Pflanzenfamilien (A.
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Jepson, W. L. 1922. A flora of California. Part VI. Liliaceae. Taylor, MacBride, and
Taylor, San Francisco, 250-321 pp.

Johansen, D. A. 1932. The chromosomes of the California Liliaceae 1. Amer. J. Bot.,
19:779-783.

— , 1940. Plant microtechnique. McGraw-Hill Book Co., New York, 523 pp.

Munz, P. a., and D. D. Keck. 1959. A California flora. Univ. California Press,
Berkeley and Los Angeles, 1681 pp.

Parsons, M. E. 1907. The wild flowers of California. Cunningham, Curtiss and Welch,
San Francisco, revised ed., 417 pp.

Raven, P. H., and D. I. Axelrod. 1978. Origin and relationships of the California
flora. Univ. California Publ. Botany, 72:1-134.

Regel, E. 1875. C. Scoliopus bigelovii Torr. Gartenflora, 24:227-228, t. 834.

Sass, j. E. 1958. Botanical microtechnique. Iowa State Univ. Press, Ames, 228 pp.

Stebbins, G. L., and j. Major. 1965. Endemism and speciation in the California
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Stewart, R. N., and R. Bamford. 1942. The chromosomes and nucleoli of Medeola
virginiana. Amer. J. Bot., 29:301-303.

Thomas, J. H. 1961. Flora of the Santa Cruz Mountains of California. Stanford Univ.
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Torrey, j. 1857. Report on the botany of the expedition. Pacific Railroad Rep., 4:1-
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Utech, F. H. 1978a. Floral vascular anatomy of Medeola virginiana L. (Liliaceae-
Parideae = TrOliaceae) and tribal note. Ann. Carnegie Mus., 47:13-28.

. 1978/?. Comparison of the vascular floral anatomy of Xerophyllum asphode-

loides (L.) Nutt, and X. tenax (Pursh) Nutt. (Liliaceae-Melanthioideae). Ann. Car-
negie Mus., 47:147-167.

— — — . 1978c’. Vascular floral anatomy of Helonias bullata (Liliaceae-Helonieae), with
a comparison to the Asian Heloniopsis orientalis. Ann. Carnegie Mus., 47: 169-191.

. \91M. Vascular floral anatomy of Pleea tenuifolia Michx. (Liliaceae-Tofiel-

dieae) and its reassignment to Tofieldia. Ann. Carnegie Mus., 47:423-454.

— . 1978£'. Vascular flora anatomy of the monotypic Japanese Metanartheciiim

luteoviride Maxim. (Liliaceae-Melanthioideae). Ann. Carnegie Mus., 47:455-477.

Utech, F. H., and S. Kawano. 1975. Biosystematic studies in Erythronium (Lili-
aceae-Tulipeae) IT Floral anatomy of E. japonicum Decne. Bot. Mag. (Tokyo),
88:177-185.

-. 1976a. Biosystematic studies on Maianthemum (Liliaceae-Polygonatae) VIII.

Vascular floral anatomy of M. dilatatum, M. bifolium, and M. canadense. Bot.
Mag. (Tokyo), 89:145-157.

— . 1976/?. Floral vascular anatomy of Convallaria majalis L. and C. keiskei Miq.

(Liliaceae-Convallarinae). Bot. Mag. (Tokyo), 89:173-182.

Watson, S, 1879. XV. Contribution to American botany. 1. Revision of the North
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apical bud chromosomes of Medeola. Bull. Torr. Bot. Club, 75:250-255.

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VOLUME 48 6 MARCH 1979 ARTICLE 5

A SYSTEMATIC REVIEW OF THE OLIVE BACKED
POCKET MOUSE, PEROGNATHUS FASCIATUS
(RODENTIA, HETEROMYIDAE)

Daniel F. Williams^

Post-doctoral Fellow, Section of Mammals

Hugh H. Genoways

Curator, Section of Mammals

Abstract

Geographic variation in Perognathus fasciatus Wied and the identities of specimens
of P. fasciatus and P. flavescens from areas of potential sympatry were investigated.
Populations of P. fasciatus from the northern Great Plains, in areas with the highest
amounts of precipitation, were the darkest colored and had proportionately the smallest
auditory bullae. Size varied clinally in the Great Plains, with larger mice being found in
the cooler, northern latitudes. Populations from the arid intermountain basins of Colo-
rado, Utah, and Wyoming were largest in size, had the longest tails, were the palest in
color, and had the largest bullae. Two races of the olive-backed pocket mouse are
recognized — P. fasciatus fasciatus Wied, 1839, from the Great Plains, and P. fasciatus
callistus Osgood, 1900, from the intermountain basins of Colorado, Utah, and Wyoming.

No evidence was found that would suggest interspecific hybridization between P.
fasciatus and P. flavescens. The two species are divergent in size and proportions and
occupy different habitats in the Great Plains. In the Unitah Basin of Utah and Colorado,
however, they are convergent structurally and appear to occupy similar habitats, but
are not known to be sympatric.

Introduction

Maximilian, Prince of Wied, obtained the first pocket mouse taken
in North America in 1833. He discovered the olive-backed pocket

‘ Permanent address: Department of Biological Sciences, California State College Stan-
islaus, Turlock, California 95380.

Submitted 13 September 1978.

73

74

Annals of Carnegie Museum

VOL. 48

mouse along the Missouri River, near the mouth of the Yellowstone
(near what is now Buford, Williams Co., North Dakota). In 1839, Wied
published his description of Perognathus fasciatus, which was excep-
tionally complete, and included a colored plate of the mouse and fig-
ures of the skull and teeth. However, for 50 years subsequent to the
naming of P. fasciatus, Wied’s (1839) excellent description was largely
ignored, and the name P. fasciatus was applied to populations now
known as P. hispidus Baird, 1858. Meanwhile, Baird (1858) included
specimens of the olive-backed pocket mouse in P.flavus Baird, 1855.

At the same time, he transferred P.flavus to Cricetodipus Peale, 1848,
which he relegated to subgeneric rank under Perognathus. Coues
(1875) later reinstated Cricetodipus to generic rank. Thus, specimens
of Perognathus fasciatus, the type species of the genus, were mistak-
enly assigned to a different genus and species.

Merriam (1889) untangled the web of nomenclatural errors woven
by Baird (1858) and subsequent authors. He was able to do so by using
a series of pocket mice collected near Maximilian’s type locality. Mer-
rian (1889) compared these specimens to Maximilian’s original descrip-
tion, and remarked that . .it allows no room whatever for difference
of opinion as to what his [Maximilian’s] animal really is^-it is the
Cricetodipus flavus (in part) of recent authors, my own specimens
from the Upper Missouri region agreeing in the minutest detail with
his careful description.” Merriam (1889) redescribed P. fasciatus from
a “duplicate type,” and named and described a new subspecies, P.
fasciatus flavescens from the Sand Hills region of Nebraska.

Shortly thereafter, Thomas (1893) described a new species, P. in-
fraluteus, from Loveland, Larimer Co., Colorado. He stated that his
new species was perhaps most nearly allied with P, fasciatus, but he
compared it in detail only with P. longimembris (Coues, 1875). In the
second and latest revision of Perognathus, Osgood (1900) assigned
infraluteus to subspecific rank under P. fasciatus, and elevated Per-
ognathus fasciatus flavescens to specific status. Osgood (1900) also
named an additional species, P. callistus, from the Green River Basin
of Wyoming, which he regarded as being most similar to P. apache '
Merriam, 1889 (= P. flavescens), but also sharing some external re- |
semblance with P. fasciatus.

Cary (191 1) described a new race of olive-backed pocket mouse from
Sun, Natrona Co., Wyoming, which he named P. fasciatus litus. He
characterized P. f. litus as being a little smaller than P. f fasciatus
and with an extremely pale color. Cary’s holotype was young, and had j
not attained mature proportions. Later, Swenk (1940) named a new
subspecies of pocket mouse, Perognathus flavescens olivaceogriseus, ■
from sandy soils of the Panhandle of Nebraska. He apparently pre- j
sumed that P. fasciatus did not occur on dune sands and Valentine f

1979

Williams and Genoways — Perognathus Systematics

75

sandy soils, so did not consider the possibility that his new form rep-
resented P. fasciatus.

Jones (1953) provided the only comprehensive review of the tax-
onomy and distribution of P. fasciatus. He showed that P. calUstus
was conspecific with P. fasciatus, but was recognizable as a subspe-
cies. He also demonstrated that P. flavescens olivaceogriseus was,
instead, a form of P. fasciatus. Jones (1953) noted that the color of
the pelage was the best character to distinguish the subspecies, and
remarked that color varied clinally from northeast (dark colored mice)
to southwest (pale colored mice). He pointed out that only the skulls
of P. f callistus could be distinguished from those of the other sub-
species. Jones (1964) later stated that specimens of P.f. fasciatus and
P. /. olivaceogriseus from Nebraska did not differ in cranial details,
and that their type localities both lie in a broad zone of intergradation
between the races.

Pefaur and Hoffmann (1974) investigated geographic variation in ex-
ternal dimensions of samples of P. f. fasciatus and P. f. olivaceogris-
eus. Their results showed no clearcut pattern of geographic variation,
but they did not draw any systematic conclusions from their study.
Williams (197&3) studied the karyotypic relationships among the
species of the subgenus Perognathus, and found a close similarity in
chromosome structure between P. fasciatus, P. flavescens, and P.
apache. Williams (1978^) investigated further the systematic relation-
ships of these three species, concentrating on those of P. apache with
P. fasciatus and P. flavescens. He concluded that P. apache and P.
flavescens were conspecific, and that P. fasciatus and P. flavescens
were structurally distinct where their geographic ranges meet in the
Uintah Basin of Colorado and Utah.

The present project was initiated in order to reexamine the patterns
of geographic variation in P. fasciatus, and to investigate the specific
identity of specimens from areas of potential sympatry between P.
fasciatus and P. flavescens in the Great Plains.

Methods

Thirty external, cranial, and dental traits were recorded for each specimen. Cranial
traits were measured with dial calipers, and were rounded to the nearest 0.05 mm.
Dental measurements were taken with an ocular micrometer, and are accurate to 0.03
mm. Unless they are otherwise defined, measurements are standard and are as illustrated
in Williams {\91%b). Traits used, their abbreviations, and appropriate remarks are listed
below.

Total length (TOTL). — Taken from specimen tags; recorded to nearest mm.

Length of tail (TL). — Taken from specimen tags; recorded to nearest mm.

Length of head and body (HBL). — Total length minus length of tail.

Length of hind foot (HFL).— Taken from specimen tags; recorded to nearest 0.1 mm
(most preparators recorded to nearest mm).

Length of ear (EL). — Taken from specimen tags; recorded to nearest 0.1 mm (most

76

Annals of Carnegie Museum

VOL. 48

Fig. L— Map, showing approximate localities of specimens of Perognathus fasciatus \

examined and localities for samples used in the analyses. Numbered areas indicate j

geographical samples included in the initial univariate and multivariate analyses (see
text). Lettered areas designate geographical groupings for the summary statistics (Table I
1). Localities with partially overlapping positions are indicated by single circles. I

collectors recorded only to nearest mm). Many preparators did not measure the ear and
some others measured the ear from the crown, rather than from the notch. Only mea-
surements that were apparently taken from the notch were included (that is, ^ 5 mm).

Greatest length of skull (GLS). — Greatest distance from the anteriormost projection j;
of the nasal bones to the posteriormost projection of the cranium. i:

1979

Williams and Genoways — Perognathus Systematics

77

Ocdpitonasal length (ONL). — Measured from the posteriormost point of the supraoc-
cipital region to the anterior tips of the nasals.

Least interorbital breadth (lOB).— Least width across interorbital constriction mea-
sured at right angle to longitudinal axis of cranium.

Alveolar length of maxillary toothrow (MXTL).— Least distance from anterior lip of
alveolus of to posterior lip of alveolus of M^.

Width across maxillary toothrows (WMXT), — Greatest distance between the labial
sides of the maxillary toothrows.

Length of bulla (BL).— Greatest length of mastoid bulla.

Width across bullae (BW)=-— Greatest width across the mastoid bullae, measured im-
mediately dorsal to the tube-like extensions of the tympanic bullae, at the anterior
borders of the external auditory meatuses.

Length of interparietal (IPL). — Greatest length of longest interparietal.

Width of interparietals (IPW),— Greatest width across the interparietals.

Length of nasal (NL).- — Length of longest nasal bone.

Width of nasals (NW). — ^Width across nasals at their widest point.

Width of rostrum (RW). — Greatest distance across rostrum at the junction between
the premaxillae and jugals.

Least interbullar distance (LID).-— Least distance between the mastoid bullae, mea-
sured on the dorsal surface of the skull.

Length of mandibular toothrow (MNTL). — Crown length of mandibular toothrow,
from front of P4 to back of M3.

Bullar extension (BE).^ — The distance the auditory bullae project posteriorly beyond
the occipital region, calculated by subtracting occipitonasal length from greatest length
of skull.

Length of articular process (LAP). — Greatest length of articular process of mandible,
measured from the posteriormost point of the process to the point of junction with the

coronoid process.

Occlusal lengths of P4, Mi, and M3.— Greatest lengths of the teeth, measured from
occlusal view. Dental abbreviations are standard; length is indicated by L.

Occlusal widths of P4, Mj, M3, P‘^, M\ and M^.— -Greatest widths of the teeth, mea-
sured from occlusal view. Abbreviations are standard; width is indicated by W.

All specimens were assigned to one of five age classes, based upon dental character-
istics which are more fully defined and illustrated in Williams (1978/?). Briefly, age classes
1 and 2 were juveniles with deciduous upper premolars. Subadults of age class 3 had
permanent premolars that had not reached occlusal level, or which were at occlusal
level, but showed no wear on the tips of the cusps. Age classes 4 and 5 were adults
with moderate (4) to heavy (5) wear on the cusps of P"*. Juveniles and subadults were
excluded from all interpopulation comparisons, and the two adult age classes were
pooled for statistical analyses.

Initially, samples of P. fasciatus were separated into 19 geographic groups (Fig. 1).
Standard univariate statistics were computed for samples of adult males and females,
both separately and combined. Some of the groups had too few individuals to make
intrasample comparisons, and two (15 and 16) were too small to include in intersample
comparisons. A single classification ANOVA (F-test, significance level = P ^ 0.05)
tested for significant differences between or among means. When means were found to
differ significantly, the sums of squares simultaneous testing procedure (SS-STP, Ga-
briel, 1964) was used to determine maximally nonsignificant subsets. Geographically
adjacent samples of P. fasciatus, without significant intersample differences, were com-
bined into four groups (Fig. 1, groups A, B, C, and D). Standard univariate statistics
were computed for these four samples, and three samples of P. flavescens from the
Great Plains. The sexes were treated both separately and combined.

Because many novice collectors had contributed to the external measurements ob-
tained from specimen labels, only cranial and dental characters (exclusive of greatest

78

Annals of Carnegie Museum

VOL. 48

length of skull and bullar extension) were utilized in discriminant function analyses
(BMDP7M; Dixon and Brown, 1977). The sexes were pooled, and only specimens with
complete measurements were used. Initially, 19 samples of P. fasciatus (samples 1-14,
16-19, plus ungrouped individuals, which were combined into a “classification only”
group) and one sample of P. flavescens were tested. In a second analysis, samples of
P. fasciatus were combined into two groups, and additional specimens were included
in the P. flavescens sample. The discriminant function/canonical analysis program
(BMDP7M) computes linear classification functions, in a step-wise manner, choosing
first the variable with the highest F value. Within and among groups variances are
recalculated for the remaining variables, a second variable is selected, and a new linear
discriminant function computed. This is repeated until all of the variables have been
entered, or until the F values are too low for further computation. Squared Mahalanobius
distance statistics and the posterior probabilities of belonging to each group are used to
place the individuals with one of the classification groups. Canonical variables are the
linear combination of variables entered that best discriminate among the groups. The
first canonical variable is the best discriminator; the second is the next best and is
orthogonal to the first. Groups submitted solely for classification do not contribute to
the computation of variable means, F values, or associated statistics. Character vectors
indicate the magnitude and direction of character effects on the canonical axes, and
were calculated by multiplying the scaled eigenvectors by the pooled within groups
standard deviation for each character (Power and Tamsitt, 1973).

Sample means for cranial and dental characters (exclusive of greatest length of skull)
of 18 samples of P. fasciatus (sample 15 was excluded because of missing character
values) were treated by a series of numerical taxonomic programs (MINT; Rohlf, 1971),
Data were standardized, taxonomic (Euclidean) distance and similarity (Q-mode cor-
relation) matrices were computed, and a principal components analysis was performed
on the matrix of correlation among characters. Phenograms were constructed from the
coefficients of the distance and similarity matrices by the UPGMA clustering routine
(unweighted pair-group method using arithmetic averages).

The specimens examined are listed in the systematic accounts, except for those of P.
flavescens copei, P.f cockrumi, and P.f. perniger, which were used in the discriminant
function analyses, but which were not otherwise included in this study. Information on
these latter taxa is available on request. The following institutions provided the speci-
mens for this study. The abbreviations preceding the institutions are used in the accounts
to identify the disposition of specimens. Further information concerning the collections
can be obtained from Choate and Genoways (1975).

BS — Biological Survey Collections, National Fish and Wildlife Laboratory (Washington,
D.C.).

BSC — Biological Survey Collections, National Fish and Wildlife Laboratory, Ft. Col-
lins, Colorado Field Station (this collection was formerly located in Denver).

CAS — California Academy of Sciences.

CM — Carnegie Museum of Natural History.

FMNH — Field Museum of Natural History (Illinois).

KU — University of Kansas, Museum of Natural History.

MMNH — University of Minnesota, James Ford Bell Museum of Natural History.

MSB — University of New Mexico, Museum of Southwestern Biology.

MVZ — University of California, Berkeley, Museum of Vertebrate Zoology.

ROM — Royal Ontario Museum (Canada).

SIUC — Southern Illinois University.

UCM — University of Colorado Museum.

UIMNH — University of Illinois, Museum of Natural History.

UM — University of Montana.

UMMZ — University of Michigan, Museum of Zoology.

1979

Williams and Genoways—Perognathus Systematics

79

UNDAK — University of North Dakota.

UNSM — University of Nebraska, State Museum.

UU- — ^University of Utah.

VMKSC— Kearney State College, Vertebrate Museum (Nebraska).

Results

Nongeographic Variation

Juveniles and subadults generally averaged smaller in external and
cranial characters (except for least interorbital breadth) than adults,
and were excluded from intersample comparisons. Old adults (age
class 5) were usually slightly larger than young adults (age class 4) but
the differences were slight. There were too few individuals in age class
5 to perform meaningful statistical tests; therefore, age classes 4 and
5 were combined for intersample comparisons.

There were no significant secondary sexual differences among the
15 geographic samples tested, and only one significant difference was
found between the sexes of the four geographic groups. Females of
group A (Fig. 1) had significantly greater lengths of interparietals than
males. As this was the only significant difference between the sexes,
they were pooled for the multivariate analyses, and only the pooled
univariate statistics are given (Table 1).

Little individual variation, other than meristic variation, was noted
in samples of P. fasciatus. All specimens of P. fasciatus were found
to have the auditory bullae welLseparated anteriorly (over the basi-
sphenoid bones). In contrast, from 20% to more than 50% of the in-
dividuals in samples of P. flavescens had bullae in contact (apposed)
over the basisphenoids. Populations of P. flavescens with the highest
percentage of apposed bullae were found in western Nebraska, but
this character was not consistent enough to positively distinguish P.
fasciatus from P. flavescens.

No accessory cusps or cusp deletions were noted on the upper pre-
molars of P. fasciatus (out of 261 specimens), whereas up to 40% of
some samples of P. flavescens exhibited such cusp anomalies (Wil-
liams, 1978Z?, and unpublished). Three (of 259) specimens of P. fas-
ciatus had the protostylid and metaconid of the P4 united into a single
cusp (see Williams, 1978i?), and one individual had these two cusps
closely apposed. Three other individuals had an accessory cuspule on
the protolophid of P4, between the protoconid and protostylid. In gen-
eral, such anomalies of the lower premolars were rare in P. fasciatus,
but were relatively common (up to 60% of some populations; Williams,
19781?, and unpublished) in P. flavescens.

Color varied individually, seasonally, and with age. The juvenile
pelage was generally darker and more uniformly gray than the adult
pelage. Specimens with fresh adult pelage had a darker appearance

80

Annals of Carnegie Museum

VOL. 48

Table 1. — Standard statistics for samples o/ Perognathus fasciatus and P. flavescens.
Abbreviations for traits are given in the text. See Fig. 1 for explanation of sample codes.
CV = coefficient of variation; SE = standard error of the mean.

Trait

N

Mean

SE

CV

Range

Perognathus fasciatus — Sample A

TOTL

15

128.2

1.143

3.45

121.0-140.0

TL

15

59.3

0.908

5.93

54.0-66.0

HBL

15

68.9

1.376

7.74

60.0-81.0

HFL

13

17.3

0.257

5.37

16.0-20.0

EL

7

6.8

0.260

10.06

6.0-8.0

GSL

14

22.11

0.145

2.45

21.30-23.20

ONL

15

22.06

0.140

2.46

21.30-23.15

lOB

16

4.86

0.041

3.36

4.50-5.15

MXTL

17

3.19

0.037

4.77

2.80-3.40

WMTR

17

4.30

0.035

3.33

3.95-4.55

BL

16

7.67

0.053

2.78

7.30-8.00

BW

16

11.68

0.068

2.35

11.15-12.20

IPL

16

2.71

0.048

7.07

2.35-3.10

IPW

16

5.01

0.070

5.55

4.60-5.60

NL

17

8.10

0.086

4.37

7.40-8.70

NW

16

2.30

0.028

4.89

2.00-2.45

RW

17

3.72

0.054

5.96

3.40-4.25

LID

16

4.82

0.067

5.58

4.40-5.50

BE

14

0.01

0.011

2.45

0.00-0.05

MNTL

17

2.78

0.030

4.46

2.50-2.90

LAP

17

2.62

0.033

5.16

2.25-2.80

PW

17

0.94

0.013

5.71

0.81-1.03

M^W

17

1.06

0.012

4.48

0.93-1.13

M^W

17

0.66

0.012

7.29

0.58-0.74

P4L

17

0.60

0.011

7.66

0.48-0.68

P4W

17

0.64

0.009

6.15

0.58-0.71

MjL

17

0.86

0.013

6.12

0.74-0.94

MjW

17

0.95

0.012

5.40

0.84-1.03

M3L

17

0.60

0.013

6.32

0.52-0.65

M3W

17

0.69

0.010

5.96

0.58-0.74

Perognathus fasciatus — Sample B

TOTL

49

130.5

0.950

5.09

112.0-146.0

TL

50

61.5

0.563

6.48

53.0-70.0

HBL

50

69.1

0.659

6.75

59.0-80.0

HFL

51

17.2

0.111

4.62

15.7-19.0

EL

30

6.9

0.091

7.24

6.0-8.0

GSL

44

22.23

0.084

2.51

20.95-23.70

ONL

44

22.22

0.084

2.49

20.95-23.70

lOB

49

4.89

0.026

3.71

4.50-5.35

MXTL

52

3.15

0.016

3.75

2.95-3.45

WMTR

49

4.33

0.016

2.56

4.00-4.55

BL

50

7.64

0.042

3.89

7.00-8.20

BW

47

11.77

0.042

2.45

11.15-12.40

IPL

49

2.67

0.029

7.65

2.30-3.00

IPW

49

4.75

0.036

5.26

4.10-5.20

1979

Williams and Genoways — Perognathus Systematics

81

Table i. — {Continued)

Trait

N

Mean

SE

cv

Range

NL

48

8.25

0.052

4.33

7.50-9.10

NW

47

2.21

0.017

5.26

2.00-2.55

RW

51

3.72

0.023

4.49

3.40-4.15

LID

49

4.60

0.040

6.15

4.10-5.20

BE

44

0.01

0.006

3.00

0.00-0.20

MNTL

48

2.79

0.015

3.78

2.50-3.00

LAP

50

2.68

0.017

4.53

2.40-2.95

FW

52

0.97

0.009

6.61

0.68-1.10

NEW

52

1.09

0.006

4.03

0.97-1.19

51

0.70

0.006

6.45

0.55-0.77

P4L

50

0.61

0.006

7.42

0.52-0.74

P4W

50

0.66

0.006

6.53

0.52-0.74

MjL

50

0.85

0.006

4.90

0.77-0.97

MiW

50

0.97

0.005

3.83

0.84-1.03

M3L

52

0.60

0.005

3.75

0.52-0.68

M3W

50

0.72

0.006

5.80

0.61-0.81

Perognathus fasciatus — Sample C

TOTL

90

131.4

0.758

5.47

113.0-147.0

TL

90

61.3

0.408

6.32

52.0-70.0

HBL

91

70.09

0.531

7.22

58.0-80.0

HFL

84

17.5

0.093

4.89

14.8-19.0

EL

52

6.95

0.085

8.79

6.0-8.0

GSL

72

22.47

0.084

3.19

20.80-24.20

ONL

71

22.47

0.085

3.18

20.80-24.20

lOB

88

4.95

0.022

4.18

4.55-5.60

MXTL

91

3.13

0.014

4.29

2.80-3.40

WMTR

79

4.36

0.017

3.43

3.90-4.70

BL

84

7.63

0.033

4.02

6.95-8.50

BW

77

11.74

0.040

3.00

10.85-12.55

IPL

86

2.62

0.022

7.74

2.15-3.05

IPW

85

4.79

0.034

6.62

4.00-5.60

NL

84

8.27

0.043

4.72

7.30-9.45

NW

86

2.24

0.014

5.78

1.85-2.50

RW

87

3.69

0.016

4.03

3.35-4.05

LID

81

4.68

0.027

5.27

4.20-5.30

BE

71

0.00

—

—

0.00-0.05

MNTL

83

2.78

0.011

3.51

2.55-3.05

LAP

87

2.72

0.019

6.76

2.35-3.05

PW

92

0.96

0.004

4.18

0.84-1.03

M^W

92

1.05

0.005

4.76

0.94-1.23

M^W

84

0.69

0.004

5.39

0.61-0.81

P4L

88

0.61

0.004

5.92

0.52-0.68

P4W

88

0.66

0.004

5.86

0.55-0.74

MjL

88

0.84

0.005

5.93

0.74-0.97

MjW

88

0.94

0.006

6.04

0.74-1.06

M3L

91

0.61

0.005

7.38

0.52-0.77

M3W

84

0.71

0.005

6.40

0.58-0.84

82

Annals of Carnegie Museum

VOL. 48

Table 1. — (Continued)

Trait

N

Mean

se

cv

Range

Perognathus fasciatus — Sample D

TOTL

75

134.6

0.863

5.58

113.0-149.0

TL

75

64.3

0.502

6.76

52.0-75.0

HBL

75

70.2

0.573

7.11

56.0-80.0

HFL

76

17.7

0.094

4.62

15.0-20.0

EL

73

6.99

0.081

9.87

5. 0-9.0

GSL

71

22.99

0.098

3.59

20.15-24.80

ONL

69

22.85

0.095

3.45

20.15-24.55

lOB

77

5.14

0.017

2.84

4.85-5.50

MXTL

77

3.21

0.014

3.78

2.95-3.50

WMTR

77

4.42

0.015

2.90

4.15-4.65

BL

76

8.53

0.039

4.01

7.45-9.35

BW

74

12.71

0.055

3.70

10.95-13.85

IPL

75

2.75

0.029

8.97

2.25-3.50

IPW

75

4.53

0.036

6.79

3.85-5.15

NL

73

8.36

0.061

6.21

6.45-9.35

NW

74

2.29

0.015

5.59

2.00-2.55

RW

77

3.67

0.016

3.89

3.40-4.00

LID

75

4.38

0.038

7.48

3.55-5.30

BE

76

0.14

0.014

8.17

0.00-0.55

MNTL

76

2.84

0.012

3.63

2.60-3.20

LAP

74

2.79

0.021

6.38

2.25-3.20

PW

74

LOO

0.005

4.40

0.90-1.10

M'W

74

1.08

0.005

4.31

0.97-1.19

M^W

77

0.70

0.005

6.62

0.58-0.81

P4L

73

0.65

0.005

6.79

0.58-0.81

P4W

74

0.68

0.005

6.61

0.58-0.77

MjL

74

0.87

0.006

5.83

0.77-l'.00

MiW

74

0.95

0.005

4.62

0.81-1.03

M3L

77

0.61

0.004

3.77

0.52-0.77

M3W

74

0.72

0.004

4.89

0.64-0.81

Perognathus flavescens flavescens

TOTL

79

124.3

0.809

5.79

105.0-141.0

TL

79

59.6

0.502

7.49

50.0-69.0

HBL

82

64.6

0.541

7.59

54.0-78.0

HFL

81

17.0

0.096

5.08

15.0-20.0

EL

62

6.5

0.079

9.49

5. 0-8.0

GSL

67

21.68

0.085

3.19

19.50-23.85

ONL

67

21.64

0.079

2.99

19.50-23.40

lOB

81

5.06

0.016

2.86

4.75-5.40

MXTL

84

3.07

0.011

3.18

2.85-3.30

WMTR

81

4.22

0.015

3.12

3.95-4.55

BL

81

7.20

0.033

4.11

6.45-8.20

BW

78

11.57

0.036

2.76

10.95-12.40

IPL

81

3.01

0.021

6.39

2.45-3.55

IPW

81

4.95

0.026

4.75

3.95-5.40

NL

70

7.79

0.042

4.47

7.05-8.75

NW

67

2.25

0.018

6.49

1.80-2.60

1979

Williams and Genoways — Perognathus Systematics

83

Table 1. — {Continued)

Trait

N

Mean

SE

cv

Range

RW

81

-h.ll

0.018

4.22

3.35-4.15

LID

79

4.88

0.034

6.14

4.00-5.60

BE

67

0.00

_

. —

0.00-0.05

MNTL

79

2.69

0.012

3.96

2.50-2.95

LAP

79

2.70

0.020

6.74

2.15-3.05

p4W

80

0.90

0.004

4.40

0.84-1.03

MiW

80

1.04

0.005

4.27

0.97-1.16

79

0.62

0.004

6.52

0.55-0.74

P4L

80

0.54

0.004

7.36

0.45-0.64

P4W

80

0.59

0.004

6.89

0.48-0.88

MjL

80

0.84

0.004

4.47

0.71-0.93

MiW

80

0.90

0.005

4.65

0.81-1.00

M3L

80

0.58

0.004

6.39

0.48-0.68

M3W

80

0.67

0.005

6.17

0.58-0.77

than those in worn pelage, due to the greater number of black-tipped
hairs. Some of the tips wear off with time, so that old pelage was
generally more buffy. The buffy (yellowish-orange) bands on the dorsal
and lateral hairs seemed to oxidize with time, becoming more reddish
(less yellowish) in color. One highly melanistic (but not completely
black) individual from 9 mi E Bismark, Burleigh County, North Dakota
(UMMZ), was noted. This specimen was completing the postjuvenile
molt, which contributed to the darkness of the pelage. In a series of
15 specimens (UMMZ) from Burleigh County, North Dakota, eight
had a mixture of buffy and dark gray color on the abdomen. Other
specimens with buffy ventral parts were noted from Benson County,
North Dakota (two of three in a KU series). Garden County, Nebraska
(one, KU), Larimer County, Colorado (six of seven from Loveland,
BS), and Carbon County, Wyoming (four of 15 in MSB series, with
only slight amounts of buffy on the ventral parts). All other specimens
had ventral parts with pure white hairs.

Individuals in age classes 1 and 2 were typically in juvenile pelage.
Some individuals of age class 1 had started the postjuvenile molt, but
more frequently this molt was initiated in age class 2. Subadults (age
class 3) were nearly all in postjuvenile molt, which was typically com-
pleted by the time the permanent was at occlusal level. However,
some individuals were still in juvenile pelage after the permanent pre-
molars began to show slight wear. Adults appear to have a single molt
per year (it is doubtful that more than a small proportion of individuals
live longer than 12 to 14 months), which generally occurs in the sum-
mer after their birth. Mature adults in obvious molt during the months
of May (four males), June (three males), July (two males, four fe-

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Table 2. — Number of significant differences fP ^ 0.05) in 29 characters for samples of
Perognathus fasciatus and P. flavescens flavescens, based upon SS-STP analysis.
See text for list of characters (length of ear is not included here). Sample codes for
P. fasciatus refer to groups shown in Fig. 1 . See list of specimens examined for localities

of P. flavescens.

Taxa

Perognathus

fasciatus A

Perognathus

fasciatus B

Perognathus
fasciatus C

Perognathus

fasciatus D

Perognathus

flavescens

Perognathus fasciatus A

—

Perognathus fasciatus B

3

—

Perognathus fasciatus C

1

2

—

Perognathus fasciatus D

16

14

15

—

Perognathus flavescens

10

21

21

26

—

males), and August (two females) were noted. These data seem to
suggest that adult males usually molt prior to adult females. However,

larger collections from the spring and autumn months are needed in
order to clearly establish the number and timing of annual molts.

Geographic Variation

Few significant differences in size of external and skeletal characters
between samples 1 through 14, and 17 were found (samples 15 and 16
were too small to include in the SS-STP analysis). Nevertheless, the
mice of samples 1-17 averaged largest in the north and smallest in the
south. No significant differences were found between samples 18 and
19, but these samples were significantly different from all others in
approximately one-half of the characters. Therefore, we decided to
combine the 19 samples into fewer and larger geographic groups. Sam-
ples 1-3 were pooled to form group A (Fig. 1). Samples 4, 5, and 13-
17, plus a few isolated specimens not previously included in any sam-
ple (Fig. 1), formed group B. Group C was composed of samples 7-
12, and group D consisted of samples 18 and 19. The groupings of
samples A, B, and C were somewhat arbitrary, especially the division
of specimens of samples 3 and 4 and of samples 10 and 14.

The standard univariate statistical summaries for the four geographic
groups of P. fasciatus comprise Table 1. A sample of P.f. flavescens
is also included in Table 1 for comparative purposes. Significant dif-
ferences between groups A-D and P. flavescens are summarized in
Table 2. Note that there were few significant differences among groups
A, B, and C, but that they differ significantly in 14 to 16 characters
from group D. All of the samples of P. fasciatus showed high numbers
of significant differences with P. flavescens, although group A (with
the smallest-sized individuals of P. fasciatus) exhibited fewer signifi-
cant differences than the others.

An analysis of correlation among characters showed that all but two

1979

Williams and Genoways — Perognathus Systematics

85

Table 3. — Factor matrix from correlation among 24 cranial and dental characters for 18
samples of Perognathus fasciatus. See text for definitions of abbreviations for traits.

Principal components

Traits

i

II

III

IV

V

ONL

0.705

-0.438

0.138

-0.357

0.017

lOB

0.633

-0.528

0.040

0.222

-0.076

MXTL

0.526

0.390

0.587

-0.041

0.257

WMXT

0.650

0.172

0.064

0.018

-0.390

BL

0.687

-0.496

-0.179

0.161

0.220

BW

0.827

-0.459

0.081

0.021

0.204

IPL

0.174

-0.129

0.800

0.304

0.076

IPW

-0.619

-0.035

0.504

0.317

-0.421

NL

0.430

-0.416

0.392

-0.263

-0.361

NW

0.105

-0.596

0.392

-0.110

0.288

RW

0.193

-0.021

0.565

-0.506

-0.150

LID

-0.619

0.388

0.361

0.273

-0.169

MNTL

0.728

0.454

0.137

0.078

0.247

P4L

0.793

-0.042

-0.282

0.228

-0.102

P4W

0.748

0.259

-0.507

0.174

-0.141

MjL

0.748

0.181

0.404

-0.005

0.155

MjW

0.409

0.594

0.220

0.402

0.277

M3L

0.339

0.385

-0.024

-0.570

-0.263

M3W

0.550

0.505

-0.263

-0.331

0.338

PW

0.786

0.184

-0.202

0.248

-0.388

M^W

0.659

0.436

0.293

0.164

-0.265

M^W

0.685

0.540

0.007

-0.200

-0.157

LAP

0.553

-0.465

-0.131

0.062

-0.436

BE

0.783

-0.371

-0.053

0.220

0.153

% variation

37.85

15.58

11.84

6.99

6.65

of the cranial and dental traits were positively associated with occip-
itonasal length, which can be taken as a character representing size.
Both width of interparietals and least interbullar distance exhibited
significant negative correlations with occipionasal length (r = -0.52
and -0.57, respectively), indicating that as skull length increased, the
interbullar region of the skull became proportionately narrower. There
were several sets of highly correlated characters in the matrix of cor-
relation coefficients, suggesting that the pattern of intersample varia-
tion could be simplified by eliminating redundancy in the character
matrix (we purposefully included some redundant characters in the
study so that alternate characters would be available for various multi-
variate analyses, particularly for future identification of fossil materi-
al).

A principal components analysis of the matrix of correlation among
characters showed that five factors accounted for nearly 80% of the
intersample variation (Table 3). Sixteen factors were needed to ac-

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VOL. 48

I

Fig. 2. — Three-dimensional projection of factor scores of 18 samples of Perognathus
fasciatus on the first three principal components, extracted from a matrix of correlation
among 24 cranial and dental characters. See Fig. 1 for key to samples.

count for all of the variation. Principal Component I was a size factor,
with all traits except width of interparietals (IPW) and least interbullar
distance (LID) exhibiting positive coefficients. Among the characters
with positive coefficients, only length of interparietal (IPL), width of
nasals (NW), and width of rostrum (RW) showed low character load-
ings.

Principal Component II can be characterized as a skull and dental
width factor. It exhibited a more complex pattern of character loadings
than did Principal Component 1. The highest positive coefficients were
dental dimensions, especially molar widths (Table 3). The greatest neg-
ative coefficients were width of nasals (NW) and least interorbital
breadth (lOB). Bullar dimensions (BL, BW) and length of articular
process also had relatively high negative coefficients.

1979

Williams and Genoways — Perognathus Systematics

87

■ 1

\ L— — — 7

I— — 2

L_ — — _ 3

“I J — — — 9

y L — ___ — ______ — _ 10

L-— — — — 12

— — _ 3

— __ 4

|— 10

_____ 14

— 6

L — _______ 13

L_ — _____ 11

— - L - — — 17

L_ — — _ 5

— — 18

”“1 — — - 19

2.0 1.5 10 0.5 0

Fig. 3. — Phenogram, computed from the taxonomic distance matrix and clustered by
UPGMA, for 18 samples of Perognathus fasciatus. See Fig. 1 for key to samples. The
cofficient of cophenetic correlation was 0.75.

Characters with the highest positive coefficients for Principal Com-
ponent III were length of interparietals (IPL), length of maxillary
toothrow (MXTL), and width of rostrum (RW). Width of P4 had a high
negative coefficient.

By plotting the factor scores of the first three principal components
for each of the samples, 65% of the intersample variation (Table 3) can
be summarized in a three-dimensional graph (Fig. 2). Samples are gen-
erally arranged on Principal Component I by size, with the largest mice
(samples 18 and 19) being located on the right in Fig. 2. Basically,
samples are arrayed on Principal Component II by the degree of con-
striction of their skulls and by the width of their molars. Samples with
relatively wide rostra (NW, RW, lOB), wide interbullar regions (LID,
IPW), and narrow molars are positioned on the positive pole of Prin-
cipal Component II. Conversely, those with narrower rostra, more

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VOL. 48

constricted interbullar regions, and wider molars are on the negative
pole. Samples with the shortest lengths of interparietals and shortest
toothrows have the highest scores on Principal Component III. Note
in Fig. 2 that samples 18 and 19 (sample A) are distinct from the others
in size (PC I), and that sample 17 is intermediate to samples 18 and 19
and the others. Some of the more northern samples, such as 6, 10, and
11 are larger sized (PC I) than the more southern samples, such as 1
and 2, but there is no logical geographic clustering of samples on the
first three principal components (Fig. 2).

The results of the cluster analysis of the distance matrix did not
contribute much to an understanding of the geographic relationships
of the P. fasciatus samples. Samples 18 and 19 were more closely
linked to each other than were any other pair of samples (d 0.68,
Fig. 3). The smallest mice tended to be placed at the top of the phe-
nogram, and size generally increased in descending order in the phe-
nogram (Fig. 3). The phenogram weakly summarizes the intersample
relationships (the coefficient of cophenetic correlation = 0.75), and we
fear that small, and not necessarily significant differences in the sam-
ples are magnified by the phenogram.

The similarity analysis was equally poor at delineating geographi-
cally related groups. Samples 18 and 19 were very similar {Q - r =
0.83), and were linked with a cluster consisting of samples 6, 9, 10,

13, 16, and 17. Samples 16 and 17 were more similar to samples 18 and
19 (linkage with 18 and 19 at the Q - r = 0.17 level) than the others,
but the similarities were not close. Overall, the relationships as shown
in the similarity phenogram were the same as those shown by the I
principal components analysis of the matrix of correlation among char- !
acters (Fig. 2). |

The best summaries of the patterns of geographic variation in P. '
fasciatus were achieved by discriminant functiou/canonical analyses. ;
In the first analysis, individuals (with complete data for cranial and j
dental traits) were submitted in 19 separate groups. Samples 18 and 19 |

showed considerable misclassification of individuals between them, !
but none with other samples (indicating their morphological distinct-
ness from the others). Among the remaining samples, misclassification j
ranged from 13% to 47% within samples. All P. flavescens were cor-
rectly placed in their group, but two P. fasciatus were phenetically
more similar (closest in to the mean position on the discriminant
functions) to P. flavescens. When these data were summarized by
canonical analysis, and the positions of all of the individuals were |
plotted on the first two canonical axes, three discrete clusters were
formed. One contained only individuals of samples 18 and 19, one
contained all other individuals of P. fasciatus except for two, and the j
third contained the individuals of P. flavescens, along with the two I

1979

Williams and Genoways — Perognathus Systematics

89

misclassified P. fasciatus. There were no obvious intracluster patterns
that would suggest that samples or groups of samples could be logically
combined into geographic subgroups. More individuals of the northern
samples of P. fasciatus tended to fall closer to samples 18 and 19 than
the others, and more individuals of the southern samples of P. fascia-
tus were positioned closer to the P. flavescens cluster, but these were
trends, not definitive patterns.

Based upon these results, we performed a second discriminant func-
tion/canonical analysis. A few more specimens of P. fasciatus were
included, along with a larger sample of P. flavescens. The individuals
of P. fasciatus were submitted as two groups. Samples 18 and 19
comprised group I, and all other P. fasciatus comprised group IT The
results were nearly identical to the earlier analysis (see above). One
individual (out of 65) from P. fasciatus group I was placed with group
IT This individual was a juvenile (age class 1) that had accidentally
been included in the analysis. Its misclassification was logical, as in-
dividuals of group I were larger than those of group II. One specimen
of group II (out of 112) was placed in group I. This individual, from
Loveland, Larimer Co., Colorado (BS), was a young adult with larger
bullae than normal, but it did not otherwise appear to differ from the
remainder of the series from Loveland. Three individuals of group II
were classified as P. flavescens, and one specimen of P. flavescens
(out of 85) was identified as a group I P. fasciatus. These interspecific
misclassifications are discussed below.

The results of the second discriminant function analysis are por-
trayed in the plot of the scores of individuals on the first two canonical
axes (Fig. 4). Note that two of the P. fasciatus, which were positioned
closest to P. flavescens, were generally intermediate between the clus-
ters of the two species. One of these individuals is from Tilyous Ranch,
27 mi above [SE] mouth Yellowstone River, Richland Co., Montana
(BS 192,208), and the other is from 2 mi S, 6.5 mi W Buffalo, 5,620 ft,
Johnson Co., Wyoming (KU 27,966). Both specimens are from outside
the known range of P. flavescens, and it seems unlikely that they
represent hybrids between P. fasciatus and P. flavescens. The skins
(external characters were not included in this analysis) are not ob-
viously different from other P. fasciatus. The third specimen, posi-
tioned near the center of the P. flavescens cluster, is from Minichaduze
River, Todd Co., South Dakota (BS 17,255/24,186). This locality is in
an area of sympatry between P. fasciatus and P. flavescens. Most
likely, this specimen represents a skin/skull mismatch. The cranial and
dental characters are clearly those of P. flavescens (Fig. 4). When the
specimen was examined at the National Museum of Natural History
(prior to this analysis) it was noted that the individual was in molt from
juvenile to adult pelage, but no notations were made that would suggest

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Annals of Carnegie Museum

VOL. 48

Fig. 4. — Two-dimensional projection of 177 individuals of Perognathus fasciatus and 85
individuals of P. flavescens on the first two canonical axes, derived from the linear
classification functions. The first canonical variable is the linear combination of variables
entered that best discriminates among the groups; the second is the next best linear
combination, orthogonal to the first. Solid symbols indicate the positions of the group
means. Circles = P. flavescens; squares = P. fasciatus group I; triangles = P. fasciatus
group II. Character vectors (upper left part of diagram) are shown only for the most
influential characters.

that it otherwise differed from P. fasciatus. The skull was an old adult
(age class 5), so, clearly, the skin and skull are from different individ-
uals. We have not looked among the P. flavescens specimens to see j
if we could locate the complementary mismatched skin and skull.

The one individual of P. flavescens, which was placed with group
I of P. fasciatus, is the one at the extreme bottom center of Fig. 4.
This specimen, from 21 mi E, 3 mi S Carlsbad, Lea Co., New Mexico
(Eastern New Mexico University collection), was larger than typical
for P. flavescens copei, but did not differ otherwise. This locality is
about 350 miles south of the southernmost locality of P. fasciatus, and ,1
there is little chance that it represents a hybrid between the two
species.

1979

Williams and Genoways — Perognathus Systematics

91

A few specimens that had previously been classified as P.flavescens
by other workers seemed to us, on the basis of their skins, to represent
P. fasciatus. The skulls of those that we were able to test by the
discriminant function analysis were clearly classified as P. fasciatus.
These include the following: BS 148,645, from 5,500 ft, Boulder Co.,
Colorado; BS 270,850, from Lindenmeier, Larimer Co., Colorado;
UIMNH 7,567 and 7,662 from 6 mi N, 1 mi W Colorado Springs, El
Paso Co., Colorado; and CAS 14,021, 14,022, 14,031, and 14,032 from
Air Force Academy, 10 mi N Colorado Springs, El Paso Co., Colo-
rado. We made no attempt to confirm the identities of all P. flavescens
from along the Rocky Mountain front in Colorado, but we did examine
most of this material. The list of specimens examined for P.f. flaves-
cens includes only specimens measured for the univariate analysis, but
we examined most of the other material in the collections of the insti-
tutions listed in this report and in Williams (1978^).

We did not quantify color variation because the differences in color
were relatively slight, and color varied individually with age and sea-
son as well as geographically. The predominant trends in color varia-
tion can be summarized verbally, however. When adult specimens in
fresh pelage were compared, the darkest colored individuals were
found in eastern and northern North Dakota. Specimens from northern
Montana and Saskatchewan had similar numbers of black-tipped hairs,
but the buffy color was somewhat paler. Specimens were progressively
paler in samples arranged from northeast to southwest. The change in
color was gradual (clinal), but was more pronounced from northeastern
to southwestern North Dakota. Specimens from throughout most of
the area encompassed by groups A and B (Fig. 1) were essentially the
same in color. Only those from the Shirley Basin, Wyoming (sample
15) were noticeably lighter than the others. Specimens of samples 18
and 19 (Group D, Fig. 1) were considerably paler than all others except
those in sample 16, with a more yellow (less orange) buffy color, and
with less of the “olive” tone that is characteristic of P. fasciatus
elsewhere.

Discussion and Conclusions

Geographic variation in size, proportions, and color in P. fasciatus
is relatively slight, especially compared to its close relative, P. fla-
vescens (Williams, 19781?). The major trend in variation noted was that
of size. As size increases, there is a concordant constriction of the
interbullar region of the skull, and an increase in relative length of the
tail. These trends are not nearly as pronounced in P. fasciatus as in
P.flavescens (Williams, 1978/?). Size varies clinally in the Great Plains,
with larger individuals being found in the colder, northern latitudes
and smaller individuals coming from the southern parts of the geo-

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Annals of Carnegie Museum

VOL. 48

A

B

20 mm

Fig. 5. — Dorsal views of skulls of Perognathus fasciatus fasciatus and P.f. callistus.
A = P. f. fasciatus, UMMZ 75,354 male from 12 mi W Mandan, Morton Co., North
Dakota; B = P.f. callistus, CM 21,187, male from Shell Creek, 25 mi S Bitter Creek,
Sweetwater Co., Wyoming.

graphic range. Proportional differences are slight in the Great Plains
populations.

The largest mice were found in the Uintah and Great Divide basins,
in the southwestern corner of the geographic range of P. fasciatus
(group D, Fig. 1). The demarkation between this population and the
others is well defined (Figs. 1 and 4). Individuals in this population
have longer tails, proportionately larger bullae, and more constricted
interbullar regions (Fig. 5). These intermountain basins are relatively

1979

Williams and Genow ays— Perognathus Systematics

93

arid (annual precipitation averages about six inches), and the relatively
larger bullae are in accord with the principle of ecogeographic variation
that predicts larger auditory bullae in drier climates.

Color varies according to the ecogeographic principle known as G!o“
ger’s rule. The darkest colored mice are from the areas of highest
rainfall and lowest evaporation in northeastern North Dakota (mean
annual precipitation >20 inches). Mean annual precipitation decreases
progressively to the southeast. The areas with the greatest difference
in available moisture are the ietermountaie basins of Colorado, Utah,
and Wyoming (<10 inches of mean annual precipitation). Specimens
from the Shirley (sample 16), Great Divide (sample 18), Bridger (Green
River) and Uintah basins (sample 19) are much lighter colored than
those from the Great Plains. The individuals from the Shirley Basin
are structurally most similar to specimens from the Great Plains.

Our data, together with those presented by Williams (1978^), show
that there is no evidence that P. fasciatus and P. flavescens hybridize
in areas of sympatry. The two species are distinct in color, structure,
and karyotypes (Williams, 1978€i). Where the geographic ranges of P.
fasciatus and F. flavescens meet in the Uintah Basin, individuals of
P. flavescens are larger than P. fasciatus, whereas in the area of sym-
patry in the Great Plains, individuals of P. fasciatus are larger than P.
flavescens. Structural convergence in size, bullar inflation, and inter-
bullar constriction has occurred in the two species in the Uintah Basin
area, but in sympatry (Great Plains populations), the two species are
divergent in size and proportions (see Williams, 19781? for additional
measurements of P. flavescens). Williams (19781?) did not find the two
species together at any locality in the Uintah Basin, although they
were captured on opposite sides of the White River, and at other lo-
calities within 5 km of each other north of the White River and west
of the Green River. He observed no apparent differences in the habi-
tats of the two species there. These data suggest that competitive ex-
clusion could be the major interspecific interaction in the Uintah Basin.
Alternately, one or both species may be recent arrivals, and interspe-
cific interactions may be just beginning (Williams, 19781?).

In southeastern Wyoming, Maxwell and Brown (1968) caught P.
fasciatus only in a Bouteloua-Stipa community with sandy loam soils
and relatively dense vegetational cover. They found P. flavescens to
be more euryecious, but most abundant on loamy sand and sandy loam
soils in sage-grass communities. They did not find the two species
together at any of their trapping stations. Based upon our experiences
(Williams, 19781?; Geooways and Jones, 1972, and unpublished), and
those of others (for example, Jones, 1964; Sweek, 1940, Nero, 1958),
P. flavescens is usually limited to relatively open habitats with sand
or sandy loam soils. P. fasciatus is more tolerant of a variety of sub-

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Annals of Carnegie Museum

VOL, 48

Fig. 6. — Map, showing approximate collecting localities and geographical distribution
of subspecies of Perognathus fasciatus. A = P. fasciatus fasciatus; B = P.f. callistus.
Concentric circles indicate type localities. Localities with partially overlapping positions
are indicated by single circles.

strates and densities of vegetational cover, but does not appear to
inhabit sand soils where P. flavescens occurs. Well-marked habitat
differences, together with size and proportional differences in the pop-
ulations suggest a long period of competitive interaction between the
species in the Great Plains.

1979

Williams and Genoway^^-Perognathus Systematics

95

It is apparent that specimens of P. fasdatus are readily separable
into two distinct groups. Individuals from the Great Plains are smaller,
with shorter tails, proportionately smaller bullae, and are darker cob
ored. This population includes the subspecies P. fasdatus fasdatus
Wied, 1839, P.f. infraluteus Thomas, 1892, P.f. Utus Cary, 1911, and
P.f. olivaceogriseus Swenk, 1940. We can see no basis for recognizing
more than a single taxon from this area. Individuals of the other pop-
ulations are larger, with proportionately longer tails, larger bullae, and
paler color, and are found in the Unitah, Bridger, and Great Divide
basins, and contiguous areas (Fig. 6). This group includes P . fasdatus
callistus Osgood, 1900, and some specimens that were previously aS“
signed to P.f litus (but not the holotype of P.f Utus, from the Shirley
Basie, which was placed among the individuals of the P. f. fasdatus
sample in the discriminant function analyses).

Williams (1978«) reported identical appearing karyotypes for indi-
viduals from samples 17, 18, and 19. Until Jones (1953) synooymized
callistus with P. fasdatus, it had been regarded as a distinct species.
The structural differences between P, f. callistus and P. f. fasdatus
are not especially great, and specimens from geographically interme-
diate areas, such as the Shirley Basin (sample 16) and Ft. Steele (sam-
ple 17) are somewhat intermediate structurally (Fig. 2). In our opinion,
the evidence fully supports Jones’ (1953) decision that callistus is a
race of P. fasdatus.

Systematic Accounts
Perogeathus fasdatus fasdatus Wied, 1839

1839. Perognathus fasdatus Wied, Nova Acta Phys.-Med., Acad. Caesr. Leop. -Carol.,
19:369.

1893. Perognathus infraluteus Thomas, Ann. Mag. Nat. Hist,, ser. 6, 11:406, May;

holotype from Loveland, Larimer Co., Colorado.

1900. Perognathus fasdatus infraluteus, Osgood, N. Amer. Fauna, 18:19, 20 Septem-
ber.

1911. Perognathus fasdatus litus Cary, Proc. Biol. Soc. Washington, 24:61, 22 March;

holotype from Sun, Natrona Co., Wyoming.

1940. Perognathus flavescens olivaceogriseus Sweek, Missouri Valley Fauna, 3:6, 5
June; holotype from Chadroo [Little Bordeaux Creek, sec. 14, T38N, R48W, 3
mi E Chadroe], Dawes Co., Nebraska.

1953. Perognathus fasdatus olivaceogriseus, Jones, Univ. Kansas PubL, Mus. Nat.
Hist., 5:520, 1 August.

Neoiype.— Adult male (age class 4), skin and skull, BS 168,599, from
Buford, Williams Co., North Dakota; obtained on 6 May 1910 by H.
E. Anthony, Both skin and skull in good condition.

Measurements of neotype .—Toidi length, 140; length of tail, 66;
length of hied foot, 18.0; occipitonasal length, 23.35; least interorbital
breadth, 5.30; alveolar length of maxillary toothrow, 3.20; width across
maxillary toothrows, 4.45; length of bulla, 7.70; width across bullae,

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VOL. 48

11.90; length of interparietal, 2.65; width of interparietals, 4.90; length
of nasal, 8.75; width of nasals, 2.15; width of rostrum, 4.00; least
interbullar distance, 4.60; crown length of mandibular toothrow, 2.85;
bullar extension, 0.00.

Distribution.— Vound on light, sandy soils in the northern Great
Plains, in areas generally receiving less than 20 inches of mean annual
precipitation. Known from near Saskatoon, Saskatchewan, southward
to near La Veta, Huerfano Co., Colorado; and from near the North
Dakota-Minnesota border west to southeastern Alberta (Fig. 6).

Diagnosis. — ^See Table 1, groups A, B, and C for measurements.
Size small for the species, but about average for the subgenus Per-
ognathus. Size largest in the northeast and smallest in the south. Tail
relatively short and nonpenicillate. Skull with relatively small bullae,
wide interbullar region, and narrow interorbital region (Fig. 5). Color
dark, with prominent olivaceous tone dorsally; face and lateral line
more buffy than other dorsal parts; ventral surfaces usually pure white,
but occasionally with some buffy and gray hairs. Color darkest in the
northeast and lightest in the southwest.

Comparisons. — Distinguishable from P. flavescens flavescens by its
larger size (Table 1), narrower interorbital region, larger bullae, shorter
and narrower interparietals, and larger molariform teeth; color much
darker than sympatric P. flavescens, and with an olivaceous tone that
is not present in P. flavescens. Size considerably smaller, color more
olivaceous, and bullae much more inflated than P. hispidus. Size no-
ticeably larger, bullae less inflated, and interbullar region significantly
wider than P. flavus. Size smaller, color darker (buffy color near Cin- I
namon-Buff or Clay-color; Ridgway, 1912), bullae less inflated, and
interbullar region relatively wider than P. fasciatus callistus (Table 1). |

Remarks. — The disposition of Maximilian’s holotype is unknown.
We have not found any reference to it in the literature. However, we
presume that Maximilian preserved a specimen, as he included a figure
of the skull and teeth, as well as a colored plate of the animal with his
description. Some of Maximilian’s collection was purchased by the
American Museum of Natural History in 1869 (Pires, 1965), but there
were apparently no specimens of Perognathus fasciatus among those j;
purchased. Merriam (1889) redescribed P. fasciatus from a ‘'duplicate |
type,” obtained from Tilyous Ranch, 27 mi above [SE] mouth Yellow-
stone River, Richland Co., Montana, on 6 October 1887. We did not
designate this specimen as the neotype because specimens were avail- ‘i
able from the type locality, and because the labial cusps of the RP4 are
compressed together, giving the tooth a subacute, triangular shape. !

Maximilian has been listed in the literature under several names. He
referred to himself as Maximilian, or Maximilian, Prince of Wied. The 1

1979

Williams and Genoway^—Perognathus Systematics

97

Century Dictionary (1910) listed Mm as “Neuwied, Maximilian Alex-
ander PMlipp, Prince of.” He was born and resided in Neuwied, Prus-
sia. His title was apparently inherited from an earlier age, as the place,
Wied, no longer existed when Maximilian was born (1782). We can
see no reason to posthumously change his name (title) to Neuwied or
Wied-Neuwied, as was done in the early part of this century.

Specimens examined.— Tot^l was 297, distributed as follows: Manitoba. Aweme, 2
(BS), 6 (ROM). Saskatchewan. 10 mi S Sceptre, 2 (ROM); Tompkins, 2 (BS). Col-
orado. Boulder Co.: 5,500 ft [no speciic locality designated], 1 (BS). Elbert Co.: 1 mi
N Ramah, 1 (UMMZ). El Paso Co.: Air Force Academy, 10 mi N Colorado Springs,
4 (CAS); 6 mi N, 1 mi W Colorado Springs, 2 (UIMNH). Huerfan.o Co. : 4 mi S La Veta,

1 (KU). Jefferson Co.: Green Mountain, 5 mi W Denver, 1 (KU)= Larimer Co.: Box
Eider Canyon, sec. 2, TON, R70W, 1 (UMMZ); 13 mi E Ft. Collins, 1 (KU); Linden-
meier, 1 (BS); Loveland, 10 (BS). Logan Co.: 4 mi W Peetz, 1 (SIUC). Montana.
Bighorn Co.: Sage Creek, Bighorn Basin, 1 (BS); Walborn Ranch, Crow Indian Reser-
vation, 1 (BSC); Walborn Ranch, middle fork Tullock Creek, Crow Indian Reservation,
3 (BSC). Carbon Co.: Clark's Fork River, 2 (BS). Carter Co.: Little Missouri River,

8 mi NE Albion, 1 (BS); 5 mi N, 3.5 mi W Camp Cook, 3,400 ft, 5 (KU); Sand Creek,
6 mi S, 4.5 mi W Camp Cook, 5 (KU); Ekalaka Hills, 4.5 mi E Ekalaka, 1 (MMNH).
Custer Co. : 13 mi E Miles City, 1 (UMMZ). Fallon Co. : 18 mi S Baker, 1 (UM). Garfield
Co.: PiQey Bettes, 1 (BS). Phillips Co.: Frenchman River, 3 (BS). Powder River Co.:

2 mi E Biddle, 1 (UM); Powderville, 2 (BS). Richland Co.: Tilyous Ranch, 27 mi above
mouth Yellowstone River, 6 (BS). Roosevelt Co.: 9 mi SE Bainviile, 4 (UMMZ); 6=5 mi
N Culbertson, 2 (UIMNH); Johnson Lake, 2 (BS); 2.25 mi W McCabe, 1 (UIMNH).
Rosebud Co.: Big Porcupine Creek, 1 (BS). Sheridan Co.: 3 mi S Medicine Lake, 1,880
ft, 1 (KU); 5 mi SW Plentywood, 1 (UM). Valley Co.: Big Porcupine Creek, 1 (BS).
Yellowstone Co. : Lake Basie, 2 (BS). Nebraska. Banner Co. : 10 mi S, 2.5 mi E Gering,

3 (VMKSC). Cherry Co.: Ft. Niobrara Game Reserve, 1 (UNSM); 10 mi E Gordon, 1
(FMNH); 12 mi ESE Gordon, 2 (FMNH); Sparks, 1 (UMMZ). Dawes Co.: 10 mi S
Chadron, 3 (UMMZ); 1 mi SW Chadron, 1 (UNSM). Garden Co.: 2 mi S Oshkosh, 1
(KU). Kimball Co.: Pole Creek, 490 mi from Ft. Riley [= Lodgepole Creek], 1 (BS).
Sheridan Co.: Mirage Township, 1 (UMMZ). Sioux Co.: 1.5 mi E, 16 mi S Agate, 4,950
ft, 1 (KU); 6 mi W Crawford, 1 (KU); 5.5 mi W Crawford, 1 (UNSM); 8 mi W Ft.
Robinson, 1 (UNSM); 3 mi N Glenn, 2 (UNSM); 3 mi N, 1 mi E Glenn, 3 (UNSM);
Glenn, 1 (UNSM). North Dakota. Barnes Co.: Glen Ullin, 2 (UMMZ). Benson Co.:
2 mi W Fort Totten, 1,400 ft, 3 (KU). Billings Co.: 5.5 mi S, 2 mi W Medora, 1 (KU);
1 mi S, 1 mi W Medora, 21 (KU). Bottineau Co.: Bottineau, 1 (FMNH). Burleigh Co.:

9 mi E Bismark, 8 (UMMZ). Dickey Co. : Oakes, 1 (BS), 1 (FMNH). Divide Co. : Crosby,
1 (BS). Golden Valley Co.: sec. 9, T14iN, R105W, 1 (UNDAK). Grant Co.: Leith, i
(UNDAK). Kidder Co.: Dawson, 1 (BS); 6 mi W Steele, 6 (UMMZ). Logan Co.: Na-
poleon, 2 (BS). McHenry Co.: near Upham, 2 (BS). McKenzie Co.: 40 mi N Medora,
1 (BS); sec. 36, T154N, R97W, 1 (UNDAK). McLean Co.: sec. 3, T144N, R82W, 1
(UNDAK). Morton Co.: 11 mi S Mandan, 1 (UMMZ); 12 mi W Mandan, 4 (UMMZ);
sec. 29, T133N, R82W, 1 (UNDAK); sec. 6, T137N, R82W, 1 (UNDAK); sec. 16,
T138N, R87W, 1 (UNDAK). Oliver Co.: Ft. Clark, 10 (BS). Pembina Co.: Week's
Farm, sec. 36, T160N, R56W, 1 (MSB). Sargent Co.: 1 mi S, 7.2 mi E Oakes, 1,200 ft,
1 (KU), Sioux Co.: Cannon Ball, 7 (BS). Slope Co.: Badlands, 1 (UNDAK); sec. 17-
18, T135N, RIOIW, 2 (UNDAK); sec. 12, T136N, RIOIW, 3 (UNDAK); sec. 11, Ti36N,
R102W, 4 (UNDAK); sec. 30, T136N, R104W, 1 (UNDAK). Stark Co.: 1 mi S Dick-
inson, 1 (UMMZ); 9 mi W Dickinson, 7 (UMMZ); 2 mi W Taylor, 7 (UMMZ). Stutsman

98

Annals of Carnegie Museum

VOL. 48

Co.: 7 mi N Jamestown, 1 (UMMZ); 14 mi W Jamestown, 1 (UMMZ). Walsh Co.: 3 mi
W Park River, 1 (KU). Wells Co.: Bowden, 2 (BS), 2 (CM). Williams Co.: Buford, 10
(BS). South Dakota. Beadle Co.: 4 mi S, 5 mi E Hitchcock, 1 (UIMNH). Custer Co.:
Quinn’s Draw, Cheyenne River, 3 (BS); Campbell’s Ranch, Elk Mountain, 4,800 ft, 1
(BSC). Jackson Co.: 7 mi SW Kadoka, 2 (MMNH). Meade Co.: Smithville, 1 (BS).
Shannon Co.: 10 mi S Imlay, 2 (BS); Pine Ridge, 3 (BS), 1 (FMNH). Todd Co.: Mini-
chaduza River, 2 (BS); 15 mi W Mission, 1 (MSB); Rosebud Agency, 1 (BS). Tripp Co.:
1 mi SE Colome, 3 (UMMZ), Walworth Co.: Molstad Lake Park, 1 (KU); Swan Creek,
13 mi S Selby, 1 (KU). Washabaugh Co.: White River Flood Plain, 7 mi S Kadoka, 1
(UMMZ). No County Designated: Black Hills, 1 (BS). Wyoming. Carbon Co.: Ft.
Steele, 1 (BS); 1 mi E Ft. Steele, 15 (MSB); 8 mi SE Lost Soldier, 2 (BS). Converse
Co.: Van Tassel Creek, 1 (CM). Crook Co.: Sundance, 1 (BS). Freemont Co.: 40 mi E
Dubois, 1 (UMMZ); Granite Mountains, 1 (UMMZ). Hot Springs Co.: Kirby Creek,
5,000 ft, 1 (BS). Johnson Co.: 2 mi S, 6.5 mi W Buffalo, 5,620 ft, 3 (KU). Natrona Co.:
Casper, 1 (BS); 5 mi W Independence Rock, 6,000 ft, 4 (KU); 16 mi S, 1 1 mi W Waltman,
6,950 ft, 1 (KU). Sheridan Co. : Arvada, 3 (BS); 5 mi NE Clearmont, 3,900 ft, 2 (KU).
Weston Co.: Newcastle, 1 (BS).

Additional records .—Ai^bert a. Foremost (Jones, 1953); Manyberries (Jones, 1953);
Medicine Hat (Jones, 1953). Manitoba. Junction Antler and Souris rivers (Jones, 1953);
Oak Lake (Jones, 1953); Treesbank (Anderson, 1947). Saskatchewan. Baildon (Nero,
1965); Beaver Creek, 9 mi S Saskatoon (Nero, 1958); 10 mi SE Beechy (Nero, 1965);
Big Muddy Valley, near Bengaugh (Nero, 1965); Ceylon (Nero, 1964); Corval (Nero,
1958); Cottonwood Creek, near Regina (Nero, 1965); 6 mi SE Elbow (Nero, 1958); 4 mi
SW Elbow (Nero, 1958); Estevan (Nero, 1965); Eyebrow Lake (Nero, 1958); 2 mi NE
Grandora (Nero, 1958); Hatfield (Nero, 1958); Imperial (Nero, 1965); Last Mountain
Lake, West of Govan (Nero, 1965); 6 mi S Meyronne (Nero, 1965); 6 mi SE Meyronne
(Nero, 1965); Moon Lake, 5 mi SW Saskatoon (Nero, 1958); Mortlach (Nero, 1965);
North Portal (Nero, 1965); Old Wives (Nero, 1958); Piapot (Nero, 1965); Quantock
(Nero, 1958); Radville, (Nero, 1965); Regina Beach (Nero, 1958); Saskatchewan Landing
(Nero, 1965); Skull Creek (Nero, 1965); Strawberry Lakes (Nero, 1965); 4 mi E Swanson
(Nero, 1958); Weyburn (Nero, 1958). Colorado. Custer Co.: 12.4 mi (by road) NE
Silver Cliff, (Armstrong, 1972); 11.2 mi (by road) NE Silver Cliff, 8,200 ft (Armstrong,
1972); 10.4 mi (by road) NE Silver Cliff, 8,025 ft (Armstrong, 1972); 9.6 mi (by road)
NE Silver Cliff, 7,989 ft (Armstrong, 1972); 9.2 mi (by road) NE Silver Cliff, 8,000 ft
(Armstrong, 1972); 8.4 mi (by road) NE Silver Cliff, 8,200 ft (Armstrong, 1972); 6.8 mi
(by road) NE Silver Cliff, 8,200 ft (Armstrong, 1972); 6.4 mi (by road) NE Silver Cliff,
8,200 ft (Armstrong, 1972); 5.6 mi (by road) NE Silver Cliff, 8,050 ft (Armstrong, 1972);
4.4 mi (by road) NE Silver Cliff, 7,950 ft (Armstrong, 1972); 3.6 mi (by road) NE Silver
Cliff, 7,975 ft (Armstrong, 1972); 2,4 mi (by road) NE Silver Cliff, 7,970 ft (Armstrong,
1972); 1.2 mi (by road) NE Silver Cliff, 7,900 ft (Armstrong, 1972). Freemont Co.: 23.7
mi (by road) NE Silver Cliff, 6,200 ft (Armstrong, 1972); 14.8 mi (by road) NE Silver
Cliff (Armstrong, 1972); 13.6 mi (by road) NE Silver Cliff, 7,980 ft (Armstrong, 1972).
Weld Co.: 2.5 mi N, 12.5 mi E Ft. Collins (Armstrong, 1972). Montana. Custer Co.:
Calf Creek (Jones, 1953); Wolf s Creek (Jones, 1953). Nebraska, Cherry Co.: Valentine
(Jones, 1953). Dawes Co.: Chadron State Park (Jones, 1953); Little Bordeaux Creek,
3 mi E Chadron (Jones, 1953). Sioux Co.: Monroe Canyon (Jones, 1953). South Da-
kota. Custer Co.: Cheyenne River (Jones, 1953); Wind Cave Canyon, Wind Cave
National Park, 4,100 ft (Turner, 1974). Shannon Co.: Corral Draw (Jones, 1953). Wy-
oming. Albany Co.: Tie Siding Picnic Grounds, 8,595 ft (Long, 1965). Campbell Co.:
1.25 mi N, 0.5 mi E Rocky Point, 3,850 ft (Jones, 1953). Goshen Co.: Muskrat Canyon
(Long, 1965). Johnson Co.: 1 mi WSW Kaycee, 4,700 ft (Jones, 1953). Laramie Co.:
15 mi ESE Cheyenne (Long, 1965). Natrona Co.: 1 mi NE Casper, 5,150 ft (Jones,
1953). Platte Co.: 2.5 mi S Chugwater (Jones, 1953). Weston Co.: I'i mi SW Newcastle,
4,500 ft (Jones, 1953).

1979

Williams and Genow ays— Perognathus Systematics

99

Perognathus fasclatus callistus Osgood, 1900

1900. Perognathus callistus Osgood, N. Amer. Fauna, 18:28, 20 September.

1953. Perognathus fasciatus callistus, Jones, Univ. Kansas Publ,, Mus. Nat. Hist.,
5:524, 1 August.

Holotype .—Adult male (age class 4), skin and skull, BS 88,245, from
Kinney Ranch [about 22 mi S Bitter Creek], Sweetwater Co., Wyo-
ming; obtained on 14 May 1897 by J. A. Coring. Both skin and skull
in good condition.

Measurements of holotype.—Toi^ length, 135; length of tail, 63;
length of hied foot, 18.0; occipitonasal length, 22.65; least interorbital
breadth, 5.05; alveolar length of maxillary toothrow, 3.10; width across
maxillary toothrows, 4.45; length of bulla, 8.55; width across bullae,
13.10; length of interparietal, 2.70; width of interparietals, 4.65; length
of nasal, 8.10; width of nasals, 2.35; width of rostrum, 3.55; least
interbullar distance, 4.50; crown length of mandibular toothrow, 2.90;
bullar extension, 0.15.

Distribution.— Distil and steppe grassland associations (Upper So-
noran) in the Uintah, Bridger, and Great Divide basins, and contiguous
areas of Colorado, Utah, and Wyoming (Fig. 6).

Diagnosis. See Table 1, group D for measurements. Size large for
the species, and near average for the subgenus Perognathus. Tail rel-
atively longer than other populations of P. fasciatus, but about average
for the subgenus; tail iionpeeicillate. Skull with relatively large bullae,
narrow interbullar region, and wide interorbital region (Fig. 5). Color
light, without prominent olivaceous tone dorsally; ventral surfaces
pure white.

-Distinguishable from P. flavescens caryi of the Uin-
tah Basin by smaller size, relatively shorter tail, and light olive-yellow,
rather than yellowish-orange lateral line; interparietal shorter, rostrum
narrower, and interbullar region wider than P. f. caryi (see Williams,
1978^, for measurements of P. f. caryi). Differs from P. parvus in
smaller size (occipitonasal length in P. parvus averages greater than
25.5 mm), more buffy (less gray) color, and much shorter, nonpeni-
cillate tail. Size larger, tail longer, color lighter (buffy color near
Cream-Buff or Chamois; Ridgway, 1912) than P.f. fasciatus, with less
of an olivaceous tone; interorbital region wider, bullae more inflated,
and interbullar region more constricted than P. f fasciatus (Fig. 5).

Remarks .Specimens previously assigned to P. fasciatus litus from
Sweetwater Co., Wyoming (Jones, 1953; Long, 1965; and Williams,
1978^) are referrable to P. f. callistus. Specimens assigned to P. f.
litus by Jones (1953), from Carbon, Freemont, and Natrona counties,
Wyoming, including the holotype of P. f. litus, are referrable to P. f
fasciatus.

100

Annals of Carnegie Museum

VOL. 48

Specimens examined. — Total was 82, distributed as follows: Colorado. Moffat Co.:
N Bank Yampa River, 5 mi NW Cross Mountain, 1 (CM). Rio Blanco Co.: 16 mi W
Meeker, 3 mi up Scenery Gulch [N of White River], 1 (CM). Utah. Dagget Co.:
Bridgeport, 1 (UU); 0.5 mi SW Clay Basin Camp, 6,300 ft, 2 (UU). Uintah Co.: 4.6 mi
N Bonanza, 4 (MSB); Bonanza, 1 (UU); 1.5 mi E Bonanza, 1 (MSB); Dead Man Bench,
West Rim (opposite Leota Flats) [W side Green River], 2 (CM); E side Green River, 3
mi S Jensen, 1 (CM). Wyoming. Carbon Co.: Loco Creek, 25 mi W Saratoga, 7,500 ft,

1 (BSC); Sane Creek, 17 mi W Saratoga, 1 (BSC). Sweetwater Co.: 18 mi S Bitter
Creek, 6,800 ft, 3 (KU); 30 mi S Bitter Creek, 2 (KU); 33 mi S Bitter Creek, 6,900 ft,

2 (KU); Green River [city], 2 (BS); 350 river miles N Green River, Utah [opposite mouth
Black’s Fork, 5,930 ft], 1 (UU); Kinney Ranch, the holotype + 3 (BS); Kinney Ranch,
21 mi S Bitter Creek, 6,800 ft, 6 (KU), 2 (MVZ); Kinney Ranch, sec. 8, T15N, R98W,
23 mi SW Bitter Creek, 1 (MVZ); the Green River, 4 mi ENE Linwood, 2 (KU); 27 mi
N, 37 mi E Rock Springs, 6,700 ft, 5 (KU); 32 mi S, 22 mi E Rock Springs, 7,025 ft, 2
(KU); Shell Creek, 25 mi S Bitter Creek, 5 (CM); 27 N Table Rock, 1 (UMMZ); 25.4
mi N Table Rock, 27 (MSB); 2.5 mi N Wamsutter, 1 (KU).

Additional records.— Colorado. Moffat Co.: [little] Snake River, 7 mi above Bear
[Yampa] River (Armstrong, 1972); Two Bar Spring [20 mi NW Junction Little Snake
and Yampa rivers] (Armstrong, 1972). Wyoming. Sweetwater Co.: 25 mi N, 38 mi E
Rock Springs, 6,700 ft (Jones, 1953).

Other Specimens Examined

Perognathus flavescens flavescens.—ToidX was 110, distributed as follows: Colo-
rado. Adams Co.: Barr, 1 (UCM). Cheyenne Co.: 1 mi N, 8 mi E Kit Carson, 1 (KU).
Kit Carson Co.: Tuttle, 1 (BS). Pueblo Co.: Pueblo, 2 (BS). Washington Co.: 8 mi W
Akron, 6 (UMMZ); Eastern Colorado Experiment Range, 2 (KU); 10 mi S, 7 mi E Otis
Wash, 1 (KU). Weld Co.: Greeley, 2 (BS); Roggen, 1 (FMNH), Nebraska. Antelope
Co.: Clearwater, 1 (UMMZ); Neligh, 1 (BS), 1 (UNSM). Banner Co.: 10 mi S, 2.5 mi
E Gering, 3 (VMKSC). Cherry Co.: Big Alkali Lake, 1 (UNSM); Hackberry Lake, 1
(BSC), 2 (KU), 18 (UMMZ); Kennedy, 4 (MVZ), 1 (UMMZ); 2 mi E Kennedy, 3 (KU);
4 mi E Kennedy, 2 (KU); 4 mi S Kennedy, 1 (UNSM); 18 mi NW Kennedy, 1 (UNSM);
11.5 mi S, 0.5 mi W Nenzel, 3,000 ft, 1 (VMKSC); Niobrara River, 10 mi S Cody, 1
(UNSM); 2 mi E Valentine, 1 (KU); 4 mi E Valentine, 1 (KU). Custer Co.: W mi S, 2
mi W Broken Bow, 2 (VMKSC). Garden Co.: Headquarters, Crescent Lake National
Wildlife Refuge, sec. 29, T21N, R44W, 3 (UNSM); 0.75 mi E Headquarters, Crescent
Lake National Wildlife Refuge, sec. 28, T2iN, R44W, 1 (UNSM); 3 mi S Headquarters,
Crescent Lake National Wildlife Refuge, 4 (UNSM); 5 mi S Headquarters, Crescent
Lake National Wildlife Refuge, 1 (UNSM). Hooker Co.: Kelso, 7 (UMMZ). Kearney
Co.: 10 mi N, 1 mi E Axtell, 2 (VMKSC); 10 mi E Axtell, 4 (VMKSC); Doby Town, 5
mi S, 3 mi E Kearney, 8 (VMKSC). Keith Co.: N side Kingsley Reservoir, 1 (UNSM).
Lincoln Co.: 1 mi N Brady, 1 (UNSM); Brady, 1 (UNSM); 2.5 mi N, 4.5 mi E North
Platte, 1 (VMKSC). Rock Co.: Perch, 1 (FMNH). Scotts Bluff Co.: 6 mi N Mitchell,
1 (UNSM). Sheridan Co.: 14 mi W Lakeside, 1 (MVZ). Thomas Co.: Halsey National
Forest, 1 (VMKSC). South Dakota. Bennett Co.: Batesland, 1 (FMNH). Potter Co.:
Whitlock’s Crossing, 8 (KU).

Acknowledgments

We would like to thank the following curators for allowing us to examine specimens
in their care: D. M. Armstrong, L. C. Binford, E. C. Birney, J. P. Farney, J. S. Findley,
R. B. Finley, Jr., P. W. Freeman, H. L, Gunderson, R. S. Hoffmann, D. F. Hoffmeister,
E. T. Hooper, R. L. Hutto, C. Jones, J. L. Patton, R. L. Peterson, R. W. Seabloom,
H. J. Stains, and D. E. Wilson. Nancy Perkins prepared the figures, and Suzanne Braun
aided us in editing drafts of this report.

1979

Williams and Genoways — Perognathus Systematics

101

Literature Cited

Anderson, R. M. 1947. Catalog of Canadian Recent mammals. Bull. Nat. Mus. Can-
ada, 102:1-238.

Armstrong, D. M. 1972. Distribution of mammals in Colorado. Univ, Kansas Mus.
Nat. Hist., Monog., 3:1-415.

Baird, S. F. 1858. Mammals, in Reports of explorations and surveys for a railroad

route from the Mississippi River to the Pacific Ocean. 8(l):xxi-xlviii + 1-757 + 43

pi.

Cary, M. 1911. A new pocket mouse from Wyoming. Proc. Biol. Soc. Washington,
24:61.

Choate, J. R., and H. H. Genoways. 1975. Collections of Recent mammals in North

America. J. Mamm., 56:452-502.

Codes, E. 1875. A critical review of the North American Saccomyidae. Proc. Acad.
Nat. Sci. Philadelphia, 1875:272-327.

Dixon, W. J., and M. B. Brown (eds.). 1977. BMDP-77: Biomedical computer pro-
grams, P-series. Univ. California Press, Berkeley, xiii + 880 pp.

Gabriel, K. R. 1964. A procedure for testing the homogeneity of all sets of means in
analysis of variance. Biometrics, 20:459-477.

Genoways, H. H., and J. K. Jones, Jr. 1972. Mammals from southwestern North
Dakota. Occas. Papers Mus., Texas Tech Univ., 6:1-36.

Jones, J. K., Jr. 1953. Geographic distribution of the pocket mouse, Perognathus
fasciatus. Univ. Kansas Publ., Mus. Nat. Hist., 5:515-526.

. 1964. Distribution and taxonomy of mammals of Nebraska. Univ. Kansas

Publ., Mus. Nat. Hist., 16:1-356.

Long, C. A. 1965. The mammals of Wyoming. Univ, Kansas Publ., Mus. Nat. Hist.,
14:493-758.

Maxwell, M. H., and L. N. Brown. 1968. Ecological distribution of rodents on the
High Plains of eastern Wyoming. Southwestern Nat., 13:143-158.

Merriam, C. H, 1889. Revision of the North American pocket mice. N. Amer. Fauna,
1:1-29.

Nero, R. W. 1958. Additional pocket mouse records. Blue Jay, 16:176-179.

— . 1965. Recent pocket mouse records for Saskatchewan. Blue Jay, 23:36-38.

Osgood, W. H. 1900. Revision of the pocket mice of the genus Perognathus. N. Amer.
Fauna, 18:1-73.

Pefaur, j. E., and R. S. Hoffmann. 1974. Notes on the biology of the olive-backed
pocket mouse Perognathus fasciatus on the northern Great Plains. The Prairie
Nat., 6:7-15.

Pires, F. D. de Avila-. 1965, The type specimens of Brazilian mammals collected by
Prince Maximilian zu Wied. Amer. Mus. Novit., 2209:1-21.

Power, D. M., and J. R. Tamsitt. 1973. Variation in Phyllostomus discolor (Chirop-
tera: Phyllostomatidae). Canadian J. Zool., 51:461-468.

Ridgway, R. 1912. Color standards and color nomenclature. Published by the author.

Washington, D.C., hi + 44 pp. + LIII plates.

Rohlf, F. j. 1971. MINT user’s manual. Unpublished manual, Dept, of Ecology and
Evolution, State Univ. of New York, Stony Brook, New York, 45 pp.

SwENK, M. H. 1940. A study of the geographical and ecological distribution of the
buffy plains pocket mouse {Perognathus flavescens flavescens), with description of
a new subspecies from Nebraska. Missouri Valley Fauna, 3:1-8.

Thomas, O. 1893. Description of a new species of Perognathus from Colorado. Ann.
Mag. Nat. Hist., ser. 6, 11:405-406.

Turner, R. W. 1974. Mammals of the Black Hills of South Dakota and Wyoming,
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Wied, Maximilian, Prince of. 1839. Uber einige Nager mit ausseren Backentaschen

102

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VOL. 48

aus dem westlichen Nord-America. I. Uber ein paar neue Gattungen der Nagethiere
mit ausseren Backentaschen. Nova Acta Physico-Medica, Academiae Caesareae

Leopoldino-Carolinae, 19:367-374.

Williams, D. F. 1978fl. Karyological affinities of the species groups of silky pocket
mice (Rodentia, Heteromyidae). J. Mamm., 59:599-612.

1978/?. The systematics and ecogeographic variation of the Apache pocket

mouse (Rodentia: Heteromyidae). Bull. Carnegie Mus. Nat. Hist., 10:1-57.

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ISSN 0097-4463

ANNALS

0/ CARNEGIE MUSEUM

CARNEGIE MUSEUM OF NATURAL HISTORY
4400 FORBES AVENUE « PITTSBURGH, PENNSYLVANIA 15213
VOLUME 48 1 June 1979 ARTICLE 6

PALEONTOLOGY AND GEOLOGY OF THE BADWATER
CREEK AREA, CENTRAL WYOMING. PART 17.

THE LATE EOCENE SNAKES

J. Alan Holman^

Abstract

Snake remains of Uintan mammal age (late Eocene) of the Badwater Creek area,
central Wyoming are identified as Dunnophis sp., Coniophis sp., Dawsonophis wyo-
mingensis n. gen. et sp., Ogmophis voorhiesi, Boavus sp. Dawsonophis is a small
boid snake quite distinct from fossil and living forms and of uncertain subfamilial rela-
tionships.

Introduction

Aside from scattered references to large sea snakes of the family
Palaeophidae (Holman, 1979), late Eocene snakes are very poorly
known. In fact, only one late Eocene snake fauna, a fauna from the
Twiggs Clay in Georgia (Holman, 1977) has previously been reported.
This fauna was of Duchesnean (latest Eocene) mammal age.

The present paper deals with snake remains from Locality 6 of the
Badwater Creek area, Natrona County, Wyoming. Locality 6 is
thought to represent the Uintan (earlier late Eocene) mammal age
(Krishtalka and Setoguchi, 1977). Thus, the present paper is the first
report of snakes of Uintan age in the New World. A Bridgerian (middle
Eocene) mammal age snake fauna was reported by Hecht (1959). The
material discussed in the present paper is housed in the collections of
the Carnegie Museum of Natural History (CM).

^ Address: The Museum, Michigan State University, East Lansing, Michigan 48824.
Submitted 15 November 1978.

103

104

Annals of Carnegie Museum

VOL. 48

Acknowledgments

I wish to thank Dr. Mary Dawson of the Carnegie Museum of Natural History for the
privilege of studying these snake fossils. Merald Clark made the drawings.

Systematic Paleontology

Class Reptilia
Order Squamata
Suborder Serpentes
Familia Incertae Sedis
Genus Dunnophis Hecht, 1959
Dunnophis sp. indet.

Material. — ^One fragmentary vertebra, CM 35003.

Remarks tiny vertebra appears to represent the genus Dun-

nophis Hecht based on its short, knobbed, neural spine which is re-
stricted to the posterior third of the dorsal portion of the neural arch.
This fragmentary bone is much smaller and has a much larger neural
canal than the nominate species Dunnophis microechinis from the
middle Eocene of the Bridger Formation of the Tabernacle Butte area
of Wyoming (Hecht, 1959).

Family Aniliidae
Genus Coniophis Marsh, 1829

Coniophis sp. indet.

Material. — Three fragmentary trunk vertebrae, CM 35004.

Remarks.-— AW of these fragmentary vertebrae lack distinct neural
spines and appear to represent juvenile individuals of the genus Con-
iophis. These vertebrae are too incomplete for specific identification.
Two species, Coniophis carinatus Hecht and Coniophis platycarina-
tus Hecht, are known from the Bridgerian (middle Eocene) of the
Tabernacle Butte area of Wyoming (Hecht, 1959).

Family Boidae
Subfamilia Incertae Sedis
Dawsonophis, new genus

Diagnosis.— A distinctive genus of rather small (estimated length
about LI m) boid snake of uncertain familial relationships distin-
guished from other boid genera on the basis of the trunk vertebrae
having (1) a long, low, neural spine about twice as long as its height,
its anterior edge strongly curved anteriorly, and its posterior part swol-
len posteriorly and flattened dorsally; (2) a thin straight zygosphenal
roof; (3) a depressed cotyle and condyle; (4) a strong oblanceolate
hemal keel; and (5) a moderately vaulted neural arch.

1979

Holman — Late Eocene Snakes

105

Fig. 1. — Holotype trank vertebra fused posteriorly with a very incomplete trunk vertebra
of Dawsonophis wyomingensis, new genus and species, from the late Eocene of the
Badwater Creek area, central Wyoming. Upper left dorsal, upper right lateral, middle
ventral, lower left anterior, lower right posterior. The line equals 2 mm.

Dawsonophis wyomingensis , new species

Holotype.— CM 14444. A trunk vertebra fused posteriorly with a
very incomplete trunk vertebra (Fig. 1).

Paratype.—A third trunk vertebra, CM 14445, collected by the same
collectors at the same locality at the same time.

Horizon. — ^Late Eocene, Uintan Mammal Age, Hendry Ranch Mem-
ber.

Loc£i//fy. —SESE, S. 14, T. 39 N., R 89 W, Natrona County, Wy-
oming. Referred to as Locality 6.

106

Annals of Carnegie Museum

VOL. 48

Collectors .--Cvdxg Black, Mary Dawson, and Peter Robinson in
1962.

Etymology. — ^The name recognizes Dr. Mary Dawson for her contributions in verte-
brate paleontology. The specific name refers to the state where the fossil material was
collected.

Description of the holotype. — -In dorsal view, the vertebra is wider than long. The
anterior border of the zygosphene is straight. The neural spine is long and is over one-
half the length of the distance between the posterior edge of the neural arch and the
anterior edge of the cotyle. The neural spine is swollen posteriorly with the swollen
portion being flattened dorsally. The neural spine is about twice as long as it is high.

The prezygapophyseal articular facets are broken off. Nevertheless, the fragmentary
vertebra that is articulated posteriorly shows, in dorsal view, a partially visible right
prezygapophyseal facet which appears to be ovoid in shape. The accessory process of
this facet is rounded and poorly developed.

In lateral view, the neural spine is long and low, being about twice as long as it is
high. Its anterior edge is strongly curved anteriorly. The posterior part of the neural
arch is upswept. The subcentral ridges are moderately curved upward.

In ventral view, the hemal keel is very well-developed and strong and has deep
grooves between it and the subcentral ridges. The hemal keel is oblanceolate in shape
(see Auffenberg, 1963: 153, fig. 1). The paradiapophysis is not strongly divided into
parapophyseal and diapophyseal portions.

In anterior view, the zygopophyseal area is moderately thick. The upper border of
the zygosphene is flat. The neural canal is rhomboidal in outline and filled with cemented
matrix. The cotylar area inscribes about one-fourth more area than the area inscribed
by the neural canal outline. The cotyle is depressed. In posterior view, the neural arch
is moderately vaulted.

Measurements of the holotype are as follows: length of vertebra from cotylar border
through posterior edge of neural arch, 6.9 mm; dorsal view length of neural spine, 3.7
mm; estimated anterior width of vertebra, 9.2 mm; height of vertebra from bottom of
hemal keel through top of neural spine, 7.2 mm.

Paraiype.— The paratype (Fig. 2) may be from a more anterior portion of the trunk
than the holotype in that the hemal keel is wider and not nearly as oblanceolate in shape
as in the holotype. This type of change in vertebral proportions toward the anterior end
of the snake vertebral column is shown in Hoffstetter and Rage (1972: 21, fig. 7b and
c). The vertebra is very fragmentary, but it is about the same size as the holotype,
shows the depressed cotyle and condyle, and has a moderately vaulted neural arch.

Remarks .—Daws onophis is here compared with several genera of
fossil boids of similar vertebral form. It can be distinguished from all
of these on the basis of its depressed cotyle and condyle.

It may be separated from Pseudoepicrates stanolseni (Vanzolini) of j
the early middle Miocene of Florida on the basis of the much lower
neural spine and much less massive zygosphene in Dawsonophis
{Pseudoepicrates is illustrated in Auffenberg, 1963: 160, fig. 8).

Dawsonophis may be separated from Ogmophis compactus Lambe
of the lower Oligocene of Saskatchewan on the basis of its less welF
developed accessory processes and its oblanceolate hemal keel. Og-
mophis compactus , a very distinctive large form of paraphyletic Og-
mophis, has relatively well-developed accessory processes for a boid
(Holman, 1972: 1625, fig. 6) and a hemal keel that is of uniform width
throughout its length.

1979

Holman — Late Eocene Snakes

107

Fig. 2.~Paratype vertebra of Dawsonophis wyomingensis, new genus and species, from
the late Eocene of the Badwater Creek area, central Wyoming. Upper left dorsal, upper
right lateral, middle ventral, lower left anterior, lower right posterior. The line equals
2 mm.

Dawsonophis may be separated from Palaeopython de Rochebrune,
which has several species in the Eocene and Oligocene of Europe, on
the basis of its differently shaped neural spine and much less massive
zygosphene (Paleopython filholi de Rochebrune is illustrated in Rage,
1974: 286, fig. 3).

Dawsonophis may be separated from Huberophis georgiensis Hol-
man of the late Eocene of Georgia on the basis of the much longer

108

Annals of Carnegie Museum

VOL. 48

neural spine, the oblanceolate hemal keel, and the shorter vertebra
{Huberophis is illustrated in Holman, 1977: 143, fig. 2).

Dawsonophis may be separated from Paraepicrates brevispondylus
Hecht from the middle Eocene of Wyoming on the basis of the differ-
ently shaped neural spine and its oblanceolate hemal keel. Paraepb
crates has a much shorter hemal spine and a wider, less distinct hemal
keel (Hecht, 1959: plate 55).

Finally, Dawsonophis may be separated from Boavus Marsh, which
has several species in the Eocene of Wyoming, on the basis of the
differently shaped neural spine, oblanceolate hemal keel, and much
less massive zygosphene. The neural spine of Boavus is much higher
and shorter, the hemal keel narrower, and the zygosphene massive.
Several species of Boavus have their vertebrae illustrated in Gilmore,
1938.

It seems quite possible that the very distinctive boid genus Daw-
sonophis might represent an undescribed boid subfamily, but one hes-
itates to describe a subfamily on trunk vertebrae alone.

Subfamily Erycinae
Genus Ogmophis Cope, 1884
Ogmophis voorhiesi Holman, 1977

Material.— Two trunk vertebrae and the consolidated tip of the tail consisting of
several fused caudal vertebrae, CM 35005.

Remarks.— As several authors have commented, the status of the
genus is questionable and it probably will not become clear until more
complete fossil skeletons are recovered. Based on its small size, the
shape of the long neural spine in dorsal view, the wide hemal keel, and
the somewhat indistinct subcentral ridges, the trunk vertebrae are as-
signed to Ogmophis voorhiesi, a form previously known only from a
single vertebra from the upper Eocene Twiggs Clay, Twiggs County,
Georgia. The wide range of this species in the late Eocene is of con-
siderable interest and one wonders if it indicates similar climates in
the two areas.

The recovery of the fused caudal vertebrae representing the solid,
blunt tail-tip is of special importance as it indicates that O. voorhiesi
is unquestionably a member of the subfamily Erycinae (see Hoffstetter
and Rage, 1972: 7).

Measurements of the Wyoming vertebrae compared with the holo-
type specimen are as follows: greatest length of centrum from para-
diapophyses through the centrum ranges from 2.2 to 2.5 mm (3.0 mm
in type); neural spine length in most complete Wyoming specimen is
1.1 mm (1.2 mm in type); length of the consolidated tip of the tail
composed of fused caudal vertebrae is 4.4 mm (unknown in type).

1979

Holman- — Late Eocene Snakes

109

Subfamily Boinae
genus et sp. indet.

Marerw/.— Fragmentary trunk vertebra, CM 35006.

Remarks .-—This vertebra lacks many diagnostic features as the pos-
terior portion of the centrum is missing and the prezygapophyses are
broken. But the vertebra appears to be from a very distinctive form
with a very robust neural spine and hemal keel. It is very possible that
this specimen represents a new form.

Genus Boavus Marsh, 1871
Boavus sp. indet.

Material. — A fragmentary trunk vertebra, CM 35007.

Remarks. — =The single vertebra appears to represent the genus Boa-
vus and it differs from Paraepicrates Hecht of the Bridgerian (middle
Eocene) Tabernacle Butte area of Wyoming in having a much thinner
hemal keel. The Badwater Creek specimen is too fragmentary for spe-
cific identification. Hecht (1959) reports this genus from the Bridgerian
(middle Eocene) Tabernacle Butte area of Wyoming.

Discussion

The study of late Eocene snakes is yet in its infancy as only two
small faunas are known— the Badwater Creek fauna of Wyoming dis-
cussed herein and thought to represent the Uintan (earlier late Eocene)
mammal age, and the Twiggs Clay fauna of Georgia (Holman, 1977)
thought to represent the Duchesnean (latest Eocene) mammal age.
Although both faunas are represented only by primitive snakes (In-
fraorder Henophidia), there are few taxonomic resemblances beyond
this point. The only taxon shared by the two faunas is the erycinine
boid Ogmophis voorhiesi.

It is interesting to note that the Badwater Creek fauna shares three
genersL—Coniophis, Dunnophis, and Boavus- — ^in common with the
Bridgerian (middle Eocene) mammal age Tabernacle Butte area fauna
of Wyoming (Hecht, 1959) that are not found in the Twiggs Clay fauna
of Georgia. I suspect that rather than being of stratigraphic importance,
this situation reflects regional and ecological differences. The paleoen-
vironment for the Twiggs Clay area during Duchesnean times is
thought to have been of a tropical estuarine nature.

Literature Cited

Auffenberg, W. 1963. The fossil snakes of Florida. Tulane Studies Zool., 10:131-
216.

Gilmore, C. W. 1938. Fossil snakes of North America. Geol. Soc. Amer., Spec. Paper,
9:1-96.

110

Annals of Carnegie Museum

VOL. 48

Hecht, M. K. 1959. Amphibians and reptiles. Pp. 130-146, in The Geology and Pa-
leontology of the Elk Mountain and Tabernacle Butte Area, Wyoming (P. O.
McGrew, ed.), Bull, Amer. Mus. Nat. Hist., 117:121-176.

Hoffstetter, R., and J. C. Rage. 1972. Les Erycinae fossilies de France (Serpentes,
Boidae) comprehension et histoire de la sous-famille. Ann. Paleontologie (ver-
tebres), 58:81-129.

Holman, J. A. 1972. Herpetofauna of the Calf Creek local fauna (lower Oligocene:

Cypress Hills Formation) of Saskatchewan. Canadian J. Earth Sci., 9:1612-1631.
— — — . 1977. Upper Eocene snakes (Reptilia, Serpentes) from Georgia. J. Herpetology,
11:141-145.

— — — . 1979. A review of North American Tertiary snakes. Publ. Mus. Michigan State
Univ., Paleont. Ser., in press.

Krishtalka, L., and T. Setoguchi. 1977. Paleontology and geology of the Badwater
Creek area, Central Wyoming. Part 13. The late Eocene Insectivora and Dermop-
tera. Ann. Carnegie Mus., 46:71-99.

Rage, J. C. 1974. Les serpentes des phosporites du Quercy. Palaeovertebrata, 6:
274-308.

ISSN 0097-4463

woy, 73

I ANNALS

I of CARNEGIE MUSEUM

; CARNEGIE MUSEUM OF NATURAL HISTORY
I 4400 FORBES AVENUE ® PITTSBURGH, PENNSYLVANIA 15213
I VOLUME 48 1 June 1979 ARTICLE 7

I SUBSTRATE SELECTION BY THREE SPECIES OF

' DESMOGNATHINE SALAMANDERS FROM

I SOUTHWESTERN PENNSYLVANIA:

AN EXPERIMENTAL APPROACH

I Anthony J. Krzysik^

Edward B. Miller^

Abstract

f'

An experimental technique is developed to quantify substrate selection by stream-
bank salamanders. Substrate particle size has been shown in field studies by the senior
author to be an important niche parameter. Interspecific differences in substrate size
selection is hypothesized to play a central role in determining coexistence patterns and
habitat selection in Desmognathus. Substrate selection experiments in the laboratory
are desirable for controlling field variables and critical to understanding community
organization in these salamanders.

Introduction

Researchers have studied interactions between species of salaman-
ders in the laboratory (Jaeger, 1974; Thurow, 1975, and others) but the
study of substrate selection has been ignored. There is theoretical
(Mac Arthur and Pianka, 1966) as well as empirical (Schoener, 1974)
justification for hypothesizing that differential microhabitat utilization
! by coexisting community members is usually the most important factor
permitting stable coexistence and minimizing actual or potential com-
petition. Krzysik (1977) found a strong intra- as well as interspecific

' Address: Pymatuning Laboratory of Ecology, University of Pittsburgh, Linesville,
Pennsylvania 16424.

^ Address: Washington and Jefferson College, Washington, Pennsylvania 15301.
Submitted 21 November 1978.

1

112

Annals of Carnegie Museum

VOL. 48

Table 1. — Substrates and their particle sizes utilized in substrate selection experiments.

Substrate

Particle size

Notes

Sand

0.05-0.5 mm

Fine construction sand^

Small gravel

0.5-1 cm

IB construction gravel

Medium gravel

1.5-3 cm

2B construction gravel

Large gravel

4-8 cm

Hand selected round and smooth glacial
deposit rocks

Rocks

8-15 cm by

1-3 cm thick

Hand selected flat sedimentary rocks,
comprising the rubble of stream-banks

^ A few small rocks (1-2 cm) were sprinkled on the surface, otherwise this substrate
was totally avoided by salamanders.

(particularly the latter) relationship between body size and substrate
selection by salamanders in the field, and showed that substrate par-
ticle size is an important microhabitat parameter determining coexis-
tence patterns within a desmognathine salamander community.

Substrate selection experiments under laboratory conditions are de-
sirable because of the large number of uncontrollable variables in the
field. Substrate size is easily controlled, and such important field vari-
ables as microhabitat moisture, distance from the stream, size and type
of cover, and others may be eliminated. A firm experimental basis for
infra- and interspecific substrate relationships among desmognathine
salamanders is important in understanding coexistence patterns and
microhabitat resource allocation in this community. This report rep-
resents a preliminary investigation of an experimental procedure for
quantifying substrate size selection by streambank salamanders, uti-
lizing three sympatric desmognathines from southwestern Pennsylva-
nia.

Materials and Methods

Nine all-glass aquaria (76 cm by 30 cm by 30 cm) were separated into three 25 cm by
30 cm compartments by plexiglass dividers. The dividers were 10 cm high and several
3 mm holes were drilled into each so that an equal water level could be maintained.
Each of the three compartments was filled to the top of the divider with the substrates
that were being compared. Water was added to each aquarium until it reached a depth
of 2 to 3 cm below the surface of the substrate. A square board 10 cm on a side and 1.3
cm thick was placed in the center of each compartment. The board was raised about 1
cm over the substrate when sand or fine gravel was utilized as the substrate. The tanks
were carefully sealed with aluminum foil containing small vents to provide limited air
flow. Thus a high humidity was constantly maintained in all tanks (>90%) and all sub-
strates utilized were constantly wet. Five different substrates, which formed a particle
size gradient, were utilized (Table 1). A sixth “substrate”, the glass sides of the aquaria,
was additionally considered.

Three species of sympatric salamanders (Desmognathus monticola, D.fuscus, and D.
ochrophaeus) were collected at Rachelwood Wildlife Research Preserve (Westmoreland
Co., southwestern Pennsylvania). Animals used in the study were anesthetized in a

1979

Krzysik and Miller — Desmognathine Ecology

13

Table 2— The species of Desmognathus and their respective body sizes used in the

study.

Species

Snout-vent length
(mm)‘

Total length
(mm)

D. monticola (6 6)

68--74

124-136

D. fuscus ((3 (?)

60-68

99-125

D. monticola (immature)

45-51

80-105

C. ochrophaeus (c3 c3)

41-45

77-93

^ Tip of snout to the posterior angle of the vent.

weak solution of tricaine methanesulfonate; snout- vent and total length were recorded,
and animals were toe-clipped for identification (Table 2). Animals were stored in moist
plastic bags in a refrigerator at 5 to 8°C. Three separate experimental runs were made
because only three of the possible substrates could be tested at a time. The three ex-
periments are: 1) sand, medium gravel, rocks; 2) small gravel, medium gravel, large
gravel; 3) medium gravel, large gravel, rocks. Tanks were arranged such that each
substrate occupied the center compartment of three of the nine tanks. Three mature
males (one of each species) were utilized during an experiment. One individual was
placed on the board in the center compartment of each tank. Thus the three individuals
of any given species were initially placed in compartments that contained each of the
three possible substrates. The nine animals were tested for three days under constant
darkness with the following routine. Individuals were introduced into the aquaria at
around 1000 hrs and were looked for around 2000 hrs (pm observation) with a red light
and with minimal disturbance to the substrate. The substrate on which each individual
was found was recorded. The following morning (around 0900 hrs — ^am observation) the
individuals were again located, their positions recorded as before and they were re-
moved. This procedure was repeated on two more consecutive days using the same
individuals. In this way each individual of each species could initially be placed on each
of the three possible substrates. A minimum of six individuals of each species was
examined. After the experiments with the mature males were completed, nine immature
Desmognathus monticola were individually placed in the nine aquaria. They were sim-
ilarly rotated daily so that each individual was initially placed on each of the three
possible substrates.

Model I three-way analysis of variance utilizing arcsine transformations on the fre-
quency data (Sokal and Rohlf, 1969) was used to determine if there were interactions
among the three factors— -species, substrate, and time of observation (am or pm),
goodness of fit (Siegel, 1956) was employed to determine whether or not substrates were
differentially utilized. The data from these three experiments were combined (appro-
priate corrections were made because all substrates were not equally represented ^ter
pooling the data) to evaluate the allocation of the substrate microhabitat resource gra-
dient among the members of this hypothetical community of Desmognathus. The Kol-
mogorov-Smirnov two-tailed and one-tailed tests for large samples (Siegel, 1956) were
used to determine if any differences or a priori determined directional differences, re-
spectively, were present in substrate selection.

Values of niche overlap (Schoener, 1968) were calculated as

Oy = 1 - 1/2^ |Plk - Pjk

where Pj^ and Pj^ are the proportions of the number of times species i and j
were found on substrate k. can possess values between 0 and 1, indicating no
overlap or complete overlap in resource utilization. Niche overlap quantifies the

114

Annals of Carnegie Museum

VOL, 48

0,4

0.3

D. fuscus

0.2

0.1

jn □

■ecki Lorga Medium Small Send ©loss

Gravel gravel Grovel fArbereoll

Substrate

Fig. 1.— Frequency histograms of four Desmognathus body sizes distributed among six
substrate categories. See Table 1 for description of substrate categories.

sharing of the substrate resource gradient between two species. Niche breadth
(Levins, 1968) was calculated as

B, = l/yp^k

where pik is as above. Niche breadth quantifies the diversity or breadth of utilization of
the substrate resource gradient by a given species.

Results

Analysis of variance of transformed frequency data confirmed that
there was no relationship between the time of day the animals were

1979

Krzysik and Miller— Desmognathine Ecology

115

Table 3. — Values of niche overlap and niche breadth for the utilization of the substrate
microhabitat resource gradient.

Desmognathus

fuscus

Desmognathus

monticola

(immature)

Desmognathus

ochrophaeus

D. monticola (mature)

Niche overlap
0.911

0.814

0.392

D. fuscus

__

0.735

0.327

D. monticola (immature)

—

—

0.572

D. monticola (mature)

D. fuscus

D. monticola (immature)

D. ochrophaeus

Niche breadth

2.75

2.43

3.30

4.86

observed and either the substrate or the species (P > 0.75); however,

there was a significant difference {P < 0.005) in the relationship be-
tween a species and the substrate on which it was found. Analyzing
all three experiments by goodness of fit showed that there was no
significant difference in the way that mature D. monticola and D.
fuscus males utilized the substrate {P > 0.10) but there was a signifi-
cant difference among all other combinations {P < 0.001). The data
for all substrates were combined and the frequency distribution of the
four desmognathines in the six substrate categories is shown in Fig.
1 . Again there is no significant difference in the utilization of substrate
categories by mature D. monticola and D. fuscus, but there are highly
significant differences in all other species combinations {P < 0.02 in
the case of mature and immature D. monticola; and P < 0.001 in all
other comparisons). Values of niche overlap and niche breadth are
given in Table 3.

Discussion

Mature males of Desmognathus monticola and D. fuscus, the larg-
est members of the southwestern Pennsylvania desmognathine com-
munity, possess a similar preference for the coarsest substrate avail-
able (Fig. 1, Table 3). Immature D. monticola can be found associated
with substrates possessing a smaller particle size than mature
males. Desmognathus ochrophaeus, the shortest and thinnest sala-
mander, was found not only utilizing a wide range of substrate sizes
but preferred smaller substrate particle sizes (Fig. 1). This was the
only species to show consistent arboreal tendencies. The adaptive sig-
nificance of substrate selection can be inferred to be the following:
large salamanders require larger openings into the substrate to escape
predators, locate overwintering cavities, and follow ground water in

116

Annals of Carnegie Museum

VOL. 48

periods of drought. Smaller species not only can utilize a wider range
of openings for these purposes, but natural selection should favor those
individuals that select substrates with smaller openings, hence mini-
mizing predation from larger vertebrates, including other salamanders.

These patterns, along with the fact that body size appears to be an
important determinant of interspecific interference competition
(Morse, 1974; Thurow, 1975; personal observations), can be used to
predict community structure in this genus. Desmognathus monticola
and D. fuscus require habitats possessing a coarse substrate, but be-
cause they possess a high overlap in substrate utilization (0.911) D.
fuscus, the smaller species, should generally be rare in habitats con-
taining the coarsest substrates (for example, those selected by D. mon-
ticola). When in sympatry with D. monticola at habitats which are
suboptimal for D. monticola, D. fuscus should exhibit a shift in mi-
crohabitat preference. Desmognathus ochrophaeus because of its
wide breadth of substrate utilization (4.86) and low overlap with the
other species (Table 3) should be an abundant widespread species
commonly associated with its congeners, and also found in habitats
that are unavailable for the other two species (those with fine sub-
strates). The presence of immature monticola in smaller substrates
reduces intraspecific competition in this species for shelter and food.
The arboreal tendencies of D. ochrophaeus have been observed in the
field (Hairston, 1949; personal observations) and may be a mechanism
of exploiting food and space resources not utilized by other members
of this community. These predicted patterns of desmognathine com-
munity organization have been substantiated by field studies (Krzysik,
1977).

Acknowledgments

Thanks are expressed to Michael Mares, who was Miller’s research director, and
Stephen Tilley for a critical review of the manuscript, and to Richard Hartman for pro-
viding research facilities. The senior author was supported by a Mellon-Rachelwood
Wildlife Research Grant and Edward Miller by an NSF Undergraduate Research Pro-
gram Grant (SMI76-03428A01) to the Pymatuning Laboratory of Ecology.

Literature Cited

Hairston, N. G. 1949. The local distribution and ecology of the plethodontid sala-
manders of the southern Appalachians. Ecol. Monogr., 19:49-73.

Jaeger, R. G. 1974. Interference or exploitation? A second look at competition be-
tween salamanders. J. Herpetol., 8:191-194.

Krzysik, A. J. 1977. Resource allocation, coexistence and the niche structure of a
stream-bank salamander community. Unpublished Ph.D. dissert., Univ. Pittsburgh,
113 pp.

Levins, R. 1968. Evolution in changing environments. Monogr. Pop. Biol., Princeton
Univ. Press, 120 pp.

1979

Krzysik and Miller — Desmognathine Ecology

117

MacArthur, R. H., and Pianka, E. R. 1966. On optimal use of a patchy environment.

Amer. Nat., 100:603-^609.

Morse, D. H. 1974. Niche breadth as a function of social dominance. Amer. Nat.,
108:818-830.

Schoener, T. W. 1968. The Anolis lizards of Bimini: resource partitioning in a complex
fauna. Ecology, 49:704-726.

— — . 1974. Resource partitioning in ecological communities. Science, 185:27-39.
Siegel, S. 1956. Nonparametric statistics. McGraw-Hill, New York, 312 pp.

SoKAL, R. R. AND Rohlf, J. 1969. Biometry. W. H. Freeman, San Francisco, 776 pp.
Thurow, G. 1975. Aggression and competition in eastern Plethodon (Amphibia, Uro-

dela, Plethodontidae). J. HerpetoL, 10:277-291.

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VOLUME 48 1 June 1979 ARTICLE 8

THE LAS AVISPAS BURIAL PLATFORM AT
CHAN CHAN, PERU

Thomas Pozorski

Assistant Curator, Section of Man

Abstract

Recent investigations at the prehispanic city of Chan Chan have uncovered evidence
of large platforms formerly used as burial structures by the kings of the Chimu empire
(A.D. 1000-1470). One such burial platform, Las Avispas, contained the body of a
Chimu king along with the bodies of at least 300 young women who were ceremonially
sacrificed at the time of or shortly after his death.

Introduction

The prehispanic city of Chan Chan, located on the north coast of
Peru, was the capital of the Chimu empire which flourished from A.D.
1000-1470. Chan Chan covers an overall area of 25 square km and has
a central core area of 6 square km. This central core consists of nine
large compounds, or ciudadelas, surrounded by smaller architectural
units (intermediate architecture) that housed the elite class and their
retainers (Klymyshyn, 1976), as well as small irregular agglutinated
rooms (SIAR) that housed the artisan class of the city (Topic, 1977).

One of the most distinctive features of each ciudadela is the burial
platform, which is an elevated rectangular structure made of adobes
or tapia (tamped earth and rocks) that contains a number of chambers
laid out in a regular pattern. The primary purpose of the structure was
to contain the body of a Chimu king and the bodies of dozens of his
retainers.

Conrad (1979) has noted that there are nine such platforms within
Submitted 23 November 1978.

119

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VOL. 48

Fig. 1.— Simplified plan of the central core of Chan Chan showing distribution of burial
platforms throughout the city.

1979

PozoRSKi— -Las Avispas Burial Platform

121

Chan Chan (Fig. 1), a fact that correlates rather well with known king
lists from Chimu ethnohistory. The ciudadelas of Chan Chan were
built sequentially, one at a time. The Chimii ruler used a ciudadela as
an administrative center and as a palace while he lived and was buried
in the associated burial platform when he died. The new Chimu king
built a new ciudadela and repeated the same pattern, while the ciu-
dadela of the dead king was closed to the main activities of the city.

Before recent excavations, past investigators such as Tschudi and
Rivero (1855), Hutchinson (1873), Squier (1877), Feet (1903),, Kroeber
(1926), Holstein (1927), Horkheimer (1965), and Rodriguez Suy Suy
(1970) had speculated on the nature and significance of the Chan Chan
burial platforms, but none of these ideas was supported by specific
data. In 1969, Kent Day, on the basis of a detailed survey of the
ciudadelas of Chan Chan, hypothesized that the structures were burial
platforms. His theory led to more intensive investigation.

Excavations conducted in 1970 at the platform called Las Avispas
offer definite evidence in support of the burial platform theory. This
platform was chosen for excavation because, compared to others in
Chan Chan, it is in relatively good condition. The data gathered from
these excavations provided a meaningful base upon which a detailed
survey of other platforms was performed.

Most burial platforms within Chan Chan are located in the southeast
portion of ciudadelas (Fig. 1). However, Las Avispas is located just
outside the northeast corner of ciudadela Laberinto. Its association
with Laberinto is strengthened by the fact that in the southeast part
of that ciudadela there is no burial platform. Instead, there is a large
low rectangle of stone and adobe that appears to have sectioned off
an area for a burial platform. For some unknown reason, the platform
was not built where planned, but just outside the ciudadela in the form
of Las Avispas.

Architectural Description

The Las Avispas burial platform has many architectural character-
istics common to all burial platforms within Chan Chan. The platform
(Fig. 2) is set within a former “entry court,” a wall enclosure with a
bench along the south side reached by a free-standing ramp (Fig. 2h)
which eventually leads into a series of rooms. This “entry court”
pattern is frequently repeated within the ciudadelas and intermediate
architecture throughout Chan Chan. The former use of the enclosure
as an entry court explains why its ramp is so close to the back of the
platform.

The entire burial platform structure measures 49 m by 32 m and
consists of the elevated burial platform complex with its chamber area,
access ramp system, a forecourt (A), and two sidecourts (B and C)

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VOL. 48

LAS AVISPAS

Fig. 2. — Plan of the Las Avispas burial platform showing architectural layout, chamber
distribution, and location of dedicatory burials within the surrounding entry court.

adjacent to the forecourt. Two nonaligned entrances exist on the north
side^ — the outer one gives access to both sidecourts (B and C), whereas
the inner one leads to the forecourt (A). On the south side of the
forecourt there is a bench in front of a small central room (Fig. 2i) and
two flanking rectangular rooms (Fig. 2j-=k). A central free-standing

1979

PozoRSKi — Las Avispas Burial Platform

123

Table \— Articulated human skeletal remains found within the Las Avispas burial plat-
form.

Chamber

Human skeletal remains

Chamber 5

1 left capitate, lesser multangular, and second metacarpal

5 right metacarpals

1 left first metatarsal with first and second cuneiforms

1 whole right hand with right radius

1 left hand, carpals only

Chamber 6

1 right tibia, talus and patella

2 whole right feet

1 right innominate with basal lumber vertebra and sacrum

Chamber 13

1 left talus and distal tibia epiphysis

1 right patella, proximal tibia epiphysis, and distal femur
epiphysis

Chamber 14

1 left capitate and lesser multangular

1 left hamate, pisiform, and triquetral

Chamber 19

1 left innominate with sacrum

1 axis and atlas

Chamber 22

2 left hands, carpals only

5 phalanges of right foot

1 right calcaneum, cuboid, and all 5 metatarsals

ramp and a lateral ramp on the east side lead up to the bench. At the
top of the central ramp is a small wall that probably blocked the view
of the small central room (Fig. 2i). Within this room, excavation un-
covered a stone-lined bin containing crushed Spondylus species shell.

The east rectangular room (Fig. 2k) contains the entrance giving
access to a lateral ramp that leads up to the top of the chamber area.

At present, the platform measures 23 m by 32 m and contains 25
chambers (Fig. 2). Excavation showed that there are two stages of
construction represented. The first stage is a central rectangular block
demarcated by fine plastered faces on the west, south, and east sides;
this block contains 14 chambers (numbers 5, 6, 7, 9, 10, 11, 13, 14, 16,
17, 18, 20, 21, and 22 in Fig. 2). All except one of these chambers is
rectangular. The exception is a central T-shaped chamber that is con-
siderably larger than the others. The second construction stage is a
U-shaped addition that filled in the gap between the platform enclosure
and the original 14-chamber block; this addition contains 11 rectan-
gular chambers (numbers 1, 2, 3, 4, 8, 12, 15, 19, 23, 24, and 25 in Fig.
2).

Excavation within the chamber area consisted mainly of defining
corners of the chambers with as little digging as possible. In most
cases, three corners of each chamber were found after 50 to 100 cm

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Annals of Carnegie Museum

VOL. 48

Skeletons 1-9

Fig. 3. — Plan of chamber 19 showing uppermost nine female skeletons.

1979

PozoRSKi — Las Avispas Burial Platform

125

10 cm

N

A

Skeletons 10-12

beneath skeletons 1-9

Fig. 4. — Plan of chamber 19 showing four female skeletons found directly underneath
the nine skeletons shown in Fig. 3.

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Annals of Carnegie Museum

VOL. 48

Skeleton 13

beneath skeletons 10-12

Fig. 5. — Plan of chamber 19 showing one female skeleton found directly underneath the
four skeletons shown in Fig. 4.

1979

PozoRSKi — Las Avispas Burial Platform

127

of overburden had been removed. In each of chambers 7, 13, 17, 21,
and 25, a narrow trench was excavated to the floor, while chamber 19
was completely cleared. In general, the depth of the chambers of the
first construction stage is about 3 m while the depth of the second-
stage chambers is about 2 m.

Skeletal Evidence from the
Elevated Platform

All of the chambers had been greatly disturbed by looting before
excavations. Few bones could be associated with one another and
attributed to the same individual, except in cases where preserved skin
still held certain bones in articulated positions (Table 1). Near the floor
of chamber 19, which was completely excavated, the remains of 13
relatively complete individuals were uncovered. Nine bodies (Fig. 3)
rested in a layer over three more bodies (Fig. 4), which in turn rested
over another one (Fig. 5). Because the majority of the skeletons were
articulated, it was not difficult to assign most of the bones to specific
individuals (the numbers in Figs. 3, 4, and 5 designate skeleton num-
bers). Thus, the evidence from chamber 19 and from other articulated
remains suggest that most, if not all, of the individuals in the platform
were buried in a flexed position. It should also be stressed that, though
most bones were found in a disturbed condition, there was no evidence
of intrusive burials. Chamber walls are still intact, and all bones were
found in loose earth in and around the chambers themselves.

A total of 5,297 bones was collected during excavation (Table 2).
The minimum number of individuals originally interred in the platform,
based on left tibiae, was 93 (Table 2). This, however, cannot be taken
as the maximum possible, for it is highly unlikely that the same bones
of all the individuals originally interred have been discovered when,
at most, only about one-fourth of the total volume of the chambers of
the platform was excavated. Previously (Pozorski, 1971:103) I have
estimated a minimum total of people originally interred in the Las
Avispas platform as 200, but, upon reexamination, this figure seems
too low. Calculations based upon the excavated volume (one-fourth of
the total) times the left tibiae bone count (93) yields a figure of 372
individuals. The excavated volume times the lowest major bone count,
sacra (54), results in 216. To compute in another manner, if the 13
individuals found in chamber 19 are representative, then 13 times the
number of chambers (25) produces an estimate of 325 individuals. Cal-
culations involving more complicated factors such as chamber size or
density of bones per cubic meter result in much higher figures. Hence,
it is likely that at least 300 persons were once buried within the Las
Avispas burial platform.

Calculations of age estimates were made, mainly on the basis of

128

Annals of Carnegie Museum

VOL. 48

epiphyseal union (Flecker, 1942; McKern and Stewart, 1957; Steven-
son, 1924). A compilation of these ages for each bone is shown in
Table 3. Often, a definite time range for the age of a bone could be
determined if one epiphysis was fused and another unfused. However,
in the case of long bones, if both epiphyses were fused or if both were
not, then only one limit, upper or lower, could be established. The
same was true for bones in which only one end was preserved.

Because the majority of human bones were found in a disarticulated
state, ages for specific individuals could not, for the most part, be
determined. However, ages based on data derived from single bones
have been compiled to ascertain the age range of the general burial
population. Table 4 lists age percentages for all significant bones based
on data presented in Table 3.

To arrive at these percentages, values for both the right and left of
each bone were combined. Minimum and maximum age limits were
determined by considering only the extremes of the ranges defined on
the basis of epiphyseal union. For example, in Table 3, humeri contain
five categories (over 16; over 13-under 24; over 13-under 20; over 13~
under 18; and over 13) that define at least a lower age limit. By con-
sidering only the extreme lower limit (over 13), these categories, even
“over 16“ and the “over 16~under 18,” can be considered as “over
13“ because any bone indicating an age of over 16 years would also
be over 13 years old. Thus, for humeri, 126 out of a total sample of
135 (humeri of unknown age were omitted from the total sample) or
93.33% are over 13 years old. The same procedure was followed for
determination of an upper age limit which, in the case of humeri, is
“under 24.“

In Table 4, the minimum age limits of the human bone sample at Las
Avispas comprise the left column and the maximum age limits are in
the right column. An examination of this table reveals that from 75%
to over 90% of the persons buried were over 13 at death while 90%
were under 31 at death. Percentages progressively diminish for ages
between 13 and 31. Significantly, though, at least two-thirds or 67% of
the total sample lie between the ages of 17 and 24 years. This indicates
that a high proportion of the people buried in the platform died during
late adolescence or early adulthood. However, the actual proportion
of individuals who died between 17 and 24 years of age may be higher
than indicated by Table 4 for several reasons.

First, there is no corroborating evidence that anyone was over 31
or even close to it. No notable signs of aging such as lipping of the
vertebrae or loss of teeth were present. Second, only slightly more
than 15% of all clavicles indicate an age over 23 years. Third, many
bones that were recorded as “fused” were actually incompletely

Oth-

Human bones 2 3 4 5 6 7 9 10 12 13 14 15 16 17 18 19 20 21 22 23 24 25 er* Total

1979

PozoRSKi — Las Avispas Burial Platform

129

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Remains associated with the burial platform but not assignable to any specific chamber.

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Annals of Carnegie Museum

VOL. 48

Table 3. — Age estimates based on epiphyseal union of human bones found within the

Las Avispas platform.

Left humeri

Right humeri

over 16

27

over 16

20

over 13, under 24

17

over 13, under 24

21

over 13, under 20

9

over 13, under 20

16

over 16, under 18

4

over 13

8

over 13

4

under 18

3

under 18

4

under 20

1

under 20

1

unknown

unknown

2

total

71

total

68

Left ulnae

Right ulnae

over 15

20

over 15

22

over 14, under 23

9

over 14, under 23

19

over 14, under 19

5

over 14, under 19

3

over 14

14

over 15, under 23

6

over 15, under 23

3

over 14

13

under 19

3

under 19

4

under 23

2

under 23

1

unknown

4

unknown

5

total

60

total

73

Left radii

Right radii

over 16

29

over 16

26

over 16, under 23

3

over 16, under 23

2

over 14, under 23

15

over 14, under 23

14

over 14, under 20

1

over 14

4

over 14

6

under 20

7

under 20

9

under 23

1

under 23

2

unknown

unknown

8

total

59

total

73

Left femora

Right femora

over 14

29

over 14

36

over 14, under 20

8

over 14, under 20

4

over 14, under 22

11

over 14, under 22

13

over 13, under 20

3

over 13, under 20

2

under 20

13

over 13, under 22

2

under 22

4

over 13

2

unknown

1

under 20

14

total

69

^ unknown

5

total

78

Left tibiae

Right tibiae

over 14

39

over 14

41

over 14, under 23

8

over 14, under 23

3

over 14, under 20

1

over 14, under 20

3

over 13, under 20

1

over 13, under 23

4

over 13, under 23

3

over 13

4

over 13

6

under 20

15

under 20

15

under 23

2

under 23

4

unknown

16

unknown

16

total

88

total

93

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PozoRSKi— Las Avispas Burial Platform

131

Table 3. — Continued.

Left fibulae

Right fibulae

over 15

22

over 15

25

over 15, under 22

5

over 15, under 22

5

over 15, under 20

3

over 15, under 20

6

over 14, under 22

7

over 14, under 22

2

over 14

4

over 14

2

under 20

17

under 20

15

under 22

2

under 22

2

unknown

_6

unknown

J2

total

66

total

69

Left innominates

Right innominates

over 16

22

over 16

14

over 16, under 23

22

over 16, under 23

25

over 16, under 24

3

over 16, under 24

2

under 23

8

under 23

9

under 18

12

under 18

14

over 10.5

over 10.5

4

total

72

total

68

Left scapulae

Right scapulae

over 17

21

over 17

21

over 17, under 23

13

over 17, under 23

13

under 23

21

under 23

17

unknown

12

unknown

_8

total

67

total

59

Left ribs

Right ribs

over 18

165

over 18

180

under 24

381

under 24

342

unknown

J9

unknown

m

total

625

total

628

Left clavicles

Right clavicles

over 23

6

over 23

4

over 23, under 31

4

over 23, under 31

2

under 3 1

36

under 31

49

unknown

9

unknown

11

total

55

total

66

Sacra

over 18

10

over 18, under 33

11

over 18, under 23

4

over 17, under 22

10

over 16, under 22

10

under 22

8

under 24

1

total

54

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Annals of Carnegie Museum

VOL. 48

Table 4.— Age percentages for human bones found within the Las Avispas burial plat-
form.

Lower age limits

Upper age limits

Bone

Age

Percent*

Bone

Age

Percent*

Femora

under 22

52.48

Humeri

over 13

93.33

Fibulae

under 22

54.70

Femora

over 13

78.01

Tibiae

under 23

39.60

Tibiae

over 13

75.84

Ulnae

under 23

44.35

Ulnae

over 14

91.94

Radii

under 23

45.38

Radii

over 14

84.03

Scapulae

under 23

64.15

Fibulae

over 14

69.23

Humeri

under 24

56.30

Innominates

over 16

67.18

Sacra

under 24

61.11

Sacra

over 16

83.33

Ribs

under 24

67.70

Scapulae

over 17

64.15

Innominates

under 24

72.52

Ribs

over 18

32.30

Clavicles

under 31

90.10

Clavicles

over 23

15.84

Sacra

under 33

81.48

* Percentages are based on the total number of bones minus the number of bones of
unknown age.

fused^ — ^clearly showing lines separating epiphyses from the main body
of the bone. I

The same is true at the lower end of the scale. The evidence also
indicates that very few individuals were under 18. No unusually small j
or immature bones were noted. The only evidence for an individual j

under 15 is one mandible whose lower second molars were not erupted; |

all others had erupted second molars. This lower limit is also supported i

by the fact that only three skulls were found that had the basilar suture ‘

still open (an open basilar suture implies an age under 21 years; a |

closed suture, an age over 16, see Table 5). |

Further support for an age range of 17 to 24 years is provided by :
the fact that the skulls and mandibles (especially skulls with mandibles) jl
show a varied state of eruption of the third molars. Several individuals |
have their third molars, several others do not, and many were in the i|
process of acquiring them (Table 5). The intermediate stage of acquir-
ing third molars is not reflected because eruption was only counted if ji
the tooth was even with or within 1 mm of the general occlusional i
plane of the teeth (Garn et al., 1957:313-315), but because this group
of skulls and mandibles appears to be in the process of third molar
eruption, the Las Avispas materials would tend to support the so- '
called “normal age” for third molar eruption of 18 to 25 years (Suk, I
1919:374-377).

To determine the sex of the skeletal remains, secondary sex char-
acteristics of the skull, innominate, and sacrum were used. According I
to these criteria, all of the examinable specimens were female. |

1979

PozoRSKi — Las Avispas Burial Platform

133

Table 5.- — Third molar eruption data from skulls and mandibles found within the Las

Avispas burial platform.

Skulls without mandibles

erupted 4

not erupted 1 1

Skulls with mandibles

both maxillary and mandibular erupted 6

both maxillary and mandibular not erupted 6*

mandibular erupted, maxillary not erupted 1

maxillary erupted, mandibular not erupted 1*

Mandibles only

erupted 1 5

not erupted 25

Fragmentary skulls 16

Fragmentary mandibles 1

Special mandible

lower second molars not erupted, under 16 years old 1

* Three skulls do not have the basilar suture closed indicating an age of under 21.

Thus, the results of both age and sex determinations suggest some
rather strong conclusions. None of the data approximate the normal
death proportion for a population. It appears that most, if not all, of
the bodies interred in the Las Avispas platform were of young females.
The death of so many people of the same age and sex is best explained
as an unnatural event. Natural deaths and even epidemics or natural
disasters result in a predictable variety of individuals of different ages
and sexes unless burial patterns were highly selective. The only other
evidence of such selective burial in Chan Chan is Calavario de los
Incas, an oblique-shaped structure to the southwest of the central core
of Chan Chan, which apparently contains only the bodies of children
(Hrdlicka, 1911). Furthermore, the elaborate burial platform construc-
tion, plus its continuous use through time as suggested by the second
construction state, argues against the idea that it might contain persons
killed by an epidemic or natural disaster. Hence, the concept of “ritual
sacrifice” best explains the Las Avispas skeletal evidence. How this
massive sacrifice was done is not known. There are no indications on
any of the bones of the cause of death.

Because of its comparatively large size, central location, and unique
shape, it is likely that the T-shaped tomb in Las Avispas contained an
important person, someone more important than the people buried
around him. Perhaps this was the tomb of a king of Chimor who, when
he died, had his retainers or his harem killed and buried with him. Or,

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VOL. 48

when he died, a great number of young females from Chan Chan (or
perhaps from other parts of the empire) were sacrificed in his honor.
The two stages of chamber construction provide evidence of a sec-
ondary dedicatory sacrificial ceremony, carried out some time after
the former ruler’s death. Why just young women; why not older wom-
en? Why no males? Clearly the Chimu placed a high value on young
women as sacrificial victims. Was it because of their virginity? Or was
it just the opposite, that they were the ruler’s harem or his favorite
women? Many of these questions cannot be answered based on the
archaeological evidence.

Close examination of the innominate bones, however, indicates that
several individuals had given birth to children before their deaths (Erik
Trinkaus, personal communication). This partially refutes the hypoth-
esis on the value of virginity and suggests that at least some of the
victims were the ruler’s wives or concubines. Further support for the
wife/concubine theory comes from the early Spanish chronicler Cieza
de Leon who wrote about the burial customs generally practiced by
the natives of prehistoric Peru: “. . . it was the general belief among
all these Yunga Indians, and also the mountaineers of this kingdom of
Peru, that the souls of the dead did not die, but lived forever, and
came together with one another in the other world, where, as I said
before, they believe that they take their pleasure and eat and drink,
which is their chief delight. And firmly believing this, they buried with
the dead their best-loved wives, and their closest vassals and servants,
and their most prized possessions and arms and feathers, and other
ornaments of their person.’” (Cieza de Leon, 1959:310).

The women buried in the U-shaped addition of chambers were likely
to have been close to the king as well, though probably not as close
as the women buried in the original 14-chamber block. The individuals
in the U-shaped addition of chambers were presumably sacrificed some
time after the original interment, probably in a burial renewal cere-
mony noted by Cieza de Leon (1959:312) who stated that ”it was the
custom in olden times to open the tombs and renew the clothing and
food that had been buried in them.” Hence, in all likelihood, the Las
Avispas evidence reflects one manner of the ancient Peruvian practice
of interment of wives, concubines, and servents along with their de-
ceased lord and subsequent expansion and renewal of burial goods.

Artifactual Evidence from the
Elevated Platform

Abundant fragments of fine blackware pottery, woven textiles,
carved wood (often with shell inlay), metal adornments, and carved
and whole shells (including Spondylus species and Conus fergusoni)
were found in and immediately around the chamber area of the platform.

1979

PozoRSKi — Las Avispas Burial Platform

135

Food remains, probably placed as burial offerings in gourd bowls, were
also frequently found. Two forms of evidence argue for the association
of especially fine funerary offerings with the platform. First, the abun-
dance of artifacts drops dramatically in areas away from the elevated
platform, indicating a strong association of the artifacts with the bones
and the raised platform. Second, the enormous amount of looting,
leaving virtually no skeletons undisturbed, suggests that this activity
had proved worthwhile.

Skeletal Evidence from Areas Outside the
Elevated Platform

Seven burial pits were found during excavations in the entry court
area outside the burial platform enclosure. Six of these (Fig. 2a~f)
were found near the north entrance to the entry court, and the seventh
(g) was found in the south entrance. All except two (b, c) had been
disturbed by looters before excavation. Each pit contained one indi-
vidual except a and d, each of which had two individuals (see Donnan
and Mackey, 1978:344-349, for details of d). Evidence indicates that
all were seated and flexed. Burials a-f are females of the same age
range as those of the platform, while burial g is a slightly younger
person, age 9-14 years, whose sex could not be determined.

The chronology of these graves is equivocal. The burial pits were
excavated into the floor of the entry court; damaged sections of the
floor were replastered after the pits had been used and refilled. It is
possible that the graves were associated with the dedication or use of
the entry court before the existence of the burial platform. However,
it seems more likely, due to the age and sex range of the individuals
buried and the few artifacts associated with them, that they were in-
terred during the use of the burial platform.

Evidence of other human burials in the vicinity of the platform is
scanty. The forecourt A (Fig. 2) contains several looting holes and
some human bone fragments. This area seems to have been a true
cemetery, but it is unclear whether it is functionally connected with
the platform or a later intrusion. If the forecourt cemetery is function-
ally connected with the elevated platform, then it probably contains
persons of a status different from that of the women interred in the
platform, perhaps more comparable to the status of the women buried
in the entry court.

In sidecourt B (Fig. 2) hundreds of whole or nearly whole camelid
bones, probably llama {Lama glama), are present on the looted sur-
face. Their nearly whole conditions suggests they were looted from
tombs and not food residues. Excavations failed to reveal any intact
tombs, but because there are so many bones, it seems likely that the
llamas were sacrificed in the same ceremonies or in similar ceremonies

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Annals of Carnegie Museum

VOL. 48

associated with the women in the elevated platform. Outside of side-
court B, camelid remains were very few and scattered.

Conclusions

The excavations at Las Avispas in 1970 clearly showed that it func-
tioned as a burial structure for an important individual accompanied
by dozens of sacrificed young females. This is evidenced by 1) the
arrangement of chambers, with a uniquely large, T-shaped chamber
centrally placed among several smaller rectangular ones; 2) the ex-
traordinary quantity of young females’ bones recovered from the
chambers; 3) the large amount of fine, though fragmentary, grave
goods recovered and the heavy degree of looting associated with the
platform. Though the bones of the hypothetical ruler (and presumably
the only male) were not encountered, it should also be noted that
looting has been most concentrated in the central T-shaped chamber,
exactly where valuable grave goods would have been placed in direct
association with an important individual buried there.

Data from the Las Avispas excavations provided valuable architec-
tural and functional information for a broad survey of burial platforms
at Chan Chan and within the Moche Valley as well as in the Chicama
and Jequetepeque Valleys (Conrad, 1974, 1977, 1979). This is not to
say that every huaca on the Peruvian coast is a burial platform, but
only that a number of them meet certain criteria established by exca-
vation and survey.

Acknowledgments

Appreciation is extended to the National Science Foundation and the National Geo-
graphic Society, which funded the Chan Chan-Moche Valley project under which the
data for this paper were gathered, and to Michael E. Moseley and Carol Mackey who
directed that project. Thanks are also given to the Instituto Nacional de Cultura of
Lima which granted permission for the excavations. William Howells deserves my grat-
itude for his aid in the bone analysis. Appreciation is due Geoff Conrad who did the
original field drawings of chamber 19 and to Eduardo Martell and Shelia Pozorski who
did the final inkings of the illustrations. Fig. 1 is redrafted from Conrad (1979). Critical
comments on this paper by Geoff Conrad and Theresa Topic are also appreciated.

i

Literature Cited

CiEZA DE Leon, P. de. 1959. The Incas of Pedro de Cieza de Leon. Translated by
H. de Onis and edited by V. W. von Hagen. Univ. Oklahoma Press, Norman,
Oklahoma, 399 pp.

Conrad, G. W. \91A. Burial platforms and related structures on the north coast of
Peru: some social and political implications. Unpublished Ph.D. dissert., Harvard !
Univ., Cambridge, Massachusetts, 691 pp.

— . 1977. Chiquitoy Viejo: an Inca administrative center in the Chicama Valley,

Peru. J. Field Arch., 4: 1-18. ,

— . 1979. The burial platforms of Chan Chan. In Chan Chan; Andean desert city

1979

PozoRSKi — Las Avispas Burial Platform

137

(K. Day and M. E. Moseley, eds,), School of American Research, Santa Fe, New

Mexico, in press.

Donnan, C. B., and C. J. Mackey. 1978. Ancient burial patterns of the Moche Valley,
Peru. Univ. Texas Press, Austin, Texas, 412 pp.

Flecker, H. 1942. Time of appearance and fusion of ossification centers as observed
by roentgenographic methods. Amer. J. Roentgenology and Radium Therapy,

47:97--!59.

Garn, S. M., K. Koski, and A, B. Lewis. 1957. Problems in determining fossil and
modern man. Amer. J. Physical Anthro., 15:313--33L
Holstein, O. 1927. Chan Chan: Capital of the Great Chimu. Geog. Review., 17:
36-61.

Horkheimer, H. 1965. Identificacion y bibliografia de importantes sitios prehispanicos
del Peru. Arqueologicas 8, Museo Nacional de Antropologia, Pueblo Libre, Lima,
51 pp.

Hrdlicka, a. 1911. Some results of recent anthropological explorations in Peru.
Smithsonian Misc. Coll., 56:1-16.

Hutchinson, T. J. 1873. Two years in Peru with exploration of its antiquities. Samson,
Low, Marston, Lowe and Searle, London, 2 voL, 660 pp.

Klymyshyn, a. M. U. 1976. Intermediate architecture, Chan Chan, Peru. Unpub-
lished Ph.D. dissert.. Harvard Univ., Cambridge, Massachusetts, 1071 pp.
Kroeber, a. L. 1926. Archaeological explorations in Peru, part I: ancient pottery from
Trujillo. Field Mus. Nat. Hist., Anthro. Mem., 2:1-43.

McKern, T. W., and T. D. Stewart. 1957. Skeletal age changes in young American
males. Technical Report Ep-45, Project Reference AE7D, Environmental Protection
Research Division, U.S. Army, Natick, Massachusetts, 179 pp.

Peet, S. D. 1903. Ruined cities of Peru. Amer. Antiq., 25:151-174.

PozoRSKi, T. G. 1971. Survey and excavation of burial platforms at Chan Chan, Peru.

Unpublished A.B. thesis. Harvard Univ., Cambridge, Massachusetts, 192 pp.
Rodriguez Suy Suy, V. A. 1970. Chan Chan: ciudad de adobe, observaciones sobre
su base ecologica. Boletin del Museo de Sitio “Chavimochic,” Cooperative Agraria
de Produccion Cartavio Ltda. No. 39, Trujillo, Peru, 1:1-26.

Squier, E. G. 1877. Peru: incidents of travel and exploration in the land of the Incas.
Harper and Brothers, New York, 599 pp.

Stevenson, P. 1924. Age order of epiphyseal union in man. Amer. J. Physical Anthro.,

old ser., 7:53-93.

SuK, V. 1919. Eruption and decay of permanent teeth in whites and negroes with
comparative remarks on other races. Amer. J. Physical Anthro., old ser., 2:351-
400.

Topic, J. 1977. The lower class at Chan Chan: a qualitative approach. Unpublished
Ph.D, dissert.. Harvard Univ., Cambridge Massachusetts, 748 pp.

Tschudi, j. j. von, and M. E. Rivero. 1855. Peruvian Antiquities. Translated by F.
C. Hawks. A. S. Barnes and Company, New York, 306 pp.

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52? 7, 73

ISSN 0097-4463

ANNALS

of CARNEGIE MUSEUM

CARNEGIE MUSEUM OF NATURAL HISTORY
I 4400 FORBES AVENUE ^ PITTSBURGH, PENNSYLVANIA 15213

i VOLUME 48 1 June 1979 ARTICLE 9

1 LATE PREHISTORIC LLAMA REMAINS FROM THE

MOCHE VALLEY, PERU

Shelia Pozorski

Research Associate, Section of Man

} Abstract

A study focusing on subsistence through time and space within the Moche Valley
[ resulted in large collections of llama bone from sites dating to the Early Intermediate
j Period, Middle Horizon, and the Late Intermediate Period. Using contextual information
available, distributions of specific bone types are evaluated both within and between
sites studied. Cultural alteration of bone fragments is examined. Burned bone propor-
tions are correlated with cooking practices. Specific information on butchering practices
in the form of cut marks is explored in detail in an effort to reconstruct ancient methods
and procedures for meat processing.

Introduction

During the period between 1 October 1973 and 1 May 1974 exca-
vations were carried out at nine archaeological sites in the Moche
Valley, Peru. These excavations were designed to gather information
on subsistence from sites of different time periods and different loca-
tions within the valley (Pozorski, 1976). Abundant camelid remains,
probably llama {Lama glama), were present only at the sites of Moche,

I Galindo, and Chan Chan. All three sites are relatively late in the Moche
Valley sequence; one is Early Intermediate Period, one Middle Hori-
zon, and one Late Intermediate Period. The spatial distribution of
these sites varies, with Chan Chan very near the coast, Moche inland
near the river, and Galindo well upvalley.

Submitted 23 November 1978.

139

140

Annals of Carnegie Museum

VOL. 48

Sites Excavated

Moche

The Early Intermediate Period site of Moche, or the Moche Huacas,
covers more than 4 square km on the south bank of the Moche River,
about 5 km from the ocean (Fig. 1). Architectural components present
at the site include two very large corporate labor adobe mounds, an
adobe burial platform, and both high and low status domestic adobe
architecture. The refuse sampled at the site dates to a time after the
main mounds fell into a state of disrepair. A detailed sedation of
Moche Ceramics makes it possible to place this later occupation late
in phase IV of five phases, or near the end of the chronological span
of A.D. 300 to 600. During these late Moche times, small crude struc-
tures were fashioned on top of the larger pyramid, Huaca del Sol,
using bricks robbed from the mound surface. Depressions created by
this reuse of adobes became filled with refuse. The stratigraphic ex-
cavation of one such depression provided material for this study.

Galindo

The Middle Horizon site of Galindo is furthest inland, some 20 km
upvalley from the coast (Fig. 1). It lies on the slopes of two rugged
hills on the north side of the valley. Its location just outside the modern
limits of cultivation has resulted in good architectural preservation
over most of its 4 square km area. Formal architecture present ranges
from a walled compound with a central mound to a massive stone
and adobe wall encircling one hill and serving to reinforce social dis-
tinctions. Both high and low status dwellings are present in the areas
of domestic architecture. Ceramically, the site has been identified as
phase V of the Moche phases and placed chronologically in the early
part of the Middle Horizon (A.D. 600-1000). No stratified midden de-
posits were encountered. Instead, refuse is concentrated within resi-
dential structures where a homogeneous 5 to 30 cm lens of ash is
present in the vicinity of hearths.

Chan Chan

The Late Intermediate Period (A.D. 1000-1470) site of Chan Chan
lies north of the Moche River with its southern edges bordering on the
ocean (Fig. 1). What remains of the site covers about 6 square km, and
within this area the architectural components vary from adobe or tapia
(tamped earth and rocks) walled compounds to small-scale cobble and
adobe domestic structures known as SIAR (Small Irregular Aggluti-
nated Rooms). Information gained by the Chan Chan-Moche Valley
Project suggests that the compounds served mainly as administrative
and redistributive centers while most of the population was housed in

1979

PozoRSKi — Llama Remains

141

Fig. 1. — Map of the Moche Valley showing the location of the sites of Chan Chan,

Galindo, and the Moche Huacas.

less elaborate structures such as the SIAR. These ideas are reinforced
by the nature and distribution of artifacts and refuse in the noncom-
pound areas. Substantial refuse deposits were almost exclusively con-
fined to areas of SIAR. Within these areas, stratified refuse deposits
were discovered 1) in filled alleyways and filled rooms interspersed
with the remaining architecture, 2) dumped in unused areas of the site
between sections of domestic architecture, and 3) as isolated middens
on the edge of the site. A series of four stratigraphic excavations in
the SIAR on the western edge of the site yielded some of the Late
Intermediate Period material used for this study. Supplemental faunal
material was provided by previous excavations by John Topic (1977)
in the two SIAR units which were stratigraphically sampled. These
additional data were used only in evaluations of age structure, bone
type frequencies, and butchering practices.

Field Methodology

The field methods used for collecting subsistence remains varied, depending on the
nature of the deposit being tested. Two kinds of refuse deposits were encountered during
this study. The more common form consists of a deep deposit within which it is possible
to distinguish a number of distinct strata. The sites of Moche and Chan Chan contain

142

Annals of Carnegie Museum

VOL. 48

GALINDO

CUT 2

Fig. 2.— Hearth area with surrounding ash within the multi-hearth structure excavated
at Galindo showing division into squares prior to Cut 2 excavation.

1979

PozoRSKi — Llama Remains

143

this stratified refuse, whereas Galindo, on the other hand, is one of only two sites where
the second refuse type was encountered. Instead of being deep and stratified, the refuse
at Galindo is confined to 5 to 25 cm homogeneous deposits which cover a considerable
area of floor surface near the cooking hearths.

At the sites of Moche and Chan Chan, stratigraphic excavation procedures reflected
efforts 1) to distinguish between the natural levels present and record them prior to
excavation, 2) to excavate a certain volume of material by removing each natural level
separately and keeping its contents distinct, and 3) to record the distribution of levels
after the excavation in order to account for changes in strata within the volume removed.

In the case of Galindo where no natural strata were visible, artificial quantitative
control was imposed in the form of a grid system which was followed during excavation
of the ash deposits. Once a hearth area with accompanying ash was detected, its entire
surface was cleaned down to the ash. The deposit was then sectioned off into 50 cm
squares which became the standard excavation units (Fig. 2).

Analysis

In addition to the differences based on deposit type and excavation techniques used
between Galindo on one hand and Moche and Chan Chan on the other, there are other
distinctions which considerably affect the analysis of faunal material from these sites.
These differences center around such factors as sample size and direct cultural context
for the bone material from each site.

Within the stratigraphic deposits, it is most reasonable at present to deal with the
llama remains from each whole cut as a unit by combining data from individual levels.
This is true for two reasons. First, since the ceramics do not change from level to level,
it is not valid to assume that each natural stratum is an isolated unit with respect to the
animal bones it contains. Second, the sample of llama bone is quite small, often resulting
in a minimum number of individual count for each cut which is well below the number
of strata. If the strata were assumed to be isolated units, then the presence of even a
small amount of llama bone would necessarily be counted as one individual, yielding a
count much higher than that calculated for the cut as a whole. Grayson (1973) discusses
such an abuse of minimum number calculations. Certainly, the number of animals con-
sumed at the site far exceeded either of these estimates. However, counts based on the
cut taken as a whole are more realistic and constitute a sounder basis for further inter-
pretations and comparisons (Pozorski, 1976).

At Moche and especially Chan Chan, where refuse is abundant, the amount actually
sampled was quite small. In each case, only a fraction of the total refuse in a filled alley,
a midden, or other deposit was removed. Also significant is the fact that the refuse
sampled can be only generally correlated with specific living areas. Least specific is the
refuse at Moche where one can be sure only that it is associated with the smaller
structures built on top of the mound. At Chan Chan, refuse within alleys and filled
rooms is probably derived from nearby living areas while refuse in the midden at the
edge of the site could have come from a larger portion of the site. However, all is clearly
associated with SIAR occupation.

At Galindo, the situation is unique. Each refuse deposit, usually near a hearth within
a room, is clearly associated with a single domestic structure and therefore with the
activities which took place within the structure. Thus, a complete excavation of the
material surrounding such a hearth yields a relatively more complete sample than was
the case for Chan Chan or Moche. This makes possible at Galindo inquiries of a type
not feasible for the other two sites. The specific locations of bones can be evaluated to
determine if a pattern of distribution exists within the room. To accomplish this, material
from each artificial square must be considered separately. Yet for comparisons between
rooms and with other sites, the refuse for each room is necessarily considered as a unit

144

Annals of Carnegie Museum

VOL. 48

Table l.Sample size of llama bone including minimum number of individuals , bone(s)
most frequently represented, total identifiable bone, and number of bones per individual
for the sites of Moche, Galindo, and Chan Chan arranged according to cut.

Site

Minimum

number

Most

frequent

bone

Total

identifiable

bone

Bones

per

animal

Moche

5

Femur

Tibia

585

117

Galindo

Cut 1

3

Radius-ulna

84

28

Cut 2

5

Femur

160

32

Cut 3

2

Femur

56

28

Cut 4

3

Left temporal

222

74

Cut 5

2

Femur

82

41

Cut 6

4

Humerus

340

85

Cut?

2

Radius-ulna

48

24

Chan Chan

Unit 1

Cut 1

1

All equal

42

42

Cut 2

2

Femur

128

64

General*

4

Humerus

60

15

Unit 2

Cut 3

2

Left calcaneum
Right scapula

78

39

Cut 4

2

Radius-ulna

56

28

General*

12

Right femur

684

57

’ Material from previous excavations within the unit. Bones per animal is occasionally
low because shaft fragments were often discarded.

because the deposit is generally homogeneous and the division into squares purely an
artificial device.

Bone Identification and Description

The procedure followed for the identification of specific bones was the same for all
sites. Camelid remains were identified as llama {Lama glama) on the basis of compar-
isons with a complete skeleton in the Lima Natural History Museum. Bones from each
context (stratum or 50 cm square) were treated as a unit and the same analytical pro-
cedures were applied uniformly. As each bag of material was examined, the diagnostic
fragments were identified. Other characteristics of each bone were recorded. If the bone
was burned, this was noted, and it was separated from the remainder of the sample.
Bones with unfused or partially fused portions such as epiphyses were described with
respect to the degree of fusion and the presence or absence of unfused portions. Finally,
all identifiable fragments were carefully checked for evidence of cultural alteration.
Virtually all long bone shafts were very fragmented, and cut marks were detected on
quite a few bones.

Quantitative Analysis

Most persons evaluating faunal samples have dealt with the problem of quantifying
bone material. Early species lists (see Baker, 1930, 1931, 1936), even elaborate ones.

1979

PozoRSKi — Llama Remains

145

are now out of date and current emphasis is on discovering the best indicator of the
true former sample size. Attempts range from counting fragments to weighing fragments
to a determination of the minimum number of individuals. Most investigators favor the
“minimum number” method which, in simplest form, involves a count of the most
common bone from each animal to ascertain the numbers of each animal actually rep-
resented by the faunal material (see for example Clark, 1971; Daly, 1969; Degerbpl,
1929; Hill, 1966; Jope, 1954; Munson et al., 1971; Parmalee, 1962, 1963, 1965; Parmalee
et al., 1972; White, 1952, 1953a, 1953^, 1955). More elaborate methods have been pro-
posed which involve adding variables such as size or age to that of bone types (Bokonyi,
1970) or a preliminary pairing of the bones (Chaplin, 1971). The minimum number meth-
od has been criticized as over representing rarer species because one bone can equal
one animal or 100 unduplicated bones can equal one animal. Shotwell (1955) and Perkins
(1964) have controlled this variable somewhat by calculating the relative completeness
of each animal. The value obtained is an expression of the average number of bones
present per individual animal.

Counts of individual bones or bone fragments for the Moche Valley sample were
possible using the descriptive lists made at the time of identification. The first quanti-
tative procedure was the establishment of a minimum number of individuals for each
cut. The method used to determine this minimum number of individuals for the Moche
Valley material was slightly more sophisticated than the simplest procedure, but cer-
tainly not so complex as the method proposed by Bokonyi (1970). First, for each cut,
the bone(s) occurring most frequently was determined. Then the age (fused versus un-
fused) groupings for each of the most common bones were considered. This often served
to raise the minimum number slightly. Finally, an evaluation of the completeness of the
animals was carried out using the formula supplied by Perkins (1964). A summary of
this information is presented in Table 1.

In a more detailed study of the Moche Valley subsistence (Pozorski, 1976), the quan-
titative analysis was taken a step further to obtain values for the amount of meat rep-
resented by each bone or bone fragment for all species for comparative purposes. How-
ever, such data are not relevant in a consideration of only camelid remains and are
therefore not included here.

Interpretation

Interpretations of faunal material are largely dependent on two fac-
tors—sample size and sample context. Many researchers (see for ex-
ample, Uerpmann, 1973, and Krautz, 1968) have considered the effect
on a sample studied of bone loss due to factors such as scattering by
the butchers, special butchering or consumption techniques, carni-
vores removing bones, manufacture into tools, and differential weath-
ering. The volume of material removed and processed by the excavator
affects the sample even more significantly. It is usually difficult to
evaluate the size and significance of a given sample while still in the
field, thus many interpretations become highly qualified.

Information on the context of faunal material is vital on two levels.
On the more specific level, contextual information is directly depen-
dent on the excavator’s skill in first knowing how much data to record
about the situation in which bones are found and, second, recording
such data in a useful form. Examples of time well spent recovering
detailed contextual data are the sites of Olsen-Chubbuck excavated by
Wheat (1967, 1972), and Glenrock, Casper, and Hawken excavated by

146

Annals of Carnegie Museum

VOL. 48

Prison (1970, 1974, 1976). Most of their conclusions would not have
been possible had the exact context of faunal material gone unrecord-
ed. At the other extreme, no contextual information is recorded, and
the investigator is presented with an undifferentiated collection of all
the bones from a site. This precludes any interpretation based on even
the general distribution of bone at a site.

On a general level, context refers to the faunal material within the
cultural framework known for the site. In many cases only general
contextual information is available. However, when much more is
known about factors such as social and political organization through
additional excavation or ethnohistorical data, more far-reaching inter-
pretations can occasionally be made.

On a specific level, context has proven most significant at Galindo
where faunal material was taken directly from food preparation areas.
On a general level, however, much is known about the sites of Galindo,
Moche, and Chan Chan, and this information should be helpful in the
final interpretations.

Bone Distribution within the Site

Galindo is the site in which specific contextual data have proven
most significant. General bone distribution was recorded through the
use of arbitrary 50 cm squares. The distribution of specific bone types
was plotted, based on the inventory for each square, but no specific
conclusions could be drawn. When the relative abundance of bone
fragments of all body parts was considered, two basic patterns
emerged. At all times the bones were confined to the floor area where
ash was present, but in most cases the distribution of bone within this
ash was patchy, with concentrations of bone discernible, but not di-
rectly within hearth areas. The second pattern reveals a concentration
of bone fragments in the immediate vicinity of the hearth.

The differences between the two patterns can be interpreted as an
indication of hearth-cleaning practices. The confinement of the bone
to ash deposits is significant here. The predominance of the patchy
pattern of scattered concentrations could be the result of haphazard
scraping of residue and ashes from the hearth area in order to rebuild
the fire. This could scatter both bones and ashes, with the heavier
bones frequently remaining in concentrations. Hearths not cleaned
near the time of abandonment would contain concentrations of burned
bone.

Burned versus Unburned Bone

In order to quantify the frequency of burning for different bones,
comparisons were expressed in percentages of burned bones of the
total bone of a given type (Table 2). In dealing with burned bone,

Table 2. — Frequency {in parentheses) of burned llama bone compared to unburned bone by body part for the sites of Moche, Galindo,

1979

PozoRSKi — Llama Remains

a

IS

q — ; q q r-;

q

wo m

q q

y o

Q-i

o

wo fo 0© 00

ro NO wo r" (N

ON 00
ro

NO WO

q q q ^ 00 q

00 oq

d ^

Shaft

e

yF

m

r4 fNi « NO 00 wo
CN wo 00 CN

©

fd NO

w w

d oc

wo ^

1

2

yr\

fN O

wo o wo

ro wo ^ ^

NO q
ro NO

o

wo r4

00 ^ ^ ^ O
(N — ' wo

o

wo d
m

wo r4
r4

15 S

c ©

q,

ye\

w

S' X S' ^

ro ri © o o
wo r- o NO

(rii)

(20.0)

(20.0)

(50.0)

(0)

a

£

Ph

TO

<U

E

ro <N wo ^

—I r4 NO eN wo (N

q

q q

r4 q

ON

^ ^ ^ r5 o

»— 1 T— 1

— d

q

o' (o' w7 o' o' NO

fd

Ofi

o

wo m Tf" O ©

O'! NO wo NO

d d

ro O
m ^

a

X

o

wo

fN wo CM

00 04 1 — ^ wo C-J

q

m WN

NO r4

io

r5 ^ ^ ^ ^

d 04

r4 d

t:

>>

^N© q q q

o'

q q

m

o

OQ

M

NO 00 o' O wo o
^ wo ^ ^

d

d wo

O M

d rd

eo

o

X

(N

wo

f«0 ON (N

C C! C ^ ^

w

wo

C ?5

r- m

Os

^ o rJ ro ^

04

r4

d ^

(N

q q q NO q w7

oq

q

Rib

ON

ro

ON

o © o ON 00 cT

wo L«- o r4 00 Tf

o r- O

NO ' r^i ri ON

r4

q

d

^ f<0

04 ON
— ' f4

dd

f<o q

fO

r3 ^ 00

do

© d;

d d

q

wo fT o' o w7 o

cT

d d

CS

q6

w

od r4 wo (N o

04 r4 NO wo 04

d d

’—1 ro

dd

4>

>

i

00 ON o

oo r4 — 1

q

q q

NO as

oo

rN

04 ro r4 00 ^ 00

j— <

ro

d d

ro

q q ^ NO q q

d d

d ^

NO 0© o' fd NO ON
NO 00 ^ NO NO — <

[0)

© NO

q q

d d

3.

m

m

r4

ro q O ^ m r4

q q

wo q

04 00 r4 ^

d

r4 d

1 04 fO wo NO

r-

c

cd

^ r4

cn

Site

Moche

Galindo

p "S "p 3 3 3

u u u u u u

3

u

-C

U

G

cd

U

Unit 1
Cut
Cut

Unit 2

3 3
U U

147

148

Annals of Carnegie Museum

VOL. 48

Table 3.~Relative age stages for progressive epiphyseal fusion in camelids.

Age

stages

Epiphysis

fused

0

Astragalus

1

Proximal scapula

2

Proximal metacarpal

Proximal metatarsal

3

Distal humerus

Distal tibia

4

Distal radius-ulna

5

Calcaneum

6

Distal metacarpal

Distal metatarsal

7

Distal femur

Deciduous premolars 2 and 3 replaced and 3rd molar erupting,
36 months (Cardoza, 1954:91)

8

Proximal femur

Proximal tibia

Proximal humerus

Proximal radius-ulna
(after Wing, 1972:330, Table 2)

elements have been grouped according to body section. For example,
the scapula, humerus, radius-ulna, and carpals are included under the
heading foreleg. This procedure provides larger, more significant sam-
ples. Nevertheless, several cuts yielded samples which are often too
small to evaluate.

A consideration of Table 2 reveals immediately the consistent high
incidence of burned bone for most cuts at Galindo. This is certainly
due largely to the context of the refuse because all of the Galindo
material was taken from hearth areas, whereas Chan Chan and Moche
samples came from middens and filled rooms or pits. It is interesting
to note that within the site of Galindo, Cuts 6 and 7 have the least
burned material. Based on studies of ceramics and other artifacts from
these structures, it is known that they housed individuals of much
higher status than those dwellings sampled by Cuts 1 through 4. How
this could relate to the percentage of burned bone is subject to spec-
ulation. Perhaps relatively little actual food preparation was done
there.

The high percentage of burned bones for some cuts at Moche and
Galindo may also reflect cooking practices. These bones may have
become charred during roasting rather than as a result of later burning
of debris in the hearth. Certainly much of the Galindo material was

1979

PozoRSKi— Llama Remains

149

Fig. 3. — Percentage of bone for each fusion stage for samples collected at Moche, Gal-
indo, and Chan Chan.

accidentally burned, and it is difficult to comment on material from
Moche because no hearths were found. Chan Chan, on the other hand,
shows a very low incidence of charred meat-bearing bones. This might
suggest that more cooking was done in containers. This correlates well
with both frequent finds of fire-blackened vessels and the nature of the
Chan Chan hearths — ^deep oval pits containing only vegetal fuel resi-
dues.

Age of Animals Consumed

An evaluation of the age of llamas for each site was made on the
basis of whether each bone was fused or unfused. All teeth were frag-
mentary and rarely had remained in the sockets, therefore, this criteria
could not be applied. Wing (1972:330, Table 2) has determined a series
of camelid age stages based on epiphyseal fusion. This information is
reproduced as Table 3. The sixth stage is correlated with a teeth erup-
tion schedule established by Cardoza (1954) to give a chronological
age equivalent of 36 months. However, all other stages are relative
and have not yet been tied to chronological ages.

Bone data from Moche, Galindo, and Chan Chan were organized
according to the stages, and the results were graphed (Fig. 3). Each
graph records the percentage of unfused bones at a given stage. The

150

Annals of Carnegie Museum

VOL. 48

age structures for Chan Chan and Galindo are quite similar, but both
differ considerably from Moche results. At Moche two aspects are
significant. Generally a greater proportion of animals are young as
indicated by the larger unfused percentages of most bones in the
Moche sample. And within this group, the animals are younger than
animals from Chan Chan or Galindo. This is evident from the per-
centages for fusion stages 3-5 for Moche compared to the two later
sites. Although the Galindo and Chan Chan samples contain more
older animals than Moche, it is significant that even in the later fusion
stages about half the bones are not fused.

According to Uerpmann (1973) and to some extent Perkins and Daly
(1968) a methodical exploitation of a specific age group is expected
among peoples who keep herds of domesticated animals for specific
purposes. In the case of the Moche Valley sample, the evidence in-
dicates that herds were kept primarily for meat. In such a situation,
after individuals are selected for breeding stock, it is most efficient to
butcher meat animals soon after they reach adult size. At this point
they no longer gain substantial weight, but do consume what must
have been extremely precious fodder. The high proportion of young
and young- adult animals in all our samples fits this situation well. The
slightly older age structure for Chan Chan and Galindo may reflect an
increased use of these herd animals for wool and as beasts of burden.

Frequencies of Bone Types

Quantitative evaluations of the distribution of various bones were
made in order to investigate aspects such as butchering practices and
associated cultural attitudes and behaviors. Identifiable meat-bearing
bones are evaluated separately with quantities expressed as percent-
ages of the total meat-bearing bones from each cut (Table 4). The
breakdown by bones correlates with divisions confirmed by butchering
marks. Foot bone (metapodials and phalanges) frequencies are ex-
pressed as a percentage of the total number of identifiable bones,
whereas shaft fragments are considered with respect to the grand total
of all bone elements recovered (Table 5).

Galindo provides a unique opportunity for evaluating the distribution
of meat-bearing bones with respect to specific structures. Certainly,
many samples are quite small. However, each represents the total
camelid inventory directly associated with a given hearth and therefore
is the most nearly complete sample available.

Data from Cuts 2, 3, and 4 come from three separate food prepa-
ration areas within a single complex living structure (Fig. 4). For Cut
2, the sample contains a large proportion of foreleg (or front body)
parts, especially the radius-ulna and the scapula as well as a moderate
amount of hind limb parts. This suggests that the shoulder girdle and

Tabie 4. — Frequency of meat-bearing llama bones relative to total sample from each of the sites of Moche, Galindo, and Chan Chan.

POZORSKI-

—Llama Remains

e «

m

NO ON

NO

fNI

G\

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m

fNI

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(Nl

CN

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©

0

© ©

0

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0

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0

0

0

©

©

cU

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H

§

1

SB

8

0

©

8

1

8

8

I

8

8

ca

0

in

q q

in

in

q

q

q

in

IB

r-'

m

r-; ^

©

0

r4

©

00

fN

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in in

in

q

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— <N

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1 =

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rl

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q

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si

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d

00

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©

'S

ON

2

2

q q

q

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cn ^

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00

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u >

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<-4

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+->

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0)

.S

U

U U

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0)

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p

p

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X

a

U

UJ

151

152

Annals of Carnegie Museum

VOL. 48

Table 5.— Frequency of llama foot bones and shaft fragments present and expected
from the sites of Moche, Galindo and Chan Chan.

Metapodial

Phalange 1

Phalange 2

Phalange 3

Shaft

Site

No.

Percentage

No.

Percent-

age

No.

Percent-

age

No.

Percent-

age

No.

Percentage

Moche

5

3.6'

10

7.3

6

4.4

2

1.4

245

41.9^

Galindo

Cut 1

4

14.8

3

11.1

2

7.4

0

0

35

41.1

Cut 2

2

3.0

6

9.2

1

1.5

5

7.6

50

31.2

Cut 3

2

11.7

1

5.8

1

5.8

0

0

27

49.0

Cut 4

4

8.6

5

10.8

3

6.5

3

6.5

112

50.0

Cut 5

1

6.2

1

6.2

1

6.2

0

0

45

54.8

Cut 6

6

7.5

6

7.5

6

6.2

0

0

170

50.1

Cut?

0

0

3

17.6

3

17.6

0

0

6

12.7

Chan Chan

Unit 1

Cut 1

0

0

2

18.1

2

18.1

0

0

36

52.9

Cut 2

0

0

1

4.0

2

8.0

0

0

64

50.3

General

5

10.2

10

20.4

2

4.0

0

0

—

Unit 2

Cut 3

2

18.1

1

9.0

1

9.0

0

0

50

63.2

Cut 4

2

10.0

0

0

0

0

0

0

24

43.6

General

49

13.6

16

4.4

6

1.6

0

0

—

Expected

8=*

6.6

8

6.6

8

6.6

8

6.6

—

* Percent of total number of identifiable bones.

^ Percent of total number of bones: fragments + identifiable bones.

^ Because of the fragmentary nature of most bones, proximal and distal metapodials
were counted separately, hence the double expected number.

foreleg was the meat unit most often consumed in this area of the
complex. Cut 3 data indicate an emphasis on hind limb parts (femur,
tibia, and innominate) and a lesser consumption of vertebrae. Foreleg
parts are conspicuously rare. At the site of Cut 4, evidence points to
a heavier reliance on meat from body parts in the skull-vertebra-im
nominate region as well as some hind limb sections.

Two aspects of the meat distribution data within this complex are
especially significant. First, the general body areas most highly rep-
resented at each hearth are different and to some extent mutually ex-
clusive. Second, meat elements, as represented by bones, are seen to
cluster in units which would have been meaningful in light of butch-
ering procedures. Such clustered units include the forelimb-shoulder
girdle, the hind limb-pelvic girdle, and the skull- vertebra-innominate
section. The systematic meat distribution system seen here may reflect
divisions such as those based on the established hierarchy within an
extended family occupying the complex structure.

STRUCTURE

Fig, 4. Plan of multi-hearth dwelling at Galindo showing locations of Cuts 2, 3, and 4.

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VOL. 48

Other data from Galindo come from structures with a single hearth.
Evidence from Cut 1 suggests a reliance on the forelimb almost to the
exclusion of other body parts. Again for Cut 5, one body portion is
very well represented. In this case it is the neck and shoulder girdle
region. Cut 6 reveals a hearth area where emphasis was again on for-
ward body parts due to the high frequency of skull, thoracic vertebra,
and humerus portions. However, these units are not so easily corre-
lated with butchering divisions. Finally, data from Cut 7 reveal a con-
siderable emphasis on the hind limb and pelvic area, but the high
frequency of an element such as the radius-ulna suggests that substan-
tial amounts of meat came from other body portions as well.

Data from single-hearth structures at Galindo indicate that a meat
distribution system is operating on this level as well. As mentioned
above, Cuts 5, 6, and 7 were located in structures containing high-
status artifacts. Perhaps the variety of meat cuts consumed by the
inhabitants is correlated with this higher status.

At Chan Chan refuse contextual information is less precise, but bone
distribution can be evaluated generally with respect to SIAR areas.
When the sample from the first unit is examined, elements of the for-
ward body portion predominate with hind limb elements secondary.
Skull and vertebral portions are well represented as are forelimb seg-
ments. An examination of the second Chan Chan SIAR unit reveals
a uniform access to limb bone meat and a deemphasis on vertebral
segments. While the consistencies in meat use at Chan Chan are less
marked than was the case for Galindo, patterns are evident which
suggest systematic distribution.

When foot bone frequency is considered (Table 5), most elements
are seen to occur at roughly the expected frequency or in greater
numbers. Third phalanges are consistently rare or absent for all time
periods represented. Such a scarcity of toe bones may correlate with
skinning procedures during which the last digits remain with the skin.
First and second phalanges are also usually “lost” due to this process,
but they are well represented at all sites.

Metapodials and first and second phalanges occur in roughly the
expected numbers at Moche and in increased frequencies at Galindo
and Chan Chan. The presence of substantial numbers of metapodial
fragments reflects their usefulness as raw material for tool manufac-
ture. In fact, two proximal fragments from Moche and one from Gal-
indo are deeply grooved where the shaft was severed in an early stage
of tool manufacture (Fig. 9h“j). It is possible that the increased me-
tapodial frequencies seen at Galindo and Chan Chan represent the
remains of deliberate accumulations of these skeletal elements for tool
manufacture. High frequencies of first and second phalanges are un-
usual because there is no evidence of their use to make tools or in any

Fig. 5.— Mandible (a) , a skull fragment (b), and cervical vertebrae (c-~g) with cut marks.
Examples a and e-g are from Moche; b is from Galindo; and c and d are from Chan
Chan.

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VOL. 48

other manner. Most likely they are a by-product of metapodial collec-
tion.

Butchering Practices

Some faunal material from all the excavations was marked with
grooves or bashes made by prehistoric knives as the carcasses were
processed. Although there is no direct evidence from excavations in
the refuse, it seems most likely that these cuts were made using both
metal and stone knives. An axis from Chan Chan (Fig. 5d) had a stone
tool fragment embedded in one of the bash marks. Both the Moche
and Chimu peoples used a wide variety of metal utensils made from
a naturally occurring copper alloy.

Interpretations of meat processing techniques proposed here are
based almost exclusively on the direct evidence provided by actual
marks on bones (Table 6). When the cut bones from each cultural and
chronological context were examined, there were virtually no indica-
tions of butchering techniques that varied through time. The one pos-
sible exception is discussed below.

The most detailed studies to date on prehistoric butchering tech-
niques deal with the large wild North American mammals: bison, deer,
elk, and pronghorn antelope (see for example Prison, 1970, 1974, 1976;
Gilbert, 1969; Guilday et al., 1962; Hill, 1966; Parmalee, 1965; Par-
malee et al., 1972; Wheat, 1967, 1972; White, 1952, 1953a, 1953^, 1955;
Wood, 1968). Three such studies based in North America have differ-
entiated two functional types of cut marks— marks due to skinning and
marks resulting from butchering (Guilday et al., 1962; Parmalee, 1965;
Parmalee et al., 1972). In this paper marks are further distinguished
functionally depending on whether they were made during the process
of skinning, disarticulation of the skeleton, or removal of the meat
from the bone.

Skinning marks are the most rare type in the Moche Valley sample.
Parmalee (1965) found skinning marks on deer skulls around antler
bases and along the jaw at the gum line as well as on the extremities
encircling foot and lower leg bones. My evidence for skinning is from
one mandible as well as lower leg and foot bones, and several of these
cuts may correlate with disarticulation activities as well as the removal
of the pelt from the legs. The most reliable evidences for skinning are
marks on a mandible and four phalanges (three first and one second)
(Fig. 13g-j) and on occasional metapodial shafts. Marks are also pres-
ent on a number of carpal and tarsal bones (Fig. 9f™g; Fig. Oa^f), but
these are more likely to have been associated with the disarticulation
procedures.

After skinning was begun (and possibly completed) the carcass was
disarticulated— usually by cutting through the joints and associated

1979

PozoRSKi — Llama Remains

157

Table 6— Distribution of cut marks for bone disarticulation for samples from the sites
of Moche, Galindo, and Chan Chan.

Joint

Moche

Galindo

Chan Chan

Skull-

0

1

0

atlas, axis

0

0

5

Cervical vertebra

8

3

7

Thoracic vertebra

2

1

8

Lumbar vertebra

2

0

6

Sacrum

0

1

2

Thoracic vertebra-

0

1

3

rib

8

1

12

Scapula-

1

0

2

humerus

1

0

1

Humerus-

0

3

3

radius-ulna

1

4

4

Radius-ulna-

1

1

0

carpal-

2

1

0

metacarpal

1

0

0

Innominate

3

0

2

Innominate-

0

0

4

femur

0

2

10

Femur-

0

1

3

patella-

0

1

0

tibia

1

1

3

Tibia-

0

0

3

tarsal-

1

4

6

metatarsal

1

0

1

tendons and ligaments (Table 6). A single mandible provides possible
evidence of cutting the jaw away from the skull (Fig. 5a). The foreleg
was cut away from the scapula (Fig. 8d, h) and also cut apart at the
humerus-radiuS"Ulna joint (Fig. 8g; Fig. 9a--b, d-e). Cuts on carpal
bones (Fig. 9T-g) were discussed as possibly due to skinning, but prob-
ably reflect efforts to disarticulate the radius-ulna from the metacarpal
which was also occasionally marked.

Data on the disarticulation of the hind leg provide the only apparent
I indications of chronological differences in butchering techniques. In
the Moche sample, two cases of cut marks on the pelvis suggest that
the hind quarter was separated from the body by cutting through thin-
ner portions of the innominate near the acetabulum (Fig. 11b, d). This
is reinforced by the absence of cuts on proximal femora. A comparable
procedure was described by Guilday et al. (1962:74) for deer. Marks

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VOL. 48

Fig. 6.— Cervical vertebrae (a-g) with cut marks. Examples a-c are from Moche; d and
e are from Galindo; and f and g are from Chan Chan.

1979

PozoRSKi— Llama Remains

159

Fig. 7.— Cervical (a-b), thoracic (c-g), and lumbar (h) vertebrae with cut marks. Ex-
amples c and h are from Moche; d and e are from Galindo; and a-b and f-g are from
Chan Chan.

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VOL. 48

Fig. 8.— Lumbar vertebrae (a-b), scapulae (c-f), and humeri (g=h) with cut marks.
Examples a and c-e are from Moche; f and g are from Galindo; and b and h are from
Chan Chan.

1979

PozoRSKi — Llama Remains

161

on an ilium (Fig. 11c) document disarticulation of the innominate from
the sacrum. The later Moche sample from Galindo, though small, con-
tains two proximal femora with cut marks probably resulting from
disarticulation efforts (Fig. 12a-b). In the case of Chimu material from
Chan Chan, 10 proximal femur heads exhibit cut marks (Fig. 12c=-d)
while the only cuts (4) affecting the pelvic area are on or very near the
acetabulum.

The hind leg was also severed at the knee (Fig. 12e-f). Cut marks
in this region are on both the anterior and posterior surfaces of the
bones— suggesting repeated efforts from different directions until the
joint was located and cut through. Further disarticulation of the hind
limb at the hock joint is suggested by cut marks on an occasional tibia,
numerous tarsals, especially astragali, and less frequently on proximal
metatarsals (Fig. Ba^f).

Separation of various parts of the spinal column seems least me-
thodical. Most marks on vertebrae are on the vertebral arches, espe-
cially in the overlapping areas where these portions articulate (Fig. 5e--
g; Fig. 6a--g; Fig. 7a-c; Fig. 8b). All classes of vertebrae had been cut.
Of the 42 vertebrae with marks, 23 are cervical, 11 thoracic, and eight
lumbar. In all classes, marks occurred on both dorsal and ventral sur-
faces—again suggesting multidirectional efforts to locate the joint
which was then cut. The relatively high frequency of cut marks on
vertebrae is certainly due to the physically overlapping arrangement
of the joints. There is some indication that the neck was consistently
severed at the same point (Fig. 5b~d). Five marks were noted on the
atlas, one on the axis; and one of the several basioccipitals encoun-
tered is marked. Most evidence for the vertebrae reflects the dividing
of the spinal column into convenient lengths for transport or cooking.

Occasional ribs have marks near the end (Fig. 10k) which probably
resulted from cuts near the sternum to open the chest cavity. Many
more (21) have cuts near the heads which are the result of their sep-
aration from the thoracic spinal column (Fig. lOa-j, 1). Occasional (3)
cuts near the rib articulation areas of thoracic vertebrae also reflect
this disarticulation procedure (Fig. 7d, g). Although cuts were discov-
ered on all surfaces near the rib head, most were found to be distrib-
uted cranially and ventrally.

Little can be said about the order of butchering procedures, espe-
cially limb disarticulation. Our only data on this aspect come from the
location of marks on some ribs and vertebrae. The presence of cut
marks on the ventral surfaces of both the ribs and thoracic vertebrae
indicates that the body cavity had been split and the internal organs
completely removed before at least this phase of butchering was be-
gun. Also, this procedure indicates that a considerable portion of the
operation was executed with the animal on its back. This differs from

Fig. 9. — Radius-ulnae (a-e), carpal bones (f-g), and metacarpals (h-k) with cut marks.
Examples a, f, h, i, and k are from Moche; and b-e, g, and j are from Galindo.

1979

PozoRSKi — Llama Remains

163

Fig. 10. — Ribs (a-l) with cut marks. Examples a-h are from Moche; i is from Galindo:
and j4 are from Chan Chan.

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Annals of Carnegie Museum

VOL. 48

Fig. 1 1. — Innominate segments (a-d) and sacra (e-f) with cut marks. Examples a-d are
from Moche; e is from Galindo, and f is from Chan Chan.

1979

PozoRSKi — Llama Remains

165

the methods described by investigators such as White (1953^) for
North American Indians. These butchers apparently placed the animal
on its belly and slit the skin down the back. Back meat was then
removed to expose the ribs which were cut through near the spine to
reach the internal organs.

Cut marks near articular surfaces seem to be of two general types.
One is a narrow slit, often repeated for several strokes, that is com
centrated in the area of the ribs and vertebrae, but also occurs fre-
quently on the long bones. The second, less common, type more re-
sembles a bash, as if the action involved were more a chopping than
cutting motion. These marks are wider and often in the form of dents.
One set was discovered on an atlas (Fig. 5d) and another on the sacrum
(Fig. lie), but the remainder occur on long bones (see Fig. 12f). Often
they seem to be associated with cuts through bone or fractures of the
bone. These differences in cut types may reflect the use of different
tools in one or both of two ways. Depending on the nature of the joints
(vertebra versus femur-tibia, for example) fine or forceful cutting pro-
cedures might be necessary to sever the articular connection. It is also
possible that the two tool types and their respective cuts correlate with
first a generalized preliminary or centralized butchering procedure fol-
lowed by the use of finer tools on a local or individual level to prepare
meat for cooking. Bash marks on shafts certainly correlate with mar-
row extraction procedures.

The final type of cut marks— -marks reflecting removal of meat from
bones— may also be correlated with preparation for cooking. This
function is attributed to marks in locations that do not reflect either
disarticulation or skinning efforts, but which are in known areas of
muscle attachment (Lesbre, 1903). Marks on three vertebrae and part
of the sacrum are probably a result of removing meat from the spinal
column (Fig. 7f, h; Fig. 8a; Fig. Ilf). A thoracic vertebra has lateral
cuts at the base of the caudal spine (Fig. If). One lumbar vertebra is
marked above and below the left lateral spine while another has cut
marks laterally along both sides of the body (Fig. 8a). Marks on the
dorsal portion of two sacral segments also reflect meat removal (Fig.
Ilf). The removal of meat from the back is indicated by dorsally po-
sitioned marks on scapula blades just on the caudal side of the spine
(Fig. 8e) and on the spine (Fig. 8c) and ventrally located marks on
another scapula (Fig 8f). Finally, a transverse cut on the ilium probably
served to free some of the thick pelvic muscles (Fig. 11a).

The final aspect of butchering— marrow extraction— is reflected in
the nature and quantity of bone fragments as well as bash marks on
long bone shafts (Fig. 9c, k; Fig. 12f~g). In Table 5, the category of
shaft fragments is consistently large and reveals that virtually all long
bones were deliberately broken to extract the marrow. This procedure

166

Annals of Carnegie Museum

VOL. 48

0 2cm

f n

Fig. 12.— Femora (a-d), a patella (e), a tibia (f), and a metatarsal shaft (g) with cut
marks. Examples f and g are from Moche; a, b, and e are from Galindo; c and d are
from Chan Chan.

Fig. 13.— Astragali (a-f), first phalanges (g-i), and a second phalange (j) with cut marks.
Example a is from Moche; b-e, g, h, and j are from Galindo; and f and i are from
Chan Chan.

168

Annals of Carnegie Museum

VOL. 48

may also have affected the number of long bone ends, but it is assumed
that the relative frequency is not significantly altered. A number of
North American faunal studies, as well as a few from other parts of
the world, cite marrow extraction as an explanation for the very frag-
mentary condition of long bones at many sites (see Degerbpl, 1929;
Hill, 1966; Parmalee, 1965; and Uerpmann, 1973). Using ethnographic
data others elaborate on the process as a systematic procedure for
extracting bone grease through fracturing and boiling (Leechman,
1951; Gilbert, 1969). Such a procedure would have greatly affected
butchering and food preparation practices because all meat would have
been removed from the long bones before cooking. Data from the
Moche Valley suggest that marrow extraction was an important part
of llama butchering and consumption, but with only the fragmented
bone as evidence, little of the operation can be reliably reconstructed.

Summary and Conclusions

The nature of the sites sampled in the Moche Valley offered consid-
erable potential for an evaluation of the use of the domesticated llama
in later prehistoric periods. Early efforts at quantifying the sample
revealed that a very small number of individual animals were repre-
sented by the samples for each site. This makes many interpretations
tentative, especially with respect to variations within sites or even
between sites that were chronologically different. However, a number
of general observations were possible and these could tentatively be
correlated with known cultural practices. Concrete information on
butchering practices in the form of cut marks on bones was explored
in detail, and possible methods and procedures were suggested.

The interpretations presented here are all too frequently based on
small samples. However, many are justified, and in fact, made possi-
ble, through the tremendous amount of information available for each
culture studied in the areas of nonsubsistence aspects of the societies.

Acknowledgments

Fieldwork resulting in these data was funded by the National Science Foundation and
the Institute of Latin American Studies of the University of Texas. Excavations were
carried out under the auspices of the Chan Chan-Moche Valley Project directed by M.
E. Moseley and C. J. Mackey with the permission of the Peruvian Instituto Nacional
de Cultura. I would like to thank J. R. Topic for making available his faunal material
from the Chan Chan SIAR and E. L. Lundelius for supervision during the original
preparation of this manuscript. Fig. 4 was redrafted from a drawing by G. Bawden, and
Figs. 5-13 are by G. Barr A.

Literature Cited

Baker, F. C. 1930. The use of animal life by the mound-building Indians of Illinois.
Trans. Illinois State Acad. Sci., 22:41-64.

1979

PozoRSKi — Llama Remains

169

— . 1931. Additional notes on animal life associated with the mound builders of
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Cardoza A., G. 1954. Auquenidos. La Paz, 284 pp,

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200 pp.

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fOh 73

piiPm^

ISSN 0097-4463

ANNALS

o/CAKNECIE MUSEUM

CARNEGIE MUSEUM OF NATURAL HISTORY
4400 FORBES AVENUE ® PITTSBURGH, PENNSYLVANIA 15213
VOLUME 48 1 June 1979 ARTICLE 10

KARYOTYPE ANALYSIS, PALYNOLOGY, AND EXTERNAL
SEED MORPHOLOGY OF TOFIELDIA TENUIFOLIA
(MICHX.) UTECH (LILIACEAE-
TOFIELDIEAE)

Frederick H. Utech^

Associate Curator, Section of Plants

Abstract

To broaden the basis for the transfer of Pleea tenuifolia Michx. to the genus Tofieldia
several additional lines of biosystematic evidence are here presented. The first chro-
mosome count for T. tenuifolia (Michx.) Utech, that is = 30, is reported here, as
well as a karyotype analysis of the somatic complement. This number as well as the
general genome morphology agrees remarkably well with the reported counts of 2« =
30 for the genus Tofieldia. {Narthecium is characterized by a somatic number of 26.)
Scanning electron microscope (SEM) observations and photographs of the rare 2-sul-
culate condition in T. tenuifolia and in other members of Tofieldia are presented. The
size and tectum of the pollen in T. tenuifolia and in Tofieldia are also similar. {Narthe-
cium has 1-sulcate pollen grains.) The external seed morphology of T. tenuifolia is
examined and is similar to that of other members of Tofieldia. The distal, caudate
appendages in T. tenuifolia agree well in both size and shape with the appendages in
the Triantha subgroup of the genus. (There are no appendages in the seeds of Narthe-
cium.)

Introduction

! In a previous paper (Utech, 1978), Pleea tenuifolia Michx. (LilL
I aceae^Tofieldieae) was transferred to the genus Tofieldia. This transfer
I was based on an extensive comparison of external floral morphology,

^ Research and publication supported by the M. Graham Netting Research Fund through
a grant from the Cordelia Scaife May Charitable Trust.

Submitted 4 December 1978.

171

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Table L— -Complement homogeneity— Pretreatment comparison of total genome length
(p) and the ratio of the longest (Lj) to the shortest (Sg) chromosome.

Total genome length (pt)

Ratio (Li/Ss)

Mean = 68.88

Mean = 2.127

= 8.75

S" - 0.002

S = 2.95

S = 0.049

62.94 - 74.35

2.003 - 2.261

Colchicine pretreated cells (0.02% at 18-20°C for 4 h).
N = 10 somatic cells.

which emphasized the three, free, stipitate carpel bases and the inter^
carpellary gland, and internal floral vascular anatomy. The former was
compared with members of the genus Tofieldia as well as its tribal
cohort, Narthecium. Numerous similarities were reported between T.
tenuifolia (Michx.) Utech and the genus Tofieldia. However, neither
exhibited any great degree of similarity to Narthecium. The unusual
nine-stamen condition in Tofieldia tenuifolia was shown to be part of
a reductional series that matched the patterns of vascularization within
the genus Tofieldia.

To broaden the basis of the transfer of Pleea to Tofieldia additional
biosystematic data on the chromosome number, the pollen grain, and
seed morphology of T. tenuifolia are reported here.

Materials and Methods

Plant materials for the chromosomal and palynological portions of this study were
collected near Wilmington, North Carolina (Brunswick Co., 1.5 km N of Orton Plan-
tation Gardens along NC highway 133, 16 September 1978, Utech 78-300 CM). From
one large population, which occupied an area of 5 by 4 m, five isolated flowering clumps,
that is, five clones, were sampled for rapidly growing root tips. This species flowers in
autumn and therefore root tip growth is greatest at this time. Excised, actively growing
root tips from each clone were pretreated in 0.02% colchicine at room temperatures (18-
20°C) for 4 h. These root tips were then bulk stained overnight in 45° acetic-orcein
following a brief (30-60 s) fixation in 1:3 acetic-ethanol. A 30 s hydrolysis at 50°C in
a 1:1 mixture of 45% acetic-orcein and 1 N HCl was followed by squashing in a 9:1
mixture of glycerin and 45% acetic acid. Measurements, drawings, and photographs
were made at l,500x using an Olympus microscope equipped with a Leitz micrometer.

Besides general counting within the five clones for euploidy, 10 cells with well spread
and condensed chromosomes were selected for detailed measurement and karyotype
analysis (Tables 1-2, Figs. 1-3). Standard deviations are indicated for the following mea-
surements: long arm (L); short arm (S); total chromosome length; centromeric index
(Table 2). The relative chromosome lengths given (Table 2) have been based on the
absolute length of the complement’s longest chromosome. The centromeric index, which
indicates the chromosomal type, is determined by the short arm (S) to long arm (L) ratio
(S/L), that is, median or metacentric (m) with an index value of 1.000-0.850; submedian
or submetacentric (sm), 0.850-0.450, and subterminal or subtelocentric (st), 0.450-0. OCX).
Terminal or telocentric chromosomes are not recognized in this system of chromosomal

1979

Utech-— Biosystematics of Tofieldia

173

Fig. 1 .—Photomicrograph of a pretreated metaphase spread of Tofieldia tenuifolia
(Michx.) Utech collected near Wilmington, North Carolina {Utech 78-300 CM). 2n =
30. Scale indicated.

classification. Large, medium, and small sized chromosomes are denoted by L, M, and
S, respectively (Utech and Kawano, 1974, 1976).

A large quantity of freshly collected pollen from our T. tenuifolia site in North Carolina
(Utech 78-300) was sent to Dr, Joan W. Nowicke (Smithsonian Institution, Department
of Botany) for SEM analysis and photographs. Pollen samples from recognized Tofieldia
species were also sent as controls. All species have voucher specimens at Carnegie
Museum of Natural History (CM). Species names, locations, and collectors are given
in the legend of Fig. 4. The SEM observational procedures at the Smithsonian Institution
are standardized (Nowicke and Skvarla, 1977). Low power to high power photographs
were taken for each sample. The low power exposures included “stacks” of pollen
grains so that polar views were possible.

Mature seeds from living specimens of T. tenuifolia (Utech 78-3(X)) under cultivation
in Pittsburgh were used for the seed illustration (Fig. 5). Comparative seed material
from other species of Tofieldia from the herbarium at Carnegie Museum of Natural
History (CM) were also illustrated. Species names, locations, and collectors are given
in the legend of Fig. 5.

Observations

Karyotype Analysis o/ Tofieldia tenuifolia

The chromatin of the interphase nuclei stained evenly through the
nucleus. Furthermore no chromocenters or heteropycnotic bodies
were observed at this stage. Metaphase figures with small chromo-
somes, about 1 to 4 that were collected by the colchicine pretreat-
ment technique were abundant within the relatively large root tips.

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VOL. 48

Because the lengths of the longest chromosomes were less than 4 fi,
this associates T. tenuifolia with those small chromosome members of
the otherwise large chromosome species of the Liliaceae Alliance
(Cave, 1970; Sato, 1942).

For each of the five clones a total of 20 counts were made. All 100
counts for this population of T. tenuifolia were 2n = 30. This is the
first chromosomal report for this species. Neither polyploidy, aneu-
ploidy, aneusomaty, or B-chromosomes, including fragments, were
encountered.

Two different checks on the degree of chromosomal condensation
were made for the colchicine pretreatment used. For each of the 10
selected cells with well spread metaphase chromosomes, the ratio of
the longest to the shortest chromosome (L/Ss) and the total genome
length (jji) were calculated (Table 1). These two parameters are very
useful as complement standardization criteria. The average genome
length was 68.88 /x with a range of 62.94 to 74.35 fju and a standard
deviation of 2.95, whereas the average L/S ratio was 2.13 with a range
of 2.003-2.261 and a standard deviation of 0.049. A representative
metaphase spread from the selected set is presented in Figs. 1-3. Fig.
3 is a diploid idiogram of the chromosomes presented in the photo-
micrograph of Fig. 1 and redrawn for Fig. 2.

Ten complements were selected for their maximal condensation,
evenness and flatness of spread. Based on 20 measures per chromo-
some, the following average chromosomal measurements (long arm,
short arm, and total length) with their respective standard deviations,
centromeric indices, and relative chromosomal lengths are presented
(Table 2).

Three groups of chromosomes have been distinguished on the basis
of centromeric indices and lengths (Table 2). The first group comprises
the three largest pairs (3.34-3.06 /x; relative length, 91.6-100.0). These
three pairs are noted as to L3. The second intermediate-sized group
(2.06-2.61 fi; relative length, 61.6-78.1) encompasses seven pairs, that
is, Ml to My. The third and smallest (1.57-1.89 /x; relative length, 47.0-
56.6) group contains the only metacentric pair, that is, pair S5. There
are five pairs in this last group, Sj to S5. No satellites or nucleolar
organizing chromosomes with secondary constrictions were observed,
notwithstanding the small size of the complement members. The anal-
ysis presented in Tables 1-2 of the karyotype of T. tenuifolia agrees
remarkably well with the idiogram presented in Fig. 3.

Table 3 is a summary of previously presented chromosome counts
and numbers in the genus Tofieldia. The most prevalent somatic num-
ber in the genus reported to date isTn = 30, and represents a probable
base number of x = 15. The Japanese T. japonica Miq. with two re-
ported counts of 2/1 = 60 (Table 2) represents a polyploid deviation
within the genus, whereas aneuploid deviations are also known. To-

1979

Utech— Biosystematics of Tofieldia

175

Tofieldia tenuifolia

Fig. 2. — Drawing of chromosomal spread presented in Fig. 1. Arrows indicated zones
of chromosomal overlap. 2n = 30. Scale indicated.

fieldia coccinea Rich, has had several reports of In = 30, but there is
also a report of = 32, whereas T. calyculata (L.) Wahlbg., on the
other hand, has had a single report of 2/2 = 30 and two of 2n = 28.
The 2n = 30 count reported here for T. tenuifolia is not dissimilar
from that known for the genus.

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Table 2. — Summary of karyotype measurements for Tofieldia tenuifolia based on 10
colchicine pretreated, somatic complements.

Length of chromosomes (pt) Centromere

Pair

no.

Long arm

Short arm

Total

Relative

length

index

(s/1)

Type

1

2.45

(0.15)

0.89

(0.10)

3.34

(0.25)

100.00

0.363

(0.05)

St

Li

2

2.10

(0.13)

1.00

(0.10)

3.10

(0.23)

92.8

0.476

(0.04)

sm

u

3

1.95

(0.11)

1.11

(0.08)

3.06

(0.19)

91.6

0.569

(0.04)

sm

u

4

1.68

(0.15)

0.93

(0.09)

2.61

(0.24)

78.1

0.555

(0.05)

sm

M,

5

1.57

(0.12)

0.89

(0.10)

2.46

(0.22)

73.6

0.566

(0.07)

sm

M2

6

1.82

(0.15)

0.64

(0.13)

2.46

(0.28)

73.6

0.351

(0.10)

St

M3

7

1.71

(0.13)

0.64

(0.09)

2.35

(0.22)

70.3

0.374

(0.04)

St

M4

8

1.48

(0.14)

0.78

(0.15)

2.26

(0.19)

67.6

0.527

(0.06)

sm

M5

9

1.25

(0.18)

1.00

(0.08)

2.25

(0.26)

67.3

0.800

(0.14)

sm

Me

10

1.16

(0.14)

0.90

(0.07)

2.06

(0.21)

61.6

0.775

(0.06)

sm

M,

11

1.43

(0.09)

0.46

(0.07)

1.89

(0.16)

56.5

0.321

(0.05)

St

Si

12

1.27

(0.09)

0.50

(0.05)

1.77

(0.14)

52.9

0.393

(0.05)

St

S2

13

0.95

(0.08)

0.71

(0.07)

1.66

(0.15)

49.7

0.747

(0.05)

sm

S3

14

0.89

(0.10)

0.71

(0.08)

1.60

(0.18)

47.9

0.797

(0.06)

sm

S4

15

0.82

(0.08)

0.75

(0.08)

1.57

(0.16)

47.0

0.914

(0.07)

m

S5

Twenty measurements of each somatic pair with standard deviations indicated. Av-
erage total genome length — 68.88 fx.

Palynology o/ Tofieldia tenuifolia

The Liliaceae sensu Engler (1888) and Krause (1930) is a compar-
atively eurypalynous plant family with typical 1-sulcate pollen grains.
Less frequent occurrences of trichotomosulcate, 2-sulculate, zonisul-

1979 Utech—Biosystematics of Tofieldia 111

Tofieldia tenuiflora

it

1

89

2

n

3

4

5

6

88

7

88

8

9

U

lO

08

11

8ft

12

00

13

8»

14

15

P

’I

Fig. 3. — Diploid idiogram of size-ranked complement presented in Figs. 1--2. In = 30.
Scale indicated.

culate, 2“porate, 3-aperturate, and nonaperturate pollen grains are
known (Erdtman, 1966). Scanning electron micrographic (SEM) ob-
servations on the pollen grains of the species of Tofieldia therefore
seemed desirable.

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VOL. 48

Table 3. — Previously published chromosome counts in Tofieldia.

Species Count

Location

Author

Tofieldia pusilla (Michx.)

Pers.

2n = 30

Canada: Arctic

Hedberg, 1967

2n = 30

U.S.S.R.: NE part

Zhukova, 1967

2n = 30

Canada: W part

Taylor and Brockman, 1966

2n = 30

Iceland

Love and Love, 1956

2n = 30

Canada: N. part

Love and Ritchie, 1966

2n = 30

Cult.

Miller, 1930

2n = 30

U.S.A.: Alaska

Johnson and Packer, 1968

2n = 30

Norway

Laane, 1969

Tofieldia coccinea Rich.

2n = 30

U.S.A.: Alaska

Johnson and Packer, 1968

2n = 30

U.S.S.R.: Korjakian

Sokolovskaya, 1968

2n = 30

Greenland

Jorgensen et al., 1958

(aneuploidy) 2n = 32

U.S.S.R.: NE part

Zhukova, 1967

2n = 30

Japan

Matsuura and Suto, 1935

(as T. nutans Willd.)

2n = 30

Japan

Sato, 1942

(as T. nutans Willd.)

Tofieldia nuda Maxim.

2n = 30

Japan

Sato, 1942

Tofieldia japonica Miq.

(polyploidy) 2n = 60

Japan

Matsuura and Suto, 1935

(polyploidy) 2n = 60

Japan

Sato, 1942

Tofieldia glutinosa (Michx.) Pers.

subsp. glutinosa

2n = 30

Canada: British Columbia

Taylor and Mulligan, 1968

subsp. brevistyla C. L. ;

Hitch.

2n = 30

Canada: British Columbia

Taylor and Mulligan, 1968

subsp. occidentalis (S. Wats.) C. L. Hitch.

n - \5

U.S.A.: California

Cave, 1970

Tofieldia calyculata (L.) Walhlbg.

2n = 30

Yugoslavia

Lovka et al., 1971

(aneuploidy) 2n = 28

Poland

Skalinska et al., 1971

(aneuploidy) 2n = 28

Cult.

Miller, 1930

Erdtman (1966:235) reported that the species of Tolfieldia and Pleea
tenuifolia Michx., now Tofieldia tenuifolia (Michx.) Utech (Utech,
1978), had 2“Sulculate pollen grains which were comparatively small,
that is, the longest axis was 30 /x or less in length. Several illustrations
including a cross-section of the 2-sulculate grains of T, calyculata were
also presented. A polar view is required in order to observe the
2-sulculate condition, and this is possible only in a tetrad condition or
if the grains are “stacked’' together.

1979

Utech — Biosystematics of Tofieldia

179

Fig. 4.— -Scanning electron micrographs for selected species of Tofieldia, A. Polar view
showing the 2-sulculate condition in T, pusilla (Michx.) Pers., Alaska: Doutt 48 CM,
US photo/sample #237 (2960x). B. Equatorial view showing one of the two pollen
grooves in T. glutinosa (Michx.) Pers., Michigan: Buker s.n. CM, US photo/sample
#238 (2330x). C. Equatorial view showing one of the two pollen grooves in T, racemosa
(Walt.) BSP, North Carolina: Mozingo s.n. CM, US photo/sample #235 (2140x). D,
Equatorial view showing one of the two pollen grooves in T. tenuifolia (Michx.) Pers.,
North Carolina: Utech 78-300 CM, US photo/sample #236 (2520x).

Pollen grains from four species of Tofieldia were examined under
the SEM. These included T. pusilla (Michx.) Pers., T. glutinosa
(Michx.) Pers., T. racemosa (Walt.) BSP, and T. tenuifolia (Michx.)
Utech. From the low magnification “stack” shots the 2-sulculate con-

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VOL. 48

dition was evident in all four species. Fig. 4A presents a polar view
of T. pusilla, which shows the 2-sulculate condition whereas Fig. 4B-
D shows a single groove which is all that is possible from an equatorial
view for the three additional sampled species. Clearly the pollen grains
of T. tenuifolia (Michx.) Utech (=Pleea tenuifolia Michx.) share the
rare liliaceous 2-sulculate condition with members of Tofieidia. “This
would support congeneric treatment; in a much more limited way the
similarity of size and of the surface of the tectum would also reinforce
that treatment, or put another way they don’t argue against it” (J.
Nowicke, written communication, 1978).

External Seed Morphology in Tofieidia:

Caudate Appendages in T. tenuifolia

Various floras of the northern hemisphere have noted that the seeds
of some, but not all Tofieidia species are appendaged (Fernald, 1950;
Gleason and Cronquist, 1963; Hitchcock and Cronquist, 1973; Hiilten,
1968; Ohwi, 1965; Komarov, 1935). Hitchcock’s treatment of the North
American Tofieidia glutinosa complex (1944) includes several seed
sketches, which clearly demonstrate that this species has definite, dis-
tal, caudate appendages. Gates (1918) noted that the difference be-
tween the then genus Triantha (Nutt.) Baker, now section Triantha,
and Tofieidia {sensu stricto) was among other characters that the for-
mer group had caudate seeds. Tofieidia glutinosa (Michx.) Pers.,
which today includes T. intermedia (Rydb.) Bates and T. occidentalis
(S. Wats.) Bates, has caudate seed appendages and was placed in the
Tofieidia subgroup by Gates (1918). The previous paper (Utech, 1978)
on T. tenuifolia described briefly the appendaged seeds in this species
and presented an outline drawing. To further demonstrate the variation
in seed appendages in the genus Tofieidia {sensu lato), as well as the
lack of appendages, a series of seed drawings (Fig. 5) is presented.

Of the five species whose seeds were selected for illustration, three
species, that is, T. racemosa (Walt.) BSP, T. glutinosa (Michx.) Pers.,
and T. tenuifolia (Michx.) Utech, are from the Triantha subgroup and
do have distal, caudate appendages, whereas the remaining two
species, that is, T. pusilla Willd. and T. calyculata (L.) Wahlbg., are
in the Tofieidia subgroup and do not exhibit the type of appendage
found in the other subgroup.

Clearly there are two subgroups within the genus Tofieidia, which
differ in the distal seed appendages. The presence or absence of this
seed character follows sectional lines. There is some variation in the
absolute length of the caudate appendages in those species where they
are present. In these species the caudate seeds attached lower in the
gynoecium, that is, embryologically older, tend to have longer ap-
pendages than those in the upper gynoecial areas which are younger.
All species of Tofieidia examined are characterized by weak longitu-

1979

Utech — Biosystematics of Tofieldia

181

Fig. S.—Illustratioo of seeds in the genus Tofieldia showing some species with the distal,
caudate appendage and some without. A. T. glutinosa (Michx.) Pers., Michigan; Buker
s.n., 26 August 1948 CM. B. T. racemosa (Walt.) BSP, North Carolina: O. and G.
Jennings s.n., 31 July 1934 CM. C. T. tenuifolia (Michx.) Utech, North Carolina: Utech
78-300 CM. D. T. calyculata (L.) Wahlbg., Sweden: Gotland, A. andH. Kahl s.n., 19
July 1927 CM. E. T. pusilla Willd., Canada; James Bay, Doutt 2275, 22 July 1935 CM.

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VOL. 48

dinal striations. Furthermore, because T. tenuifolia shares the caudate
seed characteristics with the members of the Triantha subgroup, that
is, T. glutinosa and T. racemosa (Fig. 5), Tofieldia tenuifolia should
be associated with these species. Tofieldia glabra Nutt., which occurs
in the outer coastal plain of the Carolinas (Radford et al., 1964; John-
son, 1969) and surrounds the restricted distribution of T. tenuifolia
(Johnson, 1969; Utech, 1978), does not have caudate seeds (Gates,
1918; Utech, personal observations) and, therefore, has its affinities
with the Tofieldia subgroup.

Additional Specimens Examined
(Supplement to Utech, 1978)

Alabama: Baldwin Co., ca. 2 mi W Seminole, 9 October 1972, R. Krai 49,003 (US).
Florida: s.d., Chapman s.n. (US); Bay Co., Lynn Haven, 13 October 1921, C. Bil-
lington 76 (US); Franklin Co., sandy loam of open grassy slash-pine flatwoods, W of
Florida 379, 12 mi S of Liberty-Franklin County line, 0.6 mi S of road to Wright Rec-
reation Area, 9 November 1963, D. B. Ward & E. S. Ford 3648 (US); Suwannee Co.,
moist, sandy soil, Liveoak, 17 September 1900, Biltmore Herbarium 785 a (US); Walton
Co., low open pine area, 6 mi NW of Inlet Beach, 21 September 1952, J. W. Hardin,
W. F. Humphrey & W. H. Duncan 14,135 (US).

Concluding Remarks

This paper has presented additional biosystematic data to support
the reassignment of the monotypic genus Pleea, that is, P. tenuifolia
Michx., to the genus Tofieldia Huds. A previous paper (Utech, 1978)
presented and compared both the floral morphology and the floral vas-
cular anatomy within these two groups. The present paper, on the
other hand, has documented the following: (1) the similarity in chro-
mosome number, that is, T. tenuifolia (Michx.) Utech has In = 30,
which is the prevalent number in Tofieldia ; (2) the similarity in pollen
morphology, that is, T. tenuifolia shares a similar size, a similar tectum
and the 2-sulculate condition, which is characteristic of Tofieldia; (3)
the similarity of distal, caudate, seed appendages in T. tenuifolia to
some members of Tofieldia.

Acknowledgments

The author would like to thank the M. Graham Netting Research Fund (Carnegie
Museum of Natural History) established through a grant from the Cordelia Scaife May
Charitable Trust for supporting this research. I would also like to thank Dr. Joan W.
Nowicke of the Department of Botany at the Smithsonian Institution, Washington,
D.C., for her SEM analysis of the pollen material, and Ms. Nancy J. Perkins, Scientific
Illustrator at Carnegie Museum of Natural History, for her assistance in the preparation
of the botanical illustrations.

1979

Utech— Biosystematics of Tofieldia

183

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Hulten, E. 1968. Flora of Alaska and neighboring territories. Stanford Univ. Press,
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Johnson, A. W., and J. G. Packer. 1968. Chromosome numbers in the flora of
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Krause, K. 1930. Liliaceae. Pp. 227-260, in Die natiirlichen Pflanzenfamilien (A.

Engler and K. Prantl, eds.), Englemann Verlag, Leipzig, 2(1 5a): 227-3 90.

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Polish Angiosperms. Ninth contribution. Acta Biol. Cracov., Ser. Bot., 14:198-213.

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Taylor, R. L., and R. P. Brockman. 1966. Chromosome numbers of some western
Canadian plants. Canadian J. Bot., 44, 8:1093-1103.

Taylor, R. L., and G. A. Mulligan. 1968. Flora of the Queen Charlotte Islands.
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r

97.7ji

ISSN 0097-4463

ANNALS

of CARNEGIE MUSEUM

CARNEGIE MUSEUM OF NATURAL HISTORY
4400 FORBES AVENUE ® PITTSBURGH, PENNSYLVANIA 15213
VOLUME 48 1 June 1979 ARTICLE 11

EDAPHOSAURUS (REPTILIA, PELYCOSAURIA) FROM THE
LOWER PERMIAN OF NORTHEASTERN UNITED STATES,
WITH DESCRIPTION OF A NEW SPECIES

David S Berman

Associate Curator, Section of Vertebrate Fossils

Abstract

A new species, Edaphosaurus colohistion, is based on the greater portion of a large,
presacral vertebral column from the upper part of the Pittsburgh Formation, Monon-
gahela Group, of northwestern West Virginia. E. colohistion and fragmentary Edapho-
saurus specimens from the later Lower Permian Washington and Greene Formations,
Dunkard Group, of the Tri-state area constitute a morphological series in chronological
order that in some instances conforms and in others deviates from the well-documented
evolutionary trends seen in the series of four essentially consecutively occurring Lower
Permian Edaphosaurus species of the Southwest. On the basis of 1) the evolutionary
trends exhibited by the Tri-state edaphosaur series that conform to those of the South-
west series, 2) the large size of E. colohistion, and 3) the numerous, typically Lower
Permian amphibians previously reported from the same locality as E. colohistion and
from various levels in the lower half of the Dunkard Group, it is suggested that the
Pittsburgh Formation, or at least its upper levels, and the Dunkard Group are correlative
with the Lower Permian Wichita and lower Clear Fork Groups of north-central Texas
and that the Wolfcampian-Leonardian Series boundary lies near the base of the Greene
Formation. Geographic isolation of the Dunkard basin from the Micontinental basin
complex is offered as an explanation for the deviations of the Tri-state Edaphosaurus
series from the evolutionary trends exhibited by the Southwest series. The Tri-state
series probably represents a single lineage that evolved in place and independently from
the southwestern forms.

Introduction

The herbivorous, swamp-dwelling pelycosaur reptile Edaphosaurus
is widespread both spatially and temporally. Eight species are recog-

Submitted 4 December 1978.

185

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VOL. 48

nized from deposits of Late Pennsylvanian and Early Permian age in
North America and Europe. The genus is best known, however, from
the Lower Permian of north-central Texas, where it is represented by
three, consecutively occurring species, E. boanerges, E. cruciger and
E. pogonias, from beds extending from the lower Wichita Group up
through the overlying lower Clear Fork Group. They exhibit a pro-
gressive increase in overall body size that is accompanied by a de-
crease in relative sail size, and gradual changes in the vertebral neural
spines forming their large dorsal sail. These changes suggest that they
represent a species phylum that evolved in place (Romer and Price,
1940). The poorly-known E. novomexicanus of New Mexico, the only
other Lower Permian Edaphosaurus species of North America, is
viewed as a morphological antecedent to the Texas series. In the Tri-
state area of Pennsylvania, West Virginia, and Ohio, Edaphosaurus
is known from a number of fragmentary specimens from the Upper
Pennsylvanian and Lower Permian. The earliest occurring specimen,
a small fragment of neural spine from the middle of the Conemaugh
Group, was first described by Case (1908) as Naosaurus raymondi.
Naosaurus was later synonymized with Edaphosaurus (Romer and
Price, 1940). In 1952 Romer described all the then known Edaphosau-
rus specimens from the Tri-state area. On the basis of overall body
size and neural spine structure he referred those specimens from the
Washington and the base of the Greene Formation to E. boanerges
and those from the middle and upper Greene Formation to E. cruciger.
Since Romer’s (1952) report, additional Edaphosaurus specimens have
been discovered in the Tri-state area. Most important among these is
the greater part of a presacral vertebral column, including the dorsal
sail, from the upper Pittsburgh Formation, Monongahela Group. This
is the first Edaphosaurus specimen reported from the Monongahela
Group and, though it has been identified as E. boanerges (Lund, 1972,
1975, 1976), it has not been described. It is notable for its large size,
which is equal to that of E. boanerges from the Lower Permian of
Texas. This is a principal reason for believing that the Pittsburgh For-
mation is Lower Permian and not Upper Pennsylvanian as most au-
thors believe; Pennsylvanian members of this genus are much smaller
than their Permian descendants. The combined features of overall size
and structure of the neural spines distinguish the Monongahela form
as a new species, E. colohistion.

Viewed in chronological order the Monongahela- Dunkard edapho-
saurs exhibit changes that in some instances parallel and in others
deviate from the well-established evolutionary trends of the Lower
Permian species of the Southwest. Similarities in evolutionary trends
of both groups permit stratigraphic correlations between the Pittsburgh
Formation and the overlying Dunkard Group, and the classic Lower

1979

Berman — Lower Permian Edaphosaurus

187

Permian terrestrial section of north-central Texas. Previously de-
scribed amphibians from the same locality in the Pittsburgh Formation
from which came the holotype of E. colohistion and from various
horizons in the lower half of the Dunkard Group, though broadly rein-
forcing correlations based on Edaphosaurus , necessitate some minor
reassessments. Differences in trends between the edaphosaurs of both
areas are, however, sufficient to suggest that those from the Tri-state
may represent a single lineage that evolved in place and independently
from those of the Southwest. This is accounted for by geographic
isolation of the Dunkard basin by the end of the Pennsylvanian.

Abbreviations CM and USNM are used to refer to collections of the
Carnegie Museum of Natural History and the National Museum of
Natural History.

Systematic Paleontology

Class Reptilia
Order Pelycosauria
Family Edaphosauridae
Genus Edaphosaurus Cope 1882
Edaphosaurus colohistion, new species

Holotype. — CM 23513 consists of an articulated, or nearly articu-
lated, series of 14 essentially complete presacral vertebrae believed to
include cervicals 6 and 7 and dorsals 8 through 19 (Fig. 1). Disarticu-
lated but closely associated with the partial vertebral column are four
or five intercentra, two incomplete vertebrae and numerous neural
spine fragments near both ends of the column, and numerous ribs. The
holotype was collected by Dr. Richard Lund of Adelphi University in
1969.

Horizon.— Limestone “B” of the Pittsburgh Formation, Mononga-
hela Group. The age of the Pittsburgh Formation has been generally
accepted as Late Pennsylvanian, Virgilian, but is considered here, at
least its upper levels, as probably Early Permian, Wolfcampian.

Locality .—Road cut on Interstate Highway 70 about Vi mi east of
Elm Grove, West Virginia. The deposit from which the holotype was
collected has been described by Lund (1972:51) as a meander cutoff
channel that filled very slowly.

Differs greatly in size from all Edaphosaurus species
except the Lower Permian E. boanerges and E. novomexicanus . E.
colohistion is distinguished from E. boanerges in the following features
of its neural spines: form a relatively much shorter sail; much closer
spacing of lateral tubercles; no anteroposterior expansion of distal por-
tions of posterior cervical or anterior dorsal spines; sail decreases in
height at the same rate anteriorly and posteriorly from its highest level.

188

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VOL. 48

-Edaphosaurus colohistion, holotype, CM 23513.

1979

Berman — Lower Permian Edaphosaurus

189

E. colohistion is similar to E. novomexicanus in lacking any antero-
posterior expansion of the neural spines but differs in being somewhat
larger, in its greater development of the basal tubercles of the posterior
cervical spines and in its greater cross-sectional thickening of the distal
portions of the cervical spines.

Etymology. — Greek kotos, meaning docked, shortened or stunted, and histion, mean-
ing sail, referring to its relatively small sail.

Description. — The holotype (Fig. 1) consists of a series of 14 essentially complete,
articulated, or nearly articulated, presacral vertebrae; close to either end of this series
are two incomplete vertebrae and numerous spine fragments. Four, or possibly five,
disarticulated intercentra and numerous ribs lie near the articulated column. All but a
few of the articulated vertebrae are exposed in right lateral view and their centra show
considerable lateral crushing; the centra exposed in end view retain their circular outline.
The entire specimen, especially the neural spines, exhibits numerous fractures along
which there has been in most cases some separation. Most of the spines appear to be
complete and by way of comparison with the restoration of Edaphosaurus boanerges
by Romer and Price (1940:391, Fig. 66) the longest spine of E. colohistion probably
belongs to the fourteenth vertebra and, therefore, lies near the middle of the presacral
series which in Edaphosaurus is believed to consist of somewhere between 24 and 27
vertebrae. If this identification is correct, the articulated series would include the pos-
terior two cervicals and the anterior 12 dorsals. Crushing and incomplete preservation
makes regional differentiation of the centra impossible; it can be said, however, that the
centra exhibit no noticeable differences from those of other members of this genus. Of
the centra complete enough or sufficiently exposed to take one or more of the following
measurements, the recorded size ranges are: (1) length of centrum, 30.0 to 34.5 mm;
(2) width of centrum, 24,0 to 25.0 mm; (3) height of centrum, 24.0 to 29.0 mm. Averages
for these measurements for the dorsal vertebrae are given in Table 1. Crushing of the
laterally exposed centra has undoubtedly slightly increased their height and decreased
their width; undistorted, these two dimensions would probably be nearly equal, about
25 mm. As is typical of Edaphosaurus, the intercentra are small, anteroposteriorly
narrow, low crescents. According to Romer and Price (1940) the rare occurrence of
intercentra in Edaphosaurus suggests that most of the intercentra remained cartilagi-
nous.

The structure of the neural spines in E. colohistion is in close accord with the pattern
in Edaphosaurus generally. The proximal portions of the spines are laterally compressed
with an anteroposterior length of about 18.0 to 24.0 mm and a transverse width of about
10.0 to 14.0 mm. Just above the first tubercle they taper rather abruptly to a subcircular
section, then gradually narrow to their termination. The spine of the presumed four-
teenth vertebra, for example, narrows in anteroposterior diameter to 14, 12, and 10 mm
at levels of 14, Vi, and Ya its height. For most of their length the neural spines exhibit
a prominent longitudinal ridge on the anterior face, whereas posteriorly there is a lon-
gitudinal groove. The longest spine, that of the presumed fourteenth vertebra, is 445
mm long measured from the zygapophyses. The height of the sail appears to decrease
by about the same rate anteriorly and posteriorly from its highest point. There is no
indication of anteroposterior expansion of the distal portions of the neural spines be-
lieved to belong to the posterior cervicals and anterior dorsals such as occurs in varying
degrees in Lower Permian species (Romer and Price, 1940).

As is characteristic of Edaphosaurus the lateral tubercles or crossbars of the neural
spines tend to be arranged in bilaterally symmetrical pairs that occur at rather regular
intervals along the spine and form anteroposterior rows with those of successive spines;
this pattern, however, becomes increasingly irregular toward the distal ends of the spines

190

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VOL. 48

Table 1. — Measurements (in mm) of vertebrae in Edaphosaurus; OLU refers to ortho-
metric linear unit (radius of centrum to the % power) of Romer and Price (1940). All
measurements are of individual elements except those for E. colohistion and Southwest
species (from Romer and Price, 1940: Table 5) which are averages of dorsal vertebrae.

Specimen or
species

Centrum

length

Centrum

width

Centrum

height

Antero-
posterior
length
of spine
at base

OLU

value

Tri-State Edaphosaurs

CM 8604

18.0

17.0

CM 8540

18.0

17.0

4.16

17.0

18.0

4.33

16.0

18.0

CM 23818

16.0

CM 8601

16.0

CM 8600

13.0

USNM 205488

23.0

22.0

17.0

28.0

26.0

23.0

19.0

5.53

25.0

23.0

5.38

30.0

26.0

26.0

20.0

33.0

24.0

25.0

17.0

Marietta College specimen

19.0

18.0

17.0

Edaphosaurus colohistion CM 23513

33.2

24.5

26.1

19.2

5.24

Southwest Edaphosaurs

Edaphosaurus pogonias

46.0

36.0

35.0

6.87

Edaphosaurus cruciger

41.0

34.0

34.0

6.61

Edaphosaurus boanerges

34.0

24.0

25.0

5.24

Edaphosaurus novomexicanus

33.0

21.0

21.0

4.79

and the anterior end of the column. Average intertubercular distances for the first seven
tubercles of the articulated series of presumed dorsal vertebrae of the holotype are given
in Table 3. The lateral tubercles are well developed with rounded ends and become
shorter more distally along the spine; as an example, the first six tubercles on the right
side of an anterior dorsal spine have lengths of 20, 17, 10, 8, 8, and 5 mm. The tubercles
are further reduced distally to nubbins. The highest number of tubercles that I am able
to estimate along any one side of a neural spine is about 14.

The ribs of CM 23513 are characteristic of Edaphosaurus in being greatly curved
throughout their length and in having the tubercula represented by only a slightly raised,
rugose area on the shaft.

Comparisons

Recognition of Edaphosaurus species has been based on vertebral
structure, overall size, and to some extent stratigraphic level. On the

1979

Berman — Lower Permian Edaphosaurus

191

Table 2.- — Length of longest neural spines in Edaphosaurus; OLU refers to orthometric

linear unit of Romer and Price (1940). Measurements for Southwest species from

Romer (1948).

Specimen or
species

Longest spine
in mm

Longest spine
in OLU

Tri-State Edaphosaurs

USNM 205488

524

96

420

77

Edaphosaurus colohistion CM 23513

445

85

Southwest Edaphosaurs

Edaphosaurus pogonias

645

92

Edaphosaurus cruciger

720

109

Edaphosaurus cf. boanerges

594

no

Edaphosaurus boanerges

552

108

basis of overall size and neural spine structure Edaphosaurus coloh-
istion could be confused with only two of the four species known from
the Lower Permian of the Southwest, E. boanerges and E. novomex-
icanus.

In overall size, E. colohistion falls well within the range of E. boa-
nerges, which is best known by a number of excellent specimens from
the Wichita Group of north-central Texas (Romer and Price, 1940).
Differences in the structure of their vertebral neural spines, however,
allow them to be easily separated. The dorsal sail of E. colohistion is
relatively much shorter than that of E. boanerges. As indicated in
Table 2, the maximum lengths of dorsal spines published by Romer
and Price (1940) for two specimens of E. boanerges, representing near
minimum and maximum (possibly a separate species) sized individuals,
range from over 100 to almost 150 mm longer than that of E. coloh-
istion. In terms of the orthometric linear units used by Romer and
Price (1940:8) to express linear measurements in values relative to the
animal’s overall size- — one linear unit is defined as equal to the radius
of the average sized dorsal centrum to the % power— -the longest spines
of the above E. boanerges specimens exceed that of E. colohistion by
30 units. Whereas in E. colohistion the sail decreases in height at the
same rate anteriorly and posteriorly from its highest level, in E. boa-
nerges, as figured by Romer and Price (1940: Fig. 66), it decreases at
a lesser rate posteriorly. As indicated in Table 3 the average distances
between the neural spine tubercles are noticeably smaller in E. coloh-
istion than in E. boanerges. The number of tubercles per spine appears
to be about equal in the two species; the highest count of tubercle
pairs that Romer and Price (1940) were able to see on a specimen of
E. boanerges was 16, as compared to 14 for E. colohistion. Romer
and Price (1940) point out that in E. boanerges there is a definite,
though slight, anteroposterior expansion of the cervical spine tips; this
does not appear in E. colohistion.

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Table 3. — Average distances (in mm) between first six tubercles of dorsal neural spines
in Edaphosaurus; extremes are in parentheses. Measurements for E. boanerges from
Romer and Price (1940:384).

species

1-2

2-3

3^

4-5

5-6

6-7

CM 23818

33

42

49

31

32

USNM 205488

74

(72-78)

93

(84-105)

57

(48-64)

Edaphosaurus colohistion
CM 23513

34

(28-52)

36

(23-68)

31

(14-45)

30

(15-37)

29

(13-37)

33

(26-38)

Edaphosaurus boanerges

61

(52-70)

55

(37-71)

44

(30-62)

43

(23-58)

40

(27-48)

36

(27-40)

E. novomexicanus, smallest of the North American Lower Permian
species of Edaphosaurus, is based on a fragmentary anterior half of
skeleton from the Abo Formation of northern New Mexico (Romer
and Price, 1940). This specimen is somewhat smaller than E. coloh-
istion; its orthometric linear unit value is estimated by Romer and
Price (1940) to be 4.79, whereas that calculated for E. colohistion is
5.24. The type of E. novomexicanus includes cervicals 2 through 7 and
the centrum and base of the spine of what is believed to be dorsal 12.
The cervical spines are nearly complete and show no anteroposterior
expansion at their tips; in this feature they are similar to those of E.
colohistion. Though none of the dorsal spines are preserved, Romer
and Price (1940) note that the nature of its cervical spines suggests
that the dorsal spines were at least as long as those of later American
species; if true, E. novomexicanus would differ from E. colohistion in
having a relatively larger sail. The cervical spines of E. novomexican-
us, as figured by Williston and Case (1913:77), are considerably more
slender than those of E. colohistion. The maximum anteroposterior
length of the subcircular, distal portions of the posterior cervical spines
is about 8 mm in E. novomexicanus and about 15 mm in E. colohistion',
expressed in orthometric linear units, these measurements convert to
1.7 and 2.8 units, respectively. In contrast to E. colohistion, the large
basal tubercles at the level of the transition from the proximal to the
distal portion of the spine of the posterior cervicals are far less deveh
oped in E. novomexicanus.

Evolutionary Trends in Edaphosaurus

Edaphosaurus of the Southwest.— Whsti is known about structural
trends in Edaphosaurus is based almost entirely on the three consec-
utively occurring species, E. boanerges, E. cruciger, and E. pogonias,
from the Lower Permian of north-central Texas which probably de-
scended one from another in situ and, therefore, constitute a species

1979

Berman — Lower Permian Edaphosaurus

193

phylum (Romer and Price, 1940). Romer and Price (1940) estimated
their weights at 83, 166, and 186 kg and their lengths at about 250, 272,
and 327 cm, respectively. This steady increase in overall size, how-
ever, was not accompanied by a similar increase in sail size. As Romer
(1948) pointed out and as Table 2 indicates, using greatest spine length
as an index to overall sail size, their sails increased only slightly in
absolute size, but in terms of orthometric units have ceased to grow
and, in fact, the largest member of the series possesses the smallest
sail relative to body size.

A number of gradual structural changes in the spines of the three
Texas species have been discussed or made obvious in illustrations by
Romer and Price (1940: Figs. 66, 67, 68). There is not only a tendency
toward greater cross sectional thickening of the spines, but of antero-
posterior expansion of the distal portions of the cervical spines. In E.
boanerges anteroposterior expansion of the cervical spines is slight
and extends posteriorly to include the sixth vertebra, in E. cruciger
expansion is somewhat more prominent and extends to the tenth ver-
tebra, and in E. pogonias the spines are greatly expanded to the elev-
enth vertebra and appear club-shaped in side view. A decrease in num-
ber of pairs of lateral tubercles per spine, especially conspicuous at
the anterior end of the column, is also exhibited by the Texas species.
Concomitantly, there is an increase in the spacing of the tubercles
throughout the column; this is most pronounced in the lower portions
of the spines and along the entire length of the spines of the anterior
half of the column. The three species also exhibit a progressive in-
crease in development of the tubercles. In E. boanerges the tubercles
become gradually smaller toward the distal end of the spine where
they are reduced to nubbins. In the two larger species the tubercles
are in general larger, the tips may be somewhat irregularly expanded
and those at the distal ends of the spines, particularly in the cervical
region, may be closely clustered in groups of two or three, or divide
into two or more processes.

In both overall size and morphology of the spines E. novomexican-
us, a contemporary of E. boanerges from New Mexico and Utah
(Williston and Case, 1913; Vaughn, 1963, 1966, 1969), is considered by
Romer and Price (1940) to be a close antecedent to E. boanerges. It
is smaller than E. boanerges, having an estimated weight of about 63
kg and a length of about 241 cm, and its cervical spines are narrower
and show no signs of anteroposterior expansion.

Edaphosaurus of the Tri-state area. —In 1952 Romer published a
comprehensive report on the vertebrate fossils from the Upper Penn-
sylvanian and Lower Permian of the Tri-state area; stratigraphic and
locality data for these fossils were published by Moran (1952). In Ro-
mer’s report numerous fragmentary remains, mostly vertebrae, of

194

Annals of Carnegie Museum

VOL. 48

Edaphosaurus were referred to either E. boanerges or E. cruciger.
These assignments were based on overall size and neural spine struc-
ture. In general he assigned those specimens from the Washington and
basal Greene Formations to E. boanerges and those from the middle
and upper Greene Formation to E. cruciger. Reexamination of these
specimens and study of new finds, however, indicates that the Tri-
state Edaphosaurus specimens exhibit trends that deviate from some
of those of the three consecutive Texas species but conform to others.
Only those specimens from the Monongahela and Dunkard Groups
having vertebral elements useful in comparing relative overall sizes or
in revealing differences in spine structure between individuals are dis-
cussed here; E. raymondi from the Late Pennsylvanian Conemaugh
Group of southwestern Pennsylvanian (Case, 1908) is too incomplete
to be useful. The approximate stratigraphic positions of the specimens
discussed here are indicated on the generalized geologic column of
Fig. 2; in stratigraphic sequence from the lowest occurrence, these
specimens include:

CM 23513, E. colohistion, holotype (already discussed).

Uncataloged specimen belonging to Marietta College, Ohio, de-
scribed by Whipple and Case (1930) and consisting of several partial
spines preserved in serial order and fragments of spines and ribs from
the Upper Marietta Sandstone, Washington Formation, Jackson Coun-
ty, West Virginia (Locality I of Moran, 1952).

USNM 205488, an undescribed specimen consisting of about seven
disarticulated but closely associated, well-preserved dorsal vertebrae,
some of which are nearly complete, and numerous spine fragments
from the Upper Marietta Sandstone, Washington Formation, just south
of Belpre, Washington County, Ohio.

CM 8600, proximal portion of spine from Jollytown Sandstone,
Greene Formation, Putnam County, West Virginia (Locality 12 of
Moran, 1952).

CM 8601, fragments of spine from Middle Rockport Limestone,
Greene Formation, Monongalia County, West Virginia (Locality 25 of
Moran, 1952).

CM 23818, complete spine from a level between the Middle and
Upper Rockport Limestones, Greene Formation, Monongalia County,
West Virginia (Locality 28 of Moran, 1952).

CM 8540, numerous fragments of vertebrae and ribs from just be-

Fig. 2. — Generalized geologic column of the Upper Pennsylvanian and Lower Permian
of southwestern Pennsylvania (after Berryhill and Swanson, 1962) showing approximate
stratigraphic positions of Edaphosaurus specimens discussed in text.

1979

Berman — Lower Permian Edaphosaurus

195

GROUP

FORMATION

BED

Jollytown Coal

O

UNION-

TOWN

Washinqton

Coal

Waynesburg

Coal

Uniontown

Coal

Pittsburgh Coal

CM 8604
CM 8540

CM 23818
CM 8601

CM 8600

USNM 205488
Marietta College
Specmen

CM 23513

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Annals of Carnegie Museum

VOL, 48

neath Windy Gap Coal, Greene Formation, Wetzel County, West Vir=
ginia (Locality 35 of Moran, 1952).

CM 8604, fragments of spine either from just above limestone that
lies 50 ft beneath Windy Gap Coal or from just above the Windy Gap
Limestone, Wetzel County, West Virginia (Locality 37 of Moran,
1952).

It is of course realized that there are regional size differences in the
presacral vertebrae of Edaphosaurus and the use of isolated vertebrae
to compare sizes of individuals introduces the chance of some error.
Further, among the fragmentary specimens listed above some are ob-
viously not representative of the maximum sizes reached by Edapho-
saurus at the horizons from which they were collected. It is assumed,
however, that the number of specimens considered is large enough to
reduce the effects of these errors to a level that will eliminate erro-
neous conclusions.

Although the Tri-state specimens occur over a wide stratigraphic
sequence that undoubtedly represents a considerable length of time,
they do not exhibit the marked increase in overall size with time seen
in the Texas species and remain within the size range of the earliest
occurring Texas species, E. boanerges (Table 1). On the basis of cen-
trum size and anteroposterior length of the spine base, USNM 205488
from the Washington Formation shows only a small increase in size
over E. colohistion from the Pittsburgh Formation. The same mea-
surements for the other specimens from the Washington and Greene
Formations indicate individuals of a slightly to moderately smaller size
than USNM 205488 and E. colohistion CM 23513.

Although the Tri-state edaphosaurs do not appear to increase in
overall size with time, their sails may have increased slightly in rela-
tive size. The evidence for this is limited to a comparison between
E. colohistion CM 23513 and USNM 205488 (Table 2). Spine length
is available in only two of the vertebrae of USNM 205488; in one the
spine is complete and is 420 mm long, whereas in the other the distal
half of the spine is missing a small central portion and what is believed
to be the end of the spine is represented by impression. If the extent
of the latter spine has been correctly determined, its length is 525 mm;
expressed in orthometric linear units it is 11 units longer than the
longest spine of CM 23513.

Edaphosaurus specimens of the Tri-state area appear to exhibit
structural changes in the neural spines that parallel those seen in the
southwestern species. Although there is no obvious indication of a
tendency toward greater cross sectional thickening of the spines, there
is definite evidence of a trend in anteroposterior expansion of the spine
tips and greater development of the tubercles of the spines. None of

1979

Berman — Lower Permian Edaphosaurus

197

Fig. 3. — A, and B, neural spine tips and C, a small section of neural spine presumably
from the cervical or anterioi: dorsal region of Edaphosaurus CM 8540, Greene Forma-
tion, Dunkard Group.

the spines of E. colohistion CM 23513, including those of the presumed
posterior cervicals, show expansion or unusual development of the
tubercles. The earliest occurring Tri-state specimen exhibiting expan-
sion of the spine tips is the Marietta College specimen from the Wash-
ington Formation. A series of spine tips of this specimen, presumably
from the cervical region, are somewhat expanded but the tubercles
remain as small nubbins. In these features the Marietta College spec-
imen is similar to E. boanerges. CM 8540 is the only specimen from
the Greene Formation in which the spine tips from presumably the
cervical or anterior dorsal region are preserved (Fig. 3); they exhibit
a much greater development compared to those of the Marietta College
specimen. One of the spine tips is greatly expanded but its tubercles
remain small, two others are only slightly expanded but possess closely
grouped, well-developed tubercles; a portion of a fourth spine bears
a tubercle with a greatly expanded base from which project two well-

198

Annals of Carnegie Museum

VOL. 48

developed processes. As pointed out by Romer (1952) these characters
suggest a stage of development paralleling that of E. cruciger of the
Texas edaphosaur series.

There is also limited evidence of a progressive increase in tubercular
spacing and a reduction in the number of tubercles per spine in the
Tri-state series of edaphosaurs. As Table 3 shows, the spacing of the
tubercles in E. colohistion is considerably less than in E. boanerges.
USNM 205488 from the Washington Formation exhibits a very sub-
stantial increase in tubercular spacing compared to that in E. coloh-
istion and a moderate increase compared to that in E. boanerges. A
few intertubercular distances are available for the lower portions of
four spines preserved in serial order in the Marietta College specimen
from the Washington Formation; the measurements are consistent with
those for USNM 205488. It is also estimated that the spines of USNM
205488 probably possessed at most about eight pairs of tubercles, a
number that is less than the minimums for the dorsal spines of E.
colohistion and E. boanerges (see Romer and Price, 1940: Fig. 66),
but is in line with minimum counts for the anterior dorsals of E. cru-
ciger or the middorsals of E. pogonias (see Romer and Price, 1940:
Figs. 67, 68). A complete, isolated spine from the Greene Formation,
CM 23818, deviates somewhat from the trends discussed above, but
this is probably due to its being from an immature individual. The
spine is only 380 mm long with an anteroposterior length at its base of
13 mm. It possesses six pairs of weakly developed tubercles that be-
come smaller distally until they are no longer detectable on the distal
third of the spine. The intertubercular spaces are comparable to those
in E. colohistion.

Discussion

Although most authors consider the Dunkard Group to be Lower
Permian, there persists some widely differing opinions concerning its
biostratigraphic placement. Using vertebrate, invertebrate, and plant
fossils (Barlow, 1975), some consider the entire Dunkard to be Penn-
sylvanian, others place the Pennsylvanian-Permian boundary at var-
ious levels in the Dunkard, some accept the traditional placement of
the base of the Permian at or very near the base of the Dunkard,
whereas others include all or portions of the underlying Monongahela
Group in the Permian. Views based on vertebrates, however, have
been fairly consistent and workers have assigned either the Dunkard
Group alone (Romer, 1952; Berman and Berman, 1975; Olson, 1975)
or with the upper part of the Monongahela Group (Lund, 1975, 1976)
to the Lower Permian. In this paper the latter view is supported.

Information presented here indicates that the edaphosaurs from the
upper Pittsburgh Formation, Monongahela Group, and the Dunkard

1979

Berman — Lower Permian Edaphosaurus

199

Group do not exhibit a progressive increase in size as in the south-
western species but remain, for the most part, within the size range of
E. boanerges, smallest member of the Lower Permian Edaphosaurus
series of Texas. The few exceptions are from the middle and upper
Greene Formation and they, in contrast, appear to have an overall size
that is smaller than that of E. boanerges. The fact that E. colohistion
attained a size well within the range of E. boanerges is interpreted as
evidence that the upper Pittsburgh Formation is correlative with the
lower Wichita Group. It should also be remembered that E. colohistion
is larger than E. novomexicanus , which occurs in beds in New Mexico
and Utah (Williston and Case, 1913; Vaughn, 1963, 1966, 1969) that
are considered equivalent to the lower parts of the Wichita Group
(Langston, 1953; Romer, 1960). Trends in the anteroposterior length-
ening of the upper portions of the anterior neural spines and the greater
development of the spine tubercles in the Southwest species appear to
have been paralleled in the Monongahela-Dunkard edaphosaurs. These
parallelisms suggest that the upper Pittsburgh Formation is correlative
with the lower Wichita, the Washington Formation up through the
lower and middle Greene Formation with the middle Wichita, and the
upper Greene Formation with the upper Wichita Group. These cor-
relations in turn place the Pennsylvanian-Permian boundary at about
the middle or lower Pittsburgh Formation and the Wolfcampian-Leo-
nardian Series boundary of the Lower Permian at about middle
Greene Formation. Amphibians recently discovered in the Dunk-
ard corroborate in great part the correlations based on Edapho-
saurus, except that some suggest an equivalence of the Greene
Formation and possibly the upper Washington Formation with the
Clear Fork Group of Texas. Heading this list is Trematops described
by Olson (1970) from the Creston Shale just below the Upper Marietta
Sandstone, a horizon equivalent to the Washington Coal “A” of the
upper Washington Formation. The family to which it belongs, Tre-
matopsidae, is restricted to the Lower Permian and in Texas the three
recognized species of Trematops are confined to the Clear Fork
Group. Broiliellus was reported (Berman and Berman, 1975) from the
Mount Morris Limestone, Washington Formation (Waynesburg For-
mation of the nomenclature used in Fig. 2), which is only about 60 ft
above the Waynesburg Coal, the top of the Monongahela Group. Bro-
iliellus, a well represented and moderately advanced member of the
Dissorophidae, includes four species from the Lower Permian of Texas
that have a stratigraphic range from the Putnam, lower Wichita Group,
to the Arroyo Formation, lower Clear Fork Group. From the same
site in the upper Pittsburgh Formation that yielded the holotype of E.
colohistion, Lund (1972, 1975, 1976) has also collected and identified,
or tentatively identified, the amphibians Diploceraspis burkei (CM

200

Annals of Carnegie Museum

VOL. 48

25206), Lysorophus dunkardensis (CM 25653), Edops (not cataloged),
and Zatrachys serratus (CM 25659); the specimen described here as
the holotype of E. colohistion was identified from this site by Lund as
E. boanerges. Noting that D. burkei is the most common vertebrate
from the Greene Formation, that Zatrachys is confined to Wichita and
Clear Fork equivalents of the Southwest, and that E. boanerges is
known from the Washington and Greene Formations as well as the
Wichita Group of Texas, Lund ascribed a Wolfcampian age to the
upper Pittsburgh Formation and suggested that the upper Greene For-
mation may be equivalent to the basal Leonardian Clear Fork beds of
Texas. Considering the vertebrate evidence as a whole, the Pittsburgh
Formation, or at least its upper levels, and the Dunkard Group are
probably equivalent to the Wichita and lower Clear Fork Groups of
Texas with the Wolfcampian-Leonardian boundary lying near the base
of Greene Formation.

The observations that the Tri-state edaphosaurs, in contrast to those
of the Southwest, do not show a progressive increase in overall size
but remain the same size, or even possibly become smaller, and that
their sails may have increased, rather than decreased, in relative size
may indicate that they evolved in isolation. There are other members
of the Dunkard fauna that appear to have developed in isolation from
the Lower Permian faunas of the Southwest. I (Berman, 1978) have
recently described a new species of the rare, Early Permian pelycosaur
genus Ctenospondylus, C. ninevehensis, from a very high level in the
Greene Formation, the Nineveh Limestone. Though C. ninevehensis
existed at the same time or very probably somewhat later than the
only other member of this genus, C. casei from the Lower Permian of
Texas (Romer and Price, 1940) and Utah (Vaughn, 1964), the former
exhibits a greater primitiveness in a number of features that makes it
an ideal predecessor to C. casei. Parallel evolution following long-term
separation has long been accepted (Beerbower, 1963) as the explana-
tion for the remarkable similarity between the amphibians Diplocer-
aspis from the Upper Pennsylvanian and the Lower Permian of the
Tri-state area and Diplocaulus from the Lower Permian of the South-
west, both noted for their bizarre long-horned skulls.

I (Berman, 1978) have attempted to explain on paleogeographic
grounds the possible presence of relictual or endemic forms in the
Dunkard fauna. During the Early and Middle Pennsylvanian a shallow,
northeastern arm of the Midcontinental basin complex, the Appala-
chian basin, extended into the Tri-state area. During this time an un-
broken habitat zone, or zones, probably allowed free faunal move-
ments between the Tri-state and Midcontinental regions. With the
close of the Pennsylvanian, however, expansion of areas of low relief
eliminated all of the Appalachian basin except its northeastern termi-

1979

Berman — Lower Permian Edaphosaurus

201

nation, which persisted into the Permian as the Dunkard basin. The
Dunkard basin was a restricted basin that was bordered on the east
and southeast by the active old Appalachian highlands and on the west
by the stable continental interior, specifically the Cincinnati Arch. Sep-
arated by at least 1,000 mi, it is possible to envision the Lower
Permian deltaic faunas of the Midcontinental basin complex and the
Dunkard basin as having had different evolutionary histories. With
regard to the Monongahela- Dunkard Edaphosaurus series, it seems
very probable that it represents a single lineage that evolved in situ
and independently from the Lower Permian species of the Southwest.
If this view is accepted, then it seems equally probable that the Dun-
kard edaphosaurs include two as yet undescribed species— -the Wash-
ington and lower Greene edaphosaurs pertain to one species and those
from the middle and upper Greene to a second. Though this is likely,
recognition of new Dunkard Edaphosaurus species is best delayed
until they can be based on more complete materials.

Acknowledgments

I wish to express my gratitude to Dr. Richard Lund of Adelphi University, who not
only discovered and collected the specimen described here as the type of Edaphosaurus
colohistion, but brought it to my attention. I am also indebted to Dr. Nicholas Hotton
III of the National Museum of Natural History and Dr. Dwayne D. Stone of Marietta
College for the loan of specimens in their care. Thanks are also due Dr. Mary R. Dawson
of the Carnegie Museum of Natural History and Dr. Robert Reisz of Erindale College,
University of Toronto, for critical reading of the manuscript.

Literature Cited

Barlow, J. A. (ed.). 1975. Proceedings of the First 1. C. White Memorial Symposium:

the age of the Dunkard. West Virginia Geol. Econ. Surv., 352 pp.

Beerbower, J. R. 1963. Morphology, paleontology and phylogeny of the Permo-Car-
boniferous amphibian Diploceraspis. Bull. Mus. Comp. Zool., Harvard Univ.,
130:31-108.

Berman, D. S. 1978. Ctenospondylus ninevehensis, a new species (Reptilia, Pelyco-
sauria) from the Lower Permian Dunkard Group of Ohio. Ann. Carnegie Mus.,
47:493-514.

Berman, D. S., and S. L. Berman. 1975. Broiliellus hektotopos sp. nov. (Temno-
spondyli: Amphibia) Washington Formation, Dunkard Group, Ohio. Pp. 69-78, in
Proceedings of the First 1. C. White Memorial Symposium: the age of the Dunkard
(J. A. Barlow, ed.). West Virginia GeoL Econ. Surv., 352 pp.

Berryhill, H. L., Jr., and V. E. Swanson. 1962. Revised stratigraphic nomenclature
for Upper Pennsylvanian and Lower Permian rocks, Washington County, Penn-
sylvania. U. S. Geol. Survey Prof. Pap., 4500:43-46.

Case, E. C. 1908. Description of vertebrate fossils from the vicinity of Pittsburgh,
Pennsylvania. Ann. Carnegie Mus., 4:234-241.

Langston, W., Jr. 1953. Permian amphibians from New Mexico. Univ. California
Publ. Geol. Sci., 29:394-416.

Lund, R. 1972. Notes on the vertebrate fossils of the Elm Grove area. West Virginia.
West Virginia Geol. Surv. Guidebook of I. C. White Memorial Symposium Field
Trip, p. 51.

202

Annals of Carnegie Museum

VOL. 48

. 1975. Vertebrate-fossil zonation and correlation of the Dunkard basin. Pp. 171-

178, in Proceedings of the First I. C. White Memorial Symposium: the age of the
Dunkard (J. A. Barlow, ed.), West Virginia Geol. Econ. Surv., 352 pp.

. 1976. General geology and vertebrate biostratigraphy of the Dunkard basin.

Pp. 225-239, in The Continental Permian in Central, West, and South Europe (H.
Falke, ed.), D. Reidel Publ. Co., Dordrecht, Holland, 352 pp.

Moran, W. E. 1952. Location and stratigraphy of known occurrences of fossil tetra-
pods in the Upper Pennsylvanian and Permian of Pennsylvania, West Virginia, and
Ohio. Ann. Carnegie Mus., 33:1-112.

Olson, E. C. 1970. Trematops stonei sp. nov. (Temnospondyli: Amphibia) from the
Washington Formation, Dunkard Group, Ohio. Kirtlandia, 8:121-130.

1975. Vertebrates and biostratigraphic position of the Dunkard. Pp. 155-165,

in Proceedings of the First I. C. White Memorial Symposium: the age of the Dun-
kard (J. A. Barlow, ed.). West Virginia Geol. Econ. Surv., 352 pp.

Romer, a. S. 1948. Relative growth in pelycosaurian reptiles. Pp. 45-55, in Robert
Broom Commemorative Volume (A. L. Du Toit, ed.). South Africa Roy. Soc. Spec.
Publ., 257 pp.

. 1952. Late Pennsylvanian and Early Permian vertebrates of the Pittsburgh-

West Virginia region. Ann. Carnegie Mus., 33:47-113.

. 1960. The vertebrate fauna of the New Mexico Permian. New Mexico Geol.

Soc. Guidebook of Rio Chama Country, 11th Field Conf., pp. 48-54.

Romer, A. S., and L. I. Price. 1940. Review of the Pelycosauria. Geol. Soc, America
Spec. Paper, 28:1-538.

Vaughn, P. P. 1963. The age and locality of the late Paleozoic vertebrates from El
Cobre Canyon, Rio Arriba County, New Mexico. J. Paleontol., 37:283-286.

. 1964, Vertebrates from the Organ Rock Shale of the Cutler Group, Permian of

Monument Valley and vicinity, Utah and Arizona. J. Paleontol., 38:567-583.

— . 1966. Comparison of the Early Permian vertebrate faunas of the Four Corners

region and north-central Texas. Los Angeles County Mus. Nat. Hist., Contrib. Sci.,
105:1-13.

— . 1969. Early Permian vertebrates from southern New Mexico and their paleo-

zoogeographic significance. Los Angeles County Mus. Nat. Hist., Contrib. Sci.,
166:1-22.

WiLLiSTON S. W., AND E. C. Case. 1913. A description of Edaphosaurus Cope. Pp.
71-81, in (E. C. Case, S. W. Williston, and M. G. Mehl, eds.) Permo-Carboniferous
vertebrates from New Mexico, Carnegie Inst. Washington, 181:1-81.

Whipple, R. W., and E. C. Case. 1930. Discovery of Permo-Carboniferous verte-
brates in the Dunkard Formation of West Virginia. J. Washington Acad. Sci.,
20:370-372.

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ANNALS

0/ CARNEGIE MUSEUM

CARNEGIE MUSEUM OF NATURAL HISTORY
4400 FORBES AVENUE » PITTSBURGH, PENNSYLVANIA 15213
VOLUME 48 1 June 1979 ARTICLE 12

A NEW LIOLAEMUS (SAURIA, IGUANIDAE) FROM THE
HIGH ANDES OF ARGENTINA, WITH
ECOLOGICAL COMMENTS

Arthur C. Hulse^

Research Associate, Section of Amphibians and Reptiles
Abstract

A high elevation Andean Liolaemus is described as a new species. It appears to be
related to the elongatus group, but differs from all members of the group by having
generally lower scale counts. Comments are made concerning the ecology of the species.

Introduction

Liolaemus is a large (over 50 described species), ecologically diverse
genus of iguanid lizards inhabiting Austral, Patagonian, and Andean
regions of South America (Donoso-Barros, 1966). Although the genus
is widely distributed many species are localized endemics with re-
stricted ranges (Cei, 1974; Donoso-Barros, 1966, 1971; Donoso-Barros
and Cei, 1971). The species described here seems to conform to the
pattern of restricted or local distribution because it was only found in
a small region of the Cuesta de Minas Capillitas, Catamarca. Cata-
marca Province is herpetologically poorly known; Koslowsky (1895)
reported initially on the herpetofauna of the region, but little additional
work has been done in the province. Hellmich (1964) described Lio-
laemus robertmertensi from the mountains in the vicinity of Belen,
I Catamarca. The new species also comes from the Cordillera de Ca-

I

1 ^ Biology Department, Indiana University of Pennsylvania, Indiana, Pennsylvania

I 15705.

Submitted 27 November 1978.

203

204

Annals of Carnegie Museum

VOL. 48

Fig. 1. — Lateral view of head (A) and dorsal view of body (B) of holotype of Liolaemus

capilUtas.

tamarca; however, it was found in the vicinity of Andalgala (approx-
imately 90 air km east of Belen).

This study is based on material collected during 1973-1974 and 1974-
1975 while I was involved in the Desert Scrub Subprogram of the
International Biological Program.

Liolaemus capillitas, new species

//o/ory/7^.— Carnegie Museum of Natural History (CM) 70114, an
adult male from 5 km S of Minas Capillitas, about 3,900 m, Provincia
de Catamarca, Argentina, collected 5 January 1975 by Arthur C.
Hulse.

Paratopotypes .—CM 70115-70147.

Diagnosis.— A large Liolaemus possibly of the elongatus group,
with moderately long hind limbs; dorsal scales triangular, keeled, not
pointed; preanal pores, three or four; dorsum brownish to black, with-
out distinct pattern; differing from other members of the elongatus
group in possessing generally lower scale counts around the body, 58-

1979

Hulse — New Species of Liolaemus

205

Fig. 2. — Dorsal view (A) and ventral view (B) of head of holotype of Liolaemus capil-
litas.

67; fewer supralabials, five to seven; and fewer subdigital lamellae on
the fourth toe, 24-28.

Description of the holotype .—Size, large, general form robust; adpressed hindlimb

reaching to just behind the ear; tail one and one-half times snout- vent length; head
somewhat shortened, 1.3 times as long as wide (Fig. 1). Dorsal head scales variable in

206

Annals of Carnegie Museum

VOL. 48

Table 1. — Comparison of scale counts of Liolaemus capillitas with members of the
Liolaemus elongatus complex. Data on elongatus complex are from Cei (1974). Data
on L. capillitas represent mean, range, and standard error of the mean. NA = not
available in Cei (1974). N = 34 for L. capillitas.

Character

Liolaemus

capillitas

Liolaemus

elongatus

elongatus

Liolaemus

elongatus

petrophilus

Liolaemus

austromen-

docinus

SAB

61.5 (58-67 ± 0.47)

72-90

75-92

76

Supralabials

5.8 (5-7 ±0.11)

7-8

8

7

Infralabials

5.5 (5-7 ± 0,10)

NA

NA

6

Subdigital

lamellae

26.0 (24-28 ± 0.22)

27-30

25-32

28

Preanal

pores

3.4 (3-4 ± 0.18)

NA

NA

3

size, numerous, relatively small, usually smooth, and slightly convex; temporals weakly
keeled. Rostral 2.5 times as wide as high, visible from above. One azygous frontal
present; interparietal slightly smaller than parietals; three or four enlarged supraoculars;
subocular greatly extended, separated from supralabials by a single row of scales; six
supralabials; five infralabials. Supralabials, infralabials, and adjacent scales dotted with
small dark pits. Mental 2.5 times as wide as high; postmentals in two divergent rows of
five scales each (Fig. 2). Ear opening oval, twice as high as wide, bordered anteriorly
by moderately large scales and by granular scales along the other margins. Sides of neck
with granular scales, antehumeral fold present. Dorsal scales moderate in size, imbri-
cate, not pointed, conspicuously keeled. Ventral scales inbricate, smooth, slightly larger
than dorsals. Lateral scales imbricate, diagonally keeled, smaller than either dorsals or
ventrals, and often associated with small satellite scales, the distribution of which is
variable. Dorsal caudal scales rectangular, strongly keeled, imbricate, and pointed. Ven-
tral caudal scales smooth, rectangular, imbricate, directed diagonally inward, of a size
similiar to the dorsal caudals. Arm scales variably keeled, being most heavily keeled in
the dorsal antebranchial region. Thigh scales large and keeled anteriorly and small and
granular posteriorly. Scales around the body 61; 21 dorsal scale rows contained in the
length of the head; 27 fourth toe subdigital lamellae; three preanal pores.

Dorsal ground color brownish in the preserved animal. Dorsal pattern a series of
vague transverse bars produced by lighter colored scales interspersed among the dark
background scales. A definite darkening of the body occurs along the anterior sides and
along the upper surface of the arm. Head scales brown with some dark spotting. Ventral
coloration light bluish gray. Tail light brown, lacking a pattern.

Measurments of the holotype. — Snout-vent length, 89 mm; tail length, 143 mm; head
length, 22 mm; head width, 17 mm; hindleg length, 53 mm; foreleg length, 33 mm.

Variation. — The type-series consists of 11 adult males, 16 adult females, and seven
immatures. The sexes are easily distinguished by the presence of preanal pores in males.
General body form in the females is more slender; and the head is less massive.

Variation in scale counts is presented in Table 1. The azygous frontal is absent in
some individuals. The divergent postmental scale rows vary from three to five scales
per row. In some individuals the temporal scale keeling is absent or greatly reduced.
Ground color in some animals is black and in some the transverse bars are absent. Most
mature females possess slightly reddish areas in the preanal region and along the ventral
surface of the thighs. It is possible that the coloration may be associated with specific
sexual condition, such as the breeding coloration exhibited by female Crotaphytus and
Cophosaurus.

1979

Hulse-— New Species of Liolaemus

207

Table 2. — Stomach analysis of Liolaemus capillitas by percent volume and number of

food items. N = 28.

Food item

Number of items

Percent volume

Formicidae

417

59.7

Coleoptera

13

12.9

Orthoptera

6

8.0

Hemiptera

5

7.0

Nematodes

114

4.0

Lepidoptera

4

3.6

Plant matter

—

3.3

Diptera

3

1.5

Systematic remarks of the large number of species in the
genus and the scant taxonomic information available for many of the
species exact relationships are difficult to determine; however, L. cap-
illitas appears to be most closely related to the Patagonian lizards of
the elongatus group (L. elongatus elongatus Koslowsky, L. e. petro-
philus Donoso=Barros, and L. austromendocinus Cei). It differs from
the above species in having generally lower scale counts (Table 1).
Liolaemus capillitas is similiar in general appearance and body form
to descriptions of L. austromendocinus , but is more robust than either
form of L. elongatus and lacks the distinctive dorsal pattern usually
found in L. elongatus. It occurs farther north than any of the above
species.

Ecological remarks .-—Liolaemus capillitas is an inhabitant of high
elevations. The type-series was collected at approximately 3,900 m
and specimens were never observed below 3,500 m. Typical habitat is
steep rocky hillsides covered with bunch grass, scattered low shrubs,
and small cacti. Within this general habitat L. capillitas is almost en-
tirely restricted to rocky cliff faces and exposed road cuts. During mid-
day (1000 to 1600 hr) lizards can be seen sunning on the exposed rocks.
When disturbed they attempt to escape into the cracks and crevices
in the rocks. The restriction of activity to mid-day is most likely a
function of the exposure of the rock faces where the lizards were
found. During early morning and late afternoon the faces were in shad-
ow and offered no opportunity for behavioral thermoregulation.

Stomach contents of 28 lizards were examined (Table 2). Ants com-
posed almost 60% of the total volume and represented about 80% of
the individual food items ingested. Coleoptera, Orthoptera, and He-
miptera made up a significant percentage of the remaining items. Plant
material (probably incidentally ingested) represented 3.3% of the total
volume. The lizards were heavily parasitzed by gastric nematodes. A
total of 114 nematodes was found in the stomachs of the lizards and

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VOL. 48

Fig. 3.— -Relationship between snout-vent length and clutch size in Liglaemus capillitas.

comprised 4% of the total stomach volume. Only one lizard was free
of the parasites.

Seven females contained six to 11 (mean = 8.0) oviducal eggs. There
appears to be a trend for larger lizards to have larger clutches and for
the smallest clutches to be found in the smallest lizard (Fig. 3). Spec-
imens collected in November and January contained oviducal eggs,
whereas those collected in February were spent, and contained at most
small corpora lutea. The lizards seem to be viviparous as the oviducal
eggs from November and January samples contained developing em-
bryos. The marked absence of females bearing oviducal eggs in the
February sample poses some interesting questions. Are the lizards
truly viviparous or do they simply retain the eggs in the oviducts for
varying lengths of time before oviposition? The relatively small size
difference between embryos from the November and January samples
suggests a slow developmental rate if viviparous. It would seem highly
unlikely that the eggs could have developed to parturition between 5
January and 12 February. This suggests that the eggs are merely re-
tained in the oviducts for a period of time before deposition. Further
support for this explanation of the reproductive strategy comes from
the heavy, leathery eggshell that is identical to that of oviparous
forms. Furthermore, hatchlings were not present in the February sam-

1979

Hulse — New Species of Liolaemus

209

pie and would have been expected to be in the sample if the lizards
were indeed viviparous and had given birth. The phenomenon of egg
retention has been noted in other high elevational lizards such as Sce-
loporus scalaris (Newlin, 1976) and Anolis cybotes (Huey, 1977).

The largest immature male collected was 70 mm SVL and the small-
est mature male was 79 mm. The largest immature female was 72 mm
and the smallest mature female was 75 mm. The largest individual
collected was a 93 mm female.

Etymology. — The specific epithet capillitas refers to the locality where the type-series
was collected.

Acknowledgments

Fieldwork in Argentina was supported by NSF Grant 06-27125 to the University of

Texas, Austin, for the Desert Scrub Subprogram of the Origins and Structure of Eco-
systems Program of the International Biological Program. I thank Drs. R. G. Zweifel

and G. R. Zug for allowing me to examine specimens of Liolaemus in their care.

Literature Cited

Cei, J. M. 1974. Revision of the Patagonian iguanids of the Liolaemus elongatus com-
plex. J. HerpetoL, 8:219-229.

Donoso-Barros, R. 1966. Reptiles de Chile. Ed. Univ. Chile, Santiago, 1-485 -I- i-
cxlvi pp.

. 1971. A new Liolaemus from Neuquen (Argentina). Herpetologica, 27:49-51.

Donoso-Barros, R., and J. M. Cei. 1971. New lizards from the volcanic Patagonian
plateau of Argentina. J. HerpetoL, 5:89-95.

Hellmich, W. 1964. Uber eine neue Liolaemus Art aus den Bergen von Catamarca,
Argentinien (Reptilia, Iguanidae). Senckenbergiana Biol., 45:505-507.

Huey, R. B. 1977. Egg retention in some high altitude Anolis lizards. Copeia, pp. 373-
375.

Koslowsky, j. 1895. Batracios y reptiles de Rioja y Catamarca. Rev. Mus. La Plata,
6:357-370.

Newlin, M. E. 1976. Reproduction in the bunch grass lizard, Sceloporus scalaris.
Herpetologica, 32:171-184.

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ANNALS

o/ CARNEGIE MUSEUM

CARNEGIE MUSEUM OF NATURAL HISTORY
4400 FORBES AVENUE * PITTSBURGH, PENNSYLVANIA 15213
VOLUME 48 7 SEPTEMBER 1979 ARTICLE 13

GNATHORHIZA BOTHROTRETA (OSTEICHTHYES: DIPNOI)
FROM THE LOWER PERMIAN ABO FORMATION OF
NEW MEXICO

David S Berman

Associate Curator, Section of Vertebrate Fossils
Abstract

On the basis of additional specimens, the skull roof and tooth plates of the lungfish
Gnathorhiza bothrotreta Berman (19766) from the Lower Permian Abo Formation of
New Mexico are more completely described and compared with those of other Lower
Permian species of this genus. Many of the distinguishing features of G. bothrotreta
suggest that it represents a widely divergent line that separated from the evolutionary
line, or closely paralleling lines, including the other Early Permian species of Gnatho-
rhiza during the Pennsylvanian or earlier. Recent advancements in our knowledge of
Gnathorhiza deny its possible role as an ancestor to the modern lepidosirenid lungfish.

Introduction

A new species of the lungfish Gnathorhiza, G. bothrotreta, was
described on the basis of partially articulated specimens preserved in
aestivation burrow casts from the Lower Permian Abo Formation of
New Mexico (Berman, 1976^). Prior to that report, tooth plates
(Vaughn, 1966, 1969, 1970) and sedimentary objects resembling casts
of aestivation burrows (Vaughn, 1964) from the Lower Permian of
Utah and New Mexico had been assigned to Gnathorhiza without
specific designation. In the description of G. bothrotreta (Berman,
1976^) the available specimens did not allow a complete description
of the skull roof and tooth plates. The discovery of additional sped-
mens of this species from the type locality now permits not only a

This research and publication were funded by the M. Graham Netting Research Fund
through a grant from the Cordelia Scaife May Charitable Trust.

Submitted for publication 6 March 1979.

211

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VOL. 48

complete description of the skull roof and tooth plates, but also a more
detailed comparison with other Early Permian species of Gnathorhiza.

In addition to G. bothrotreta, four other species of this genus have
been recognized. G. pusillus, from the Upper Pennsylvanian of Illi-
nois, is based on an upper tooth plate (Cope, 1877), so there is little
morphological grounds for comparison with other species of Gnatho-
rhiza. The remaining three species, G. serrata, G. dikeloda, and G.
noble nsis, have been identified from Lower Permian deposits of the
northeastern United States (except G. noble nsis; Lund, 1975, 1976)
and the midcontinental states of Texas and Oklahoma. Only in the
latter region are they known from skulls rather than from tooth plates
alone. Using disarticulated and unassociated elements from the Vale
Formation of Texas, Olson (1951) attempted the first restoration of the
skull of this genus in his description of a new species, G. dikeloda.
Though his restoration contains some minor errors or omissions, it is
valid in its gross aspects. G. serrata was first recognized on the basis
of tooth morphology nearly a century ago (Cope, 1883), yet its skull
was not described until comparatively recently (Carlson, 1968). Partial
skulls preserved in aestivation burrow casts from the Wellington For-
mation of Oklahoma provided the basis of the accurate reconstruction
of its skull. G. noble nsis (Olson and Daly, 1972) includes specimens,
some of which are partial skulls, from the Hennessey Formation of
Oklahoma; only specific areas of the skull have been illustrated.

A large skull of Gnathorhiza, restored from disarticulated remains
preserved in an aestivation burrow cast from the Arroyo Formation of
Texas, was not referred to a previously described species because of
structural and proportional differences of the skull (Berman, 1976<3).
It was considered equally unwise to assign it to a new species because
almost all of the differences might be the result of widely separated
growth stages. This skull, CM 26576, about 12.5 cm in midline length,
is about four times larger than any other known Gnathorhiza skull and
many of its seemingly unique features may reflect an advanced growth
stage. Despite this taxonomic uncertainty, CM 26576 is important to
this discussion because it is known in great detail and exhibits features
that make it quite distinct from G. bothrotreta.

Systematic Paleontology

Class Osteichthyes
Order Dipnoi

Gnathorhiza bothrotreta Berman, 1976

Revised diagnosis.— ThQ following features distinguish Gnathorhiza
bothrotreta from other known species of this genus. The supraorbital
canals: 1) enclosed in bone K-M and open to surface via a series of

1979

Berman — New Mexican Lungfish

213

six, possibly seven, large, oval apertures; 2) have a much longer extent
on bone E+F, where they extend about three times farther anteriorly
than posteriorly from medialmost level of their arch-like, inward flex-
ure. Bone X fused with Yj+Yg to form compound element X+Y,
which 1) contains three-way junction of infraorbital, supraorbital, and
main canals, 2) possesses a very narrow and attenuated descending or
opercular component, and 3) extends to posterolateral corner of skull
roof and contains junction of main canal and its occipital commissure.
Pronounced shortening of posterior portion of skull roof has resulted
in 1) transverse widths of bones B and I exceeding their longitudinal
lengths by nearly two times and more than two times, respectively,
and 2) portion of opercular chamber bordered by X+Y being very
shallow in anteroposterior depth. Transverse width of bone C is about
75% of its longitudinal length. Posteriormost blade of prearticular
tooth blade consists of a smooth, proximal ridge from whose distal end
two rows of four and five large denticles radiate and have a single,
large denticle lying between them distally.

Holotype. — CM 26558 (collections of the Carnegie Museum of Nat-
ural History, Section of Vertebrate Fossils), a nearly complete skull
roof with numerous fragmentary ribs and left lower tooth plate closely
associated (see Berman, 19766, for description).

Referred specimens. — CM 26551 , fragments of skull and postcranial skeleton, both
pterygoid tooth plates, left prearticular tooth plate and left vomerine tooth; CM 26560,
fragments of skull; CM 30741, nearly complete skull with closely associated but poorly
preserved postcranial debris; CM 35884, left prearticular tooth plate.

Horizon and locality.— Ml specimens from lower part of Abo Formation, Lower
Permian, SE^A sec. 22, T2N, R4E, Socorro County, New Mexico, except CM 30764
from SE!4 sec. 36, T6N, R3W, Valencia County.

Description

The skull roof of Gnathorhiza bothrotreta is well represented in only two specimens,
the holotype, CM 26558, and the referred specimen CM 30741 found subsequent to the
description (Berman, \916b) of the type. Though the skull roofs of both specimens are
incomplete, together they complement each other in making possible the restoration
seen in Fig. 5. The skull CM 30741 was preserved in a dense, extremely hard limestone
and was exposed, for the most part, by fracturing away the surrounding matrix in small
blocks, exposing it as seen in Fig. 1. Unfortunately, much of the skull roofing bone
spalled off with the enclosing blocks of matrix. Though, as indicated in Fig. 2, the
general outline of the skull is preserved, most of the sutures and lateral lines have been
obscured or lost. As in the study of the type skull (Berman, 19766), most of the sur-
rounding blocks of matrix were reassembled into a single, counterpart-like block. The
adhering bone was removed from the counterpart block so as to create a natural mold
of the outer surface of the skull roof. From this, the silastic rubber cast seen in Fig. 3
was made; it faithfully reproduces many of the structural details of the skull roof, which
are indicated in Fig. 4. The right, posterolateral region of the skull roof has been crushed
inward somewhat with the loss or disarticulation of some elements. The remainder of
the skull roof, however, is well articulated and appears to retain its proper curvature.

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VOL. 48

Fig. l.~~Gnathorhiza bothrotreta, referred specimen CM 30741. A-C, dorsal, lateral,
and occipital views of skull roof.

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215

Fig. 2.— =A-C, outline sketches of views A-C of Fig. 1 showing sutures and supraorbital
canal (soc), B, C, E+F, I, J, K-M, and X+Y are skull roof bones.

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1 I

Fig. 3. — A~C, dorsal, lateral, and occipital views of silastic rubber cast made from
natural mold of skull roof of Gnathorhiza bothrotreta CM 30741.

1979

Berman— New Mexican Lungfish

217

Fig, 4.— A-C, outline sketches of views A-C of Fig. 3 showing sutures and lateral-line
canals. Abbreviations: clc, main canal commissure; ioc, infraorbital canal; Ic, main
canal.

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VOL. 48

The midline length of the skull roof is 34 mm and its greatest transverse widths measured
at the level of bone X+ Y, is about 24 mm.

The occipital commissure of the main lateral-line canal extends along the very pos-
terior edge of B, gently curves anteriorly as it follows the occipital margin of I, then
swings abruptly anteriorly upon entering X+Y as the main lateral line. In X-l-Y and a
short distance from its contact with K-M, the supraorbital and infraorbital canals branch
off the main canal. The infraorbital canal, seen for the first time in this species in CM
30741, extends ventrally a short distance to exit at a very small, angular notch located
just above the midheight of the free anterior margin of X+Y. The supraorbital lateral
line of X+Y is restored as a groove, but on entering K-M it becomes enclosed within
that bone and communicates with the surface via a row of six, possibly seven, evenly
spaced oval apertures. Arching slightly medially as it extends anteriorly near the lateral
border of K-M, the supraorbital lateral line leaves the skull roof just posterior to the
contact of K-M with E+F. Only some of the supraorbital apertures are detectable on
CM 30741 and these are poorly preserved; in the restoration of Fig. 5 their size, number,
and arrangement are based on the type, where they measure 1.1 mm in height and 0.8
mm in width. In the description of the type (Berman, 19766) an aperture was also noted
at the anterior end of the lateral-line canal extending anteroposteriorly across the right
Y1+Y2 (X+Y here), thought then to be the main canal but now recognized as part of
the supraorbital canal. On the basis of CM 30741, it is now believed that the supraorbital
canal of this bone is of the normal channel structure. The supraorbital canals of the
rostral bone E+F (E of most authors) of CM 30741 are complete enough to restore their
entire pattern. Entering E+F at about its midlength, the supraorbital canal turns abruptly
anteriorly as it nearly reaches the midline, then continues with a very slight lateral
curvature to reach the extreme anterior end of the lateral border of E + F. Anterior and
posterior pit lines cannot be seen in either the type or referred skulls.

Bone B is rectangular in outline and its transverse width is nearly two times its
longitudinal length. The referred skull CM 30741 exhibits the same wide jog in the
midline suture between the paired C bones seen in the type (Berman, 19766). E+F is
complete in CM 30741, but only its posterior third is preserved in the type. Widest at
its posterior end, E + F narrows by about a third at its midlength, then tapers to about
one-half its maximum width at its bluntly rounded, rostral end. Bone I has a narrow
exposure along the occipital margin of the skull roof and its transverse width is over
two times its longitudinal length. Bone I does not extend laterally to the posterolateral
corner of the skull roof. From the medial half of the occipital margin of I a rectangular
flange projects steeply posteroventrally and slightly medially to form the lateral side of
the foramen magnum.

The X+Y bone is best represented on the left side of skull CM 30741, where it is
essentially complete; it is best described as consisting of basically two components: 1)
a transversely narrow component that is contained mainly in the dorsal platform of the
skull roof, and 2) a long, narrow, descending, or opercular, component that is directed
nearly vertically and slightly antero ventrally, and separates the opercular and orbital
chambers. The dorsal component extends posteriorly to just include the posterolateral
comer of the skull roof and, therefore, appears to contain the junction between the main
lateral line and its occipital commissure. A very small, rounded process projects pos-
teriorly from the occipital margin of X+Y near its contact with I; the process is poorly
preserved in both the type and CM 30741. The posterior wall of the lateral-line canal
immediately anterior to the small occipital process of X+Y is very low and most likely
marks the point where the main lateral line of the body entered the skull roof. The
occipital process of X+Y apparently lies below the level of the dorsal surface of the
skull and was not scored by the main lateral line.

Both pterygoid tooth plates and the left vomerine tooth of the palate, and the left
prearticular tooth plate of the lower jaw are preserved in CM 26557; the right pterygoid
and the left prearticular tooth plates are essentially complete and shown in Fig. 6.

1979

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219

Fig. 5.— A, dorsal, and B, lateral views of reconstructed skull roof of Gnathorhiza
bothrotreta based on type CM 26558 and referred specimen CM 30741. Approximately

x2.

Because tooth plates have been used for differentiation of not only Gnathorhiza species,
but species belonging to genera believed to be closely related to Gnathorhiza on the
basis of tooth plate morphology, those of CM 26557 are described here in some detail.
The right pterygoid tooth plate, the more complete of the pair, measures about 11.6 mm
in greatest length and is typical of the genus in consisting of four radiating blades, which
are numbered in Fig. 6 for convenience of description. Blade 1, the anteriormost and
longest, measures about 9.0 mm long from its juncture with blade 2 and possesses only
two distinct cusps at its distal end. The blade is directed anteriorly along the margin of
the symphysial contact of the paired pterygoids (more correctly entoptery golds). Blades
2, 3 and 4 are about 4.0, 4.0 and 5.0 mm long. The proximal halves of the blades are
worn smooth and distally only worn remnants of cusps are present, making it impossible

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VOL. 48

Fig. 6.—Gnathorhiza bothrotreata. CM 26557, A, right pterygoid tooth plate with tooth
blades numbered, and B, left prearticular tooth plate. Labial is to the left and anterior
is to the top.

to give accurate counts of their numbers. The lower tooth plate consists essentially of
three blades radiating from a single point and has a greatest length of about 12.0 mm.
The longest blade, about 8.8 mm, extends anteriorly; it exhibits no cusps due to wear.
The central, anterolaterally directed blade measures 4.4 mm in length and possesses
three distinct cusps at its distal end. The posteriormost blade, measuring about 5.6 mm
long, is directed laterally. This blade consists of a proximal smooth ridge, at the end of
which two rows of large denticles narrowly diverge; the more anterior row consists of
four denticles and the other five. The denticles increase in size distally and most of the
denticles of each row only barely contact one another basally. Lying between the distal
ends of the two rows of denticles is a single, large denticle. The denticles of the distal
portion of the posterior blade sit on a flat surface rather than arise from discrete blades.
Only two other lower tooth plates, both of the left jaw, can be noted here; one belongs
to the type and the other, CM 30764, was found isolated. They resemble the lower plate
of CM 26557 except that the denticles of the distal portion of their posterior blades have
been lost due to preparation in the former and wear in the latter. Of the vomerine tooth of
CM 26557, only the greater part of the narrow, fan-shaped crown is preserved; the base
of the tooth is lost. In life the crown had a greatest length of about 4.5 mm. The crown,
formed by a flat inner surface and a slightly convex outer surface, thins somewhat as
it decreases in height posteriorly. The worn cutting edge possesses no denticles.

1979

Berman — New Mexican Lungfish

221

Fig. 7. — Diagrammatic restorations of the skull roofs of Gnathorhiza sp. CM 26576 in
A, dorsal, and B, lateral views (redrawn from Berman, 1976a) and Gnathorhiza s errata
in C, dorsal, and D, lateral views (redrawn from Carlson, 1968). 1, 4, 5, 6, and 7 are
numbered, circumorbital bone series. Abbreviations: apl, anterior pit line; poc, pre-
opercular canal; ppl, posterior pit line; qp, quadrate process. Not drawn to scale.

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Comparison

Gnathorhiza bothrotreta possesses a number of features that easily
distinguish it from other members of this genus. Not least among these
is the enclosure of its supraorbital laterahline canal within bone K-M
to form a duct that opens to the surface via six, possibly seven, rela-
tively large, evenly spaced, oval apertures; in all other species of Gna-
thorhiza the lateral line of this bone is of the expected, gutter-like
structure. Further, in G. serrata, G. dikeloda, and CM 26576 the su-
praorbital canal has a short, simple, arch-like inflection near the an-
terior end of the lateral margin of E+F (Fig. 7A, B), whereas in G.
bothrotreta this pattern differs in that the canal is much longer and
extends approximately three times farther anteriorly than posteriorly
from the medialmost point of its inward flexure on E+F.

The fate of bone X in G. bothrotreta appears to have been different
from that in other species of Gnathorhiza. Bone X is identified as a
small element, having a posteromedial contact with J and containing
the junction of the main and the infraorbital canals (Thomson and
Campbell, 1971). In CM 26576 the junction of the main and infraorbital
canals lies at the sutural intersection of K^M, 4, and Yi+ Yg (Fig. 7A,
B). This was interpreted (Berman, 1976fl) to mean that X had probably
been eliminated by the invasion of at least the latter two elements
rather than by fusion with a neighboring bone. In G. serrata, Carlson
(1968) illustrates a very small, curved notch at the lateral margin of the
contact of bones Yi+ Yg and K-M that creates a narrow break in the
lateral-line system where undoubtedly the supraorbital and infraorbital
canals would have branched off the main canal (Fig. 7C, D). It is not
likely that this notch was occupied by a bone such as X and the lateral-
line pattern in G. serrata would seem to indicate that the X bone was
also lost by invasion, most probably by Yj+Yg. The situation is not
clear in either G. dikeloda (Olson, 1951) or G. noblensis (Olson and
Daly, 1972). The fact that in G. bothrotreta the junction of the infraor-
bital with the main canal lies well within the boundaries of the bone
typically referred to as Yi+Yg is reason to believe that this bone has
fused with bone X. The junction of the two canals also lies in the
approximate area where the X bone is normally located.

The compound bone Yj+Yg+X of G. bothrotreta, referred to here
as simply X+Y, exhibits distinguishing features that are not the direct
result of the inclusion of X. X+Y extends posteriorly to the postero-
lateral corner of the skull roof and, therefore, not only appears to
capture a very small portion of the occipital commissure of the main
lateral line, but also the area in which the main lateral line of the body
entered the skull roof. In G. serrata, G. noblensis, and CM 26576, the
occipital commissure of the main lateral line is contained within bones

1979

Berman— New Mexican Lungfish

223

B and I and the main lateral-line canal of the body appears to have
entered the skull roof at the lateral edge of I near its contact with
Yi+Yg. The descending, or opercular, component of Y1+Y2 of CM
26576 and X+Y of G. bothrotreta extend a considerable distance an-
teroventrally and, as indicated in the description of the former (Ber-
man, 1976fl), their posterior margin was probably closely bordered by
the upper half of the anterior margin of the operculum. The opercular
component of G. bothrotreta differs from that of CM 26576 in being
much narrower and in lacking the small, elongate quadrate process on
its postero ventral border. In both forms the opercular component, or
at least its distal portion, was undoubtedly applied against the lateral
surface of the quadrate cartilage and, thus, provided strength to the
jaw articulation. According to Carlson (1968), the anterolateral corner
of Yi+Yg of G. s errata is expanded somewhat anterolaterally and
projecting anterolaterally from its ventral surface is a narrow process,
convex on the outer surface, and flat on the inner surface, which
sheathed the outer surface of the cartilaginous suspensorium. The an-
terolateral corner of Y1+Y2 in G. serrata, therefore, resembles in gen-
eral the opercular component and its quadrate process of this bone in
CM 26576.

Compared to G. serrata, G. dikeloda, G. noblensis, and CM 26576,
there is a pronounced, relative shortening of the posterior portion of
the skull roof of G. bothrotreta that involves mainly bones B and 1.
In the New Mexico form, the transverse widths of B and I exceed their
longitudinal lengths by nearly two times and more than two times,
whereas in the other forms the longitudinal lengths greatly exceed the
transverse widths. As a result of this shortening, the portion of the
opercular chamber bordered by the skull roof in G. bothrotreta, spe-
cifically bone X+Y, is also much shallower in anteroposterior depth
than in the other Gnathorhiza species.

In Gnathorhiza the presence of the series of circumorbital bones 1
and 4 through 7, carrying the infraorbital canal and a short branch of
the preopercular canal, has been reported only in CM 26576 (Fig. 7A,
B). Because these elements are very small and contact each other by
simple abutment sutures, they could have been easily overlooked or
lost in most specimens. Further, because CM 26576 is much larger
than the Gnathorhiza specimens studied by other workers and, there-
fore, may represent an advanced stage of growth, it is also possible
that its circumorbital bones were detectable merely as the result of
more complete ossification. Carlson (1968) described a small indenta-
tion on the lateral edge of bone K-M immediately posterior to the
curved rim of the orbit in G. serrata (Fig. 7C, D) that he reasoned
might be associated with the articulation of the lower jaw. It is possible
that the first of the numbered circumorbital series, bone 4, might have

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Annals of Carnegie Museum

VOL. 48

articulated here. No elements have been found in G. bothrotreta that
can be referred to the numbered circumorbital bones; however, the
very small, angular notch at the exit of the infraorbital canal from
X+Y could conceivably, though not likely, have held the first element
of this series.

In G. bothrotreta the transverse width of bone C is about 15% of its
longitudinal length, whereas in G. serrata, G. noblensis, and CM 26576
the width is about 32 to 36% of the length. In G. dikeloda, Olson (1951)
describes a single, median D element in place of the paired C bones.
In this species one-half of the transverse width of bone D is about 42%
of its longitudinal length.

The tooth plates and vomerine tooth of G. bothrotreta conform to
the general pattern seen in all species of Gnathorhiza except for one
major difference. The denticulated, heeblike structure of the distal
portion of the posteriormost blade of the lower tooth plate is an unique
feature of this species.

Discussion

The skull roof and tooth plates of G. bothrotreta exhibit a number
of features that set it widely apart from all other Early Permian mem-
bers of this genus, which in general exhibit a strong conformity in
these structures. Some of the distinguishing features of G. bothrotre-
ta are of particular interest because they strongly indicate that it rep-
resents an early, widely divergent offshoot from the evolutionary line
or closely paralleling lines that comprise the other Early Permian
species of Gnathorhiza. Undoubtedly the most interesting of its unique
features is the structure of the supraorbital lateral-line canal of bone
K-M. The enclosure of the supraorbital canal is most easily explained
as the retention of a primitive feature, because it is well known that
among the dipnoans there has been a general evolutionary trend for
the dermal skull bones to become increasingly more deeply seated in
the skin and less associated with the lateral-line system (Westoll,
1949). The enclosure of only a specific portion of the cranial lateral-
line system, however, does appear to be a feature not seen in other
dipnoans, and the alternative interpretation that it is a secondary mod-
ification cannot be ruled out entirely. It can be said, however, that the
precise arrangement and number, and the uniform size and dimensions
of the apertures by which the supraorbital canal of G. bothrotreta
communicates with the exterior is a feature not duplicated in other
lungfish, fossil or Recent. Large supraorbital canal pores are described
in Megapleuron zangerli from the Middle Pennsylvanian of Illinois
(Schultze, 1977), however, they are randomly arranged along the canal
course and exhibit varying dimensions and shapes. In the Pennsylva-
nian-Lower Permian Conchopoma, the lateral-line system of the skull

1979

Berman — New Mexican Lungfish

225

is entirely enclosed (Schultze, 1975), but the pores are relatively small
compared to G. bothrotreta and randomly distributed along the course
of the canals. The overall structure of the supraorbital canal of K-M
in G. bothrotreta does approach the condition seen in actinopterygian
fishes and on the basis of this similarity was interpreted by me (Ber-
man, 19761?: 1037) as an unique refinement among the dipnoans of the
sensory function of the lateraMine system, “whose primary function,
it is generally agreed, is to detect currents or vibrations (sound) of low
frequency in the water in order to aid the fish in navigation, seeking
prey, avoiding predators, or communication.” However, it has been
pointed out to me (Lund, personal communication) that the lateraMine
system in fish does not function in directional hearing in the traditional
sense, but is a boundary layer detection system that responds to water
displacements and functions as a directional locator.

The denticle-bearing, heel-like structure of the posterior blade of the
lower tooth plate of G. bothrotreta is viewed as probably either indi-
cating a specialized function of its tooth plates or simply representing
an alternative, structural pattern for accomplishing the same function
as those of other members of this genus. During the early evolution of
the lungfishes there was a trend among many independent phylogenetic
lines toward reduction of the number of radiating rows of denticles of
the tooth plates and for the separate denticles of each row to become
confluent and wear into continuous ridges; Gnathorhiza represents an
advanced stage of this trend (Denison, 1974). The two narrowly di-
verging rows of denticles, as well as the denticle lying between them
distally, that form the distal portion of the posterior blade of the lower
tooth plate of G. bothrotreta, may have originally been parts of two,
or possibly three, separate radiating rows of denticles. Further, the
denticles, for the most part, barely meet at their bases and are sup-
ported by a flat surface rather than arise from a ridge or blade. The
unusual structure of the posterior blade of G. bothrotreta is not inter-
preted as a transitional stage leading to the single-ridged condition seen
in other species of Gnathorhiza, because it is not likely that the general
trend toward reduction of the number of radiating rows of denticles
was accomplished by a simple merging of the rows. As in other species
of Gnathorhiza, the posterior blade of the prearticular tooth plate of
G. bothrotreta inserted between the posterior two blades of the op-
posing pterygoid tooth plate during occlusion (Berman, 1968).

Two features of the skull of G. bothrotreta reflect a more advanced
level of structural organization over that of other Gnathorhiza species.
In the evolution of the dipnoans there has been a general trend toward
shortening of the posterior region of the skull. Within the genus Gna-
thorhiza this trend has been carried to a greater extreme in G. both-
rotreta mainly by a shortening of bones I and B; this, in turn, has

226

Annals of Carnegie Museum

VOL. 48

resulted in a much shallower, posterior emargination of the cheek re^
gion of the skull that accommodates the operculum. The apparent ab-
sence of the numbered series of circumorbital bones in G. bothrotreta
is also in accord with general evolutionary trend within the dipnoans
of the reduction in the number of dermal elements of the skull (Westoll,
1949); this feature of G. bothrotreta is judged as a marked advance-
ment over the condition seen in CM 26576 (Fig. 7A, B). As already
noted, there is reason to believe that such a series may have been
present in G. s errata, emanating from bone K-M rather than from
Y1+Y2 as in CM 26576. The skulls of G. dikeloda and G. noblensis
are too poorly known to speculate whether they possessed these ele-
ments.

If one accepts the argument that in G. bothrotreta bone X has fused
with Y1+Y2, whereas in all other species of this genus it has been lost
by invasion of neighboring bones, then it follows that the ancestral
form to the lines leading to the Early Permian Gnathorhiza species
must have possessed an unfused X bone. A separate X bone is typical
of Devonian lungfishes, but is extremely rare in late Paleozoic forms;
the only examples of the latter group definitely known to me include
the Early Pennsylvanian to Early Permian Cte nodus, in which a sep-
arate X bone is always present, and the Mississippian to Early Permian
Sagenodus, in which it is normally fused with Y2 and only rarely
separate (Westoll, 1949). Therefore, the phylogenetic line or lines lead-
ing to the Early Permian Gnathorhiza species of the midcontinent and
eastern United States, on the one hand, and that leading to G. both-
rotreta on the other, most likely extended back to the Pennsylvanian
or earlier. Further, because the presence of a separate X bone can be
considered as a very primitive feature, it seems also reasonable to
assume that the taxonomic gap between the Early Permian Gnatho-
rhiza species and their common ancestral form was probably generic
rather than specific. To date, there is no genus in which the skull is
known that can be reasonably considered as directly ancestral to Gna-
thorhiza. It should also be pointed out that the specimens of G. both-
rotreta from the Abo Formation occur at an earlier time, Wolfcampian
(earliest Permian), than the Gnathorhiza species of the midcontinent
(Olson and Vaughn, 1970) and the eastern United States (Lund, 1975;
Olson, 1975), which are of Leonardian age (later Early Permian). In
light of this, the possession by G. bothrotreta of features that indicate
a greater advancement over other Gnathorhiza species helps to rein-
force the idea that it had an independent evolutionary history during
the Early Permian and earlier.

Recently, Lund (1970, 1973, 1975) has described two species of new
genus, Monongahela stenodonta and M. dunkardensis, on the basis
of very small tooth plates and their supporting jaw elements, and vo-

1979

Berman— New Mexican Lungfish

227

merine teeth from the Upper Pennsylvanian and Lower Permian of
southwestern Pennsylvania. On the basis of chronological changes in
tooth plate morphology M. stenodonta is envisioned as the parent
stock of not only M. dunkardensis and an unnamed species of Mo-
nongahela, but of Gnathorhiza serrata and G. dikeloda. Despite the
clear similarity between the tooth plates of M. stenodonta and Gna-
thorhiza species in general, an ancestor-descendant relationship based
on dentitions alone can only be viewed as tentative in light of recent
studies. Thomson and Campbell (1971:100) have cast doubt on the use
of dentition to distinguish phyletic lines among the dipnoans, arguing
that “The character of the teeth is clearly related to mode of feeding.
There seems to be no inherent reason why animals of a single stock
should not diversify their feeding habits and therefore their dentition
.... Given our present knowledge of the Dipnoi, the nature of the
dentition alone is not necessarily a reliable guide to phylogenetic re=
lationship . ...” In a detailed discussion of the evolution of dipnoan
dentition based on gross and histological structure, Denison (1974:53)
responded to their remarks by commenting that “teeth are adaptive,
of course, but surely this is true also of the pattern of the skull roof
and the form of the parasphenoid to which they [Thomson and Camp-
bell, 1971] give great phylogenetic weight. Similar dental apparatuses
or skull roof patterns may possibly have evolved separately and con-
vergently in different phyletic lines, so in our present state of knowl-
edge a single character cannot be accepted as proof of phyletic rela-
tionship.” Most important to the discussion here, however, are his
conclusions (p. 56) regarding the histology and manner of growth of
the tooth plates of Gnathorhiza, Monongahela and the two modern
lepidosirenid lungfish, Protopterus of Africa and Lepidosiren of South
America: “In most genera the tooth plates enlarge by the addition of
new denticles of trabecular or tubular dentine at the distal ends of the
ridges. One of the modifications that led to the origin of a new family,
the Lepidosirenidae, was the formation in the tooth plates of columns
of very hard dentine (‘petrodentine’) surrounded by softer trabecular
dentine, a condition foreshadowed by juvenile tooth plates of Monon-
gahela .... The late Paleozoic Gnathorhiza has been considered to
be related to and ancestral to the Lepidosirenidae, but the histology
of the tooth plates does not support this.” Though Denison’s obser-
vations allow the possibility of a Monongn/ze/fl- lepidosirenid ances-
try, they also strengthen other lines of evidence that refute the long-
accepted misconception of a Gnnt/ior/iizn -lepidosirenid ancestry.

Despite advancements in our knowledge of Gnathorhiza, there per-
sists the notion that it is ancestral to the Lepidosirenidae. This rela-
tionship was originally based on two lines of evidence: 1) their sup-
posed similarities in tooth plate design and function (Romer ard Smith,

228

Annals of Carnegie Museum

VOL. 48

1934), and 2) their exclusive ability to aestivate (Romer and Olson,
1954). A G nathorhiza Atpidosiitnid ancestry was denied by me (Ber™
man, 1968) on the basis of a study of the tooth plates, vomerine teeth
and lower jaws of Gnathorhiza, probably G. serrata, from the Lower
Permian of Texas, It was demonstrated that with regard to these struc-
tures, Gnathorhiza is in close accord with other late Paleozoic dipno-
ans, particularly Sagenodus, and with the Recent Neoceratodus of
Australia. Further, it was argued that the general resemblance of Gna-
thorhiza and Sagenodus to Neoceratodus in these structures suggests
that these two extinct genera were primarily herbivorous, as is Neo-
ceratodus. It was also pointed out that the similarities between the
tooth plates of Gnathorhiza on the one hand, and Protopterus and
Lepidosiren on the other are superficial. Close examination indicates
that the dentitions of the two living genera have undergone extensive
remodeling away from the typical dipnoan condition, suiting them for
a type of actively predaceous existence not suspected in any other
dipnoan group. These observations also hold true for the dentition of
Monongahela.

The cranial osteology of the Lepidosirenidae is so highly modified
that it offers little or no opportunity for comparison with other lungfish.
One unique feature of the lepidosirenid skull is its obvious kinesis; all
other dipnoans have akinetic skulls. It seems reasonable to speculate
that cranial kinesis in the lepidosirenids is related, at least in part, to
their actively predaceous habits. As an example, both Protopterus and
Lepidosiren possess a pair of slender, conical, sharp-pointed and
slightly recurved teeth that project downward from the anterior end of
the anteriormost medial dorsal roofing bone, referred to as the inter-
maxillary bone by Owen (1839) and the dermal ethomoid by Bridge
(1898) in Lepidosiren. The posterior margin of this bone is joined to
the skull by tough fibrous tissue. The snout, with its fang-like teeth,
was probably capable of elevation and depression, allowing the si-
multaneous seizure of prey by the upper and lower jaws. This sort of
adaptation is common in a large variety of vertebrate groups (Thom-
son, 1969 and references therein).

Evidence for a Gnathorhiza-ltpidosirQuid ancestry based on aesti-
vation seems equally as poor as that based on dentition. I (Berman,
1968:834) have argued that “The over 200 million year span between
the Lower Permian aestivators and the earliest known true lepidosi-
renids seems more than adequate time for this feature to have arisen
a second time. There is also no evidence to assume that these earliest
known lepidosirenids, Oligocene of Africa (Stromer, 1910), could aes-
tivate or that aestivation did not arise independently in the two modern
genera.” As noted by Thomson (1969), only Protopterus is capable of
true aestivation, Lepidosiren does not undergo a full aestivation, and

1979

Berman-— New Mexican Lungfish

229

Neoceratodus does not aestivate at all. Interestingly, he has presented
physiological evidence that the absence of aestivation in Neoceratodus
and its redaction in Lepidosiren is secondary.

Finally, Thomson (1972) has provided an unique approach to the
problem of differentiating evolutionary lineages within the Dipnoi. By
demonstrating a direct relationship between cell size and DNA com
tent, he has reconstructed evolutionary changes in the cellular DNA
content of lungfish and has been able to discern several phyletic trends.
Of importance here is Thomson’s conclusion (1972:36) that the data
“seem to exclude the Permian genus Gnathorhiza from the ancestry
of the modern families.”

Acknowledgments

Thanks are extended to Dr. Mary Dawson, Carnegie Museum of Natural History, for
critically reading the manuscript. This research was funded in part by the M. Graham
Netting Research Fund through a grant from the Cordelia Scaife May Charitable Trust.

Literature Cited

Berman, D. S. 1968. Lungfish from the Lueders Formation (Lower Permian), Texas
and the GnathorhizaAepidosirQnid ancestry questioned. J. Paleontol., 42:827-835.

. 1976fl. Cranial morphology of the Lower Permian lungfish Gnathorhiza (Os-

teichthyes: Dipnoi). J. Paleontol., 50:1020-1033.

. 1976^. Occurrence of Gnathorhiza (Osteichthyes: Dipnoi) in aestivation bur-
rows in the Lower Permian of New Mexico with description of a new species. J.
Paleontol., 50:1034-1039.

Bridge, T. W. 1898. On the morphology of the skull in the Paraguayan Lepidosiren
and in other dipnoids. Trans. Zool. Soc. London, 14:325-376.

Carlson, D. J. 1968. The skull morphology and estivation burrows of the Permian
lungfish Gnathorhiza serrata. J. Geol., 76:641-663.

Cope, E. D. 1877. Descriptions of extinct Vertebrata from the Permian and Triassic
formations of the United States. Proc. Amer. Philos. Soc., 17:182-193.

— . 1883. Fourth contribution to the history of the Permian formation of Texas.

Proc. Amer. Philos. Soc., 20:628-636.

Denison, R. H. 1974, The structure and evolution of teeth in lungfishes. Fieldiana,

Geol., 33:31-58.

Lund, R. 1970. Fossil fishes from southwestern Pennsylvania, Part I: Fishes from the
Duquesne Limestones (Conemaugh, Pennsylvanian). Ann. Carnegie Mus., 41:231-
261.

1973. Fossil fishes from southwestern Pennsylvania, Part II: Monongahela

dunkardensis, new species, (Dipnoi, Lepidosirenidae) from the Dunkard Group.
Ann. Carnegie Mus., 44:71-101.

-. 1975. Vertebrate-fossil zonation and correlation of the Dunkard basin. Pp. 171-

178, in Proceedings of the first I. C. White Memorial Symposium: The age of the
Dunkard (J. A. Barlow, ed.). West Virginia Geol. Econ, Surv., 352 pp.

— . 1976. General geology and vertebrate biostratigraphy of the Dunkard basin.

Pp. 225-239, in The Continental Permian in central, west, and south Europe (H,
Falke, ed.), D. Reidel Publ. Co., Dordrecht, Holland, 352 pp.

Olson, E. C. 1951. Fauna of upper Vale and Choza: 1-5. Fieldiana, Geol. 10:89-128.
1975. Vertebrates and biostratigraphic position of the Dunkard. Pp. 155-165,

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Annals of Carnegie Museum

VOL. 48

in Proceedings of the First 1. C. White Memorial Symposium: The age of the
Dunkard (J. A. Barlow, ed.), West Virginia Geol. Econ. Surv., 352 pp.

Olson, E. C., and E. Daly. 1972. Notes on Gnathorhiza (Osteichthyes, Dipnoi). J.
PaleontoL, 46:371-376.

Olson, E. C., and P. P. Vaughn. 1970. The changes of terrestrial vertebrates and
climates during the Permian of North America. Forma et Functio, 3:113-138.

Owen, R. 1839. Description of the Lepidosiren annectens. Trans. Linn. Soc. London,
18:327-361.

Romer, a. S., and E. C. Olson. 1954. Aestivation in a Permian lungfish, Breviora.
30:1-8.

Romer, A. S., and H. J. Smith. 1934. American Carboniferous dipnoans. J. Geol.,
42:700-719.

ScHULTZE, H.-P. 1975. Die Lungenfisch-Gattung Conchopoma (Pisces, Dipnoi).
Senckenb. Leth., 56:191-231.

. 1977. Megapleuron zangerli a new dipnoan from the Pennsylvanian, Illinois.

Fieldiana, Geol., 33:375-396.

Stromer, E. 1910. Ueber das Gebiss der Lepidosirenidae und die Verbreitung tertiarer
und mesozoischer Lungenfische. R. Hertwig Festschrift, Munchen, 2:613-624.

Thomson, K. S. 1969. The biology of the lobe-finned fishes. Biol. Rev. 44:91-154.

— . 1972. An attempt to reconstruct evolutionary changes in the cellular DNA

content of lungfish. J. Exp. Zool., 180:363-372.

Thomson, K. S., and K. S. W. Campbell. 1971. The structure and relationships of
the primitive Devonian Xungfish—Dipnorhynchus sussmilchi (Etheridge). Bull. Pea-
body Mus. Nat. Hist., Yale University, 38:1-109.

Vaughn, P. P. 1964. Evidence of aestivating lungfish from the Sangre de Cristo For-
mation, Lower Permian of northern New Mexico. Contrib. Sci., Los Angeles Co.
Mus. Nat. Hist., 80:1-8.

— . 1966. Comparison of the Early Permian vertebrate faunas of the Four Comers

region and north-central Texas. Contrib. Sci., Los Angeles Co. Mus. Nat. Hist.,
105:1-13.

— . 1969. Early Permian vertebrates from southern New Mexico and their paleo-

zoogeographic significance. Contrib. Sci., Los Angeles Co. Mus. Nat Hist., 166:1-

22.

— . 1970. Lower Permian vertebrates of the Four Corners and the Midcontinent

as indices of climatic differences. Proc. 1969 North Amer. Paleont. Conv., Chicago,
d:388-408.

Westoll, T. S. 1949. On the evolution of the Dipnoi. Pp. 121-148, in Genetics, pa-
leontology and evolution (G. L. Jepsen, E. Mayr, and G. G. Simpson, eds.), Prince-
ton Univ. Press, 474 pp.

5'(? 7. 73
fHKlHSL

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ANNALS

of CARNEGIE MUSEUM

CARNEGIE MUSEUM OF NATURAL HISTORY
4400 FORBES AVENUE ® PITTSBURGH, PENNSYLVANIA 15213
VOLUME 48 7 SEPTEMBER 1979 ARTICLE 14

THE YELLOW-BELLIED SEA SNAKE, PELAMIS PLATURUS
(REPTILIA: HYDROPHIIDAE), IN THE
PHILIPPINES

C. J. McCoy

Curator, Section of Amphibians and Reptiles

Donald E. Hahn^

Abstract

A collection of hydrophiid snakes from Marinduque Island, Philippines, includes the
only sizable sample (73 specimens) of Peiamis platurus reported from the archipelago.

Introduction

Although a dozen species of sea snakes have been found in Philip-
pine waters information on distribution, ecology, and relative abun-
dance of these snakes remains fragmentary (Dunson and Minton,
1978). This is especially true for the yellow-bellied sea snake, Peiamis
platurus (Linnaeus). Conflicting reports on the occurrence and abun-
dance of Peiamis in the Philippines lead us to record observations on
a collection of hydrophiids from Marinduque Island.

Observations

Marinduque Island lies in the northern Philippines, in the center of
Tayabas Bay which is enclosed by the southern tip of Luzon, Mindoro
to the west, and Tablas and Sibuyan to the south. The collection,
including five species of hydrophiids, Acrochordus granulatus and

^ Marcus J. Lawrence Memorial Hospital, Cottonwood, Arizona 86326.
Submitted for publication 13 February 1979.

231

232

Annals of Carnegie Museum

VOL. 48

Fig. 1. — Map of Marinduque Island, showing the hydrophiid collecting area (arrow),
and the 50- and 100-foot isobaths.

Cerberus r. rynchops, and a quantity of banded eels resembling Hy-
drophis, was obtained by purchase from native fishermen at various
times during 1971 and 1972. Most of the hydrophiids were taken in a
large, shallow bay on the north coast of Marinduque that is partially
enclosed by Banet Island (Fig. 1). The area is known as “Botilao,”
from the name of a nearby village.

The remarkable feature of the hydrophiid sample is the large number
of Pelamis platurus (73 specimens; CM 66892-66961, 68009-68011),
which outnumber the other four species. There are nine males 385 mm
to 630 mm total length, and 64 females ranging from 343 mm to 693
mm total length. The size class distribution is strongly bimodal (Fig.
2). Forty-eight specimens (46 females, two males) form a cohesive
class between 343 mm and 437 mm. The remaining 25 (18 females,
seven males) range from 476 mm to 693 mm. According to the de-

1979 McCoy and Hahn — Philippine Pelamis 233

TOTAL LENGTH (mm.)

Fig. 2.---Size class distribution of the Marinduque Pelamis, in 10 mm increments. Solid
bars represent females, hollow bars males. The smallest vertical unit represents one

individual.

mography reported for Eastern Pacific Pelamis (Kropach, 1975), those
in the smaller group are subadults and specimens over 500 mm are
adults. The presence of a well-defined subadult size class confirms
strongly seasonal breeding in Western Pacific Pelamis populations, as
suggested by Cogger (1975:129).

The other hydrophiids in the collection are 26 Laticauda colubrina
(CM 66964-66977, 68803-68814), four Hydrophis cyanocinctus (CM
68006, 68744-68746), 43 Hydrophis fasciatus atriceps (CM 68007,
68747-68788), and 12 Hydrophis inornatus (CM 68008, 68789-68799),

Discussion

Published records of Pelamis in the Philippines are numerous and
scattered, but portray Pelamis as one of the least abundant hydro-
phiids in the area. In his monograph of Philippine snakes Taylor (1922)
reported a single specimen of Pelamis, and noted that the species is
“rare in the Philippines.” Malcolm Smith (1926) quoted Belcher’s ob-
servation of “thousands” of Pelamis in the Mindoro and Sulu seas,
but saw no Philippine specimens. Herre (1942) noted that Pelamis is
“not taken as often as some (other) species” in the Philippines. Gen-
eral reviews (Tu and Tu, 1970; Cogger, 1975) usually include the Phil-
ippines in the range of Pelamis, but specific locality records are scarce.
Perhaps this is why Hecht et al. (1974) stated that Pelamis is “also
recorded from the Philippines . . . from the Sulu Sea and not from the
main island area.” Finally, Dunson and Minton (1978) reported only
three Pelamis among a sample of 421 sea snakes from shoals near the
Gigante Islands.

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Annals of Carnegie Museum

VOL. 48

Our observations from Marinduque Island demonstrate that Pelamis
is not necessarily a rare snake in nearshore habitats in the Philippines,
but may be locally or seasonally abundant. The reason for the relative
abundance of the Marinduque Pelamis, however, remains a mystery.
The snakes were taken at various times of the year (January-February
1972; April 1971; May 1972) but comparable collections are not avail-
able for other months, so seasonality cannot be confirmed. The ab-
sence of newborn snakes or gravid females in the sample argue against
a reproductive congregation.

Dunson and Ehlert (1971) described concentrations of Pelamis in
surface slicks formed by horizontal convergence of waterflow and
winds. Such slicks frequently form in the zone of transition from shal-
low, turbid nearshore water to deep, offshore water. Marinduque Is-
land has a narrow shallow zone, and is surrounded by deep waters
(Fig. 1), providing potentially ideal conditions for the formation of
slicks and concentration of Pelamis.

Acknowledgments

We thank Dr. Harold Voris for assistance in identifying the Hydrophis species, Dr,
William A. Dunson for access to a then-unpublished manuscript on Philippine sea
snakes, and Ms. Laura Arney for technical assistance. Specimens are deposited in
Carnegie Museum of Natural History (CM),

Literature Cited

Cogger, H. 1975. Sea snakes of Australia and New Guinea. Pp. 60-139 in The biology
of sea snakes (W. A. Dunson, ed.). University Park Press, Baltimore, Maryland,
X + 530 pp.

Dunson, W. A., and G. W. Ehlert. 1971. Effects of temperature, salinity, and
surface water flow on distribution of the sea snake Pelamis. Limnol. Oceanogr.
16:845-853.

Dunson, W. A., and S. A. Minton. 1978. Diversity, distribution, and ecology of
Philippine marine snakes (Reptilia, Serpentes). J. Herpetol. 12:281-286.

Hecht, M. K., C. Kropach, and B. M. Hecht. 1974. Distribution of the yellow-
bellied sea snake, Pelamis platurus, and its significance in relation to the fossil
record. Herpetologica 30:387-396.

Herre, a. W. C. T. 1942. Notes on Philippine sea-snakes. Copeia, 1942:7-9.
Kropach, C. 1975. The yellow-bellied sea snake, Pelamis, in the Eastern Pacific. Pp.
185-211, in The biology of sea snakes (W. A. Dunson, ed.). University Park Press,
Baltimore, Maryland, x + 530 pp.

Smith, M. 1926. Monograph of the sea-snakes (Hydrophiidae). British Museum (Nat-
ural History), London, xvii + 130 pp.

Taylor, E. H. 1922, The snakes of the Philippine Islands. Philippine Bureau of Science
Publ., 16:1-312.

Tu, A. T., and T. Tu. 1970. Sea snakes from Southeast Asia and Far East and their
venoms. Pp. 885-903, in Poisonous and venomous marine animals of the world (B.
W. Halstead, ed.), U.S. Government Printing Office, Washington, 3:xxv + 1-1006.

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VOLUME 48 7 SEPTEMBER 1979 ARTICLE 15

TAPHONOMY OF MICROVERTEBRATE FOSSIL
ASSEMBLAGES

William W. Korth

Rea Fellow, Section of Vertebrate Fossils

Abstract

Two principal modes of origin have been suggested for microvertebrate deposits — 1)
the scatological hypothesis, and 2) the fluvial hypothesis. These two hypotheses were
tested by 1) examining remains found in modern carnivore feces and owl pellets, 2)
experimentally determining the susceptibility of various skeletal elements of small mam-
mals to stream sorting, and 3) analyzing microvertebrate fossil assemblages.

The abraded rather than sharply broken condition of the fossil bone and the frequency
of preservation of individual elements most closely resembled that of the bone that had
been experimentally abraded rather than bone from predator scats.

Although the hydraulic properties of skeletal elements are dependent to some extent
on shape, empirically derived settling curves show that the settling velocity of mammal
bones is strongly size dependent. As predicted from the curves, it was found that the
sediment particle size of the fossil bearing deposits had equivalent hydraulic properties
to the bone contained in the sediment. It was concluded that the accumulations inves-
tigated were of alluvial origin.

The paleoecologic applications of alluvially sorted microvertebrate fossil accumula-
tions are limited because the effects of stream flow can markedly alter the composition
and relative representation of members of the fauna.

Introduction

Numerous microvertebrate fossil assemblages have been described
from the Tertiary of North America, but few studies have dealt with
the taphonomy of these accumulations.

Taphonomy, as defined by Efremov (1940), is the study of events
between the death of an organism and its subsequent deposition and

Submitted for publication 1 February 1979.

235

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fossilization. Studies of taphonomy are concerned with the processes
which ultimately are reflected in the condition, orientation, and selec-
tive preservation of fossil materials. Such studies are necessary before
accurate paleoecological reconstructions can be made from the avail-
able fossil assemblage.

Until recently, the taphonomy of fossil sites has been largely ignored
by vertebrate paleontologists. Taphonomic studies of vertebrate fossil
localities have mostly been restricted to fish (McGrew, 1975) and larger
animals such as large Paleozoic reptiles (Olson, 1962), dinosaurs (Dod-
son, 1971; Gradzinski, 1970; Langston, 1976; Lawton, 1977), and larger
mammals (Clark et al., 1967; Voorhies, 1969; Shotwell, 1955, 1958,
1963; Behrensmeyer, 1975). Few studies dealing specifically with an-
imals of rabbit size or smaller have been made. Only Wolff (1973) and
Estes and Berberian (1970) have undertaken taphonomic studies of
microvertebrate assemblages. Estes and Berberian (1970) applied
Shotwell’s techniques and agreed with the latter that preferential pres-
ervation was partly dependent on distance of transport. Wolff (1973)
explained the disproportions of representation of particular taxa as
sorting bias, but offered no experimental evidence,

Dodson (1973) and Wolff (1973) have proposed that hydraulic sorting
may be a major factor contributing to the selective preservation of
microvertebrate fossils. Dodson (1973) concluded that such assem-
blages are therefore poor indicators of paleocommunities and are more
useful as sources of information about paleocurrent velocities,

Mellett (1974:349) suggested a quite different theory for the origin
of microvertebrate fossil accumulations: “I propose that most or all
microvertebrate fossil accumulations first passed into or through the
digestive tracts of carnivores (mainly mammalian, but including pre-
daceous fish, reptiles and birds) and were deposited as fecal droppings
(scat) in or near a stream, lake, or other basin, where they were sub-
sequently covered by sediment.” He offered the term coprocoenosis
for such accumulations, and suggested that fracture and abrasion of
bone, which had been previously explained as results of stream trans-
port, are due to effects of the ingestive and digestive processes of
predators.

Experimental work, although essential to the verification of taphon-
omic hypotheses, has seldom been undertaken. Experiments on the
hydraulic behavior of bone have been conducted (Voorhies, 1969; Beh-
rensmeyer, 1975) but have been restricted to larger mammals, the
smallest of which was a coyote. Dodson (1973) conducted flume ex-
periments with the skeleton of a mouse.

The durability of bone on a land surface and the time necessary for
decomposition or destruction of soft tissues and bone tissue itself have
been discussed by Clark et al. (1967), Voorhies (1969), Clark and

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Korth — Microvertebrate Taphonomy

237

Guensburg (1970), Behrensmeyer (1975), and Payne (1965). Only Voor-
hies (1969), Clark and Guensburg (1970), and Payne (1965) reported
experimental results on the time necessary for scavenger and insect
activity to strip the flesh from a carcass. Voorhies (1969) reported on
carcasses of larger animals, the smallest of which were rabbits and the
largest of which were calves, whereas Payne (1965) was concerned
with the carcass of a baby pig. No study has previously been made of
such activities on animals covering the size range reported for mi-
crovertebrate fossil faunas. Dodson (1973) studied the decomposition
of mouse, toad, and frog bodies submerged in pond water. This test
was conducted in quite a different situation from that used by the
former two authors and is not directly comparable. An understanding
of decomposition is essential in the interpretation of fossil assemblages
because it will allow a worker to estimate biases of preservation cre-
ated by such activity.

The effect of stream transport on the condition of bone has not been
tested, although some authors have speculated that it may explain the
scarcity of some elements of the skeleton at certain fossil localities.

The purpose of this investigation was to devise tests of existing
theories of accumulation and preservation of microvertebrate fossils
and to apply these tests to actual fossil localities.

Methods and Materials

Experiments and observations were made on skeletons and fecal material of Recent
animals from the following sources: 1) Bam Owl pellets — Keith County, Nebraska; 2)
Great Horned Owl pellets— Garden County, Nebraska; 3) coyote feces— Brown and
Keya Paha counties, Nebraska; 4) skeletons of Recent small mammals— Brown, Ante-
lope, and Lancaster counties, Nebraska.

The settling rates of Recent bones were determined as follows: 1) bones of small
mammals (shrew to rabbit size) were dropped in a settling tube 10 cm in diameter,
allowed to settle for 50 cm, and then timed with an electric timer during settling for
another 50 cm; 2) bones of intermediate size (raccoon) were similarly tested in a settling
tube 25 cm in diameter over a distance of 60 cm; 3) large bones (horse and sheep) were
settled in a swimming pool and timed with a stopwatch over a distance of 3 m. Results
were recorded in tenths of seconds. Bones were released beneath the surface of the
water and allowed to reach a preferred settling orientation before timing began. Only
naturally cleaned (by insects, owls, bacteria) bones were used in the settling experiments
because it was found that bones that were boiled in preparation settled at a slower rate
than fresh ones, probably because of decreased density.

Bones of small mammals were removed from dry owl pellets with the use of forceps.

The abrasion experiments were conducted using two rubber-lined tumbling barrels 12
cm in diameter. The barrels were half filled with quartz grains 2 to 4 mm in diameter
which served as an abrasive.

The fossils analyzed were recovered by the screen washing method described by
Hibbard (1949) and McKenna (1962). Approximately 1,000 kg of matrix were processed
from each locality. The fossils were identified by the writer and deposited in the collec-
tion of the University of Nebraska State Museum. Sediment samples from the fossil
quarries were analyzed in the sedimentology laboratory of the Geology Department of
the University of Nebraska.

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Table \.— Percentage of representation of skeletal elements of small mammals (all taxa
combined) preserved in predator feces or owl pellets, and fossil assemblages (percent-
age representation = observed number of elementslexpected number of elements in

complete skeletons).

Predators

Fossil assemblages

Element

Barn

Owl

Great

Horned

Owl

Coyote

Bw 110

Kx 110

Rodeo

Ranch*

(Pleisto-

cene)

Mandible (ramus)

92.5

86.7

97.6

7.9

61.8

15

Maxilla

87.5

90.0

78.6

6.9

29.7

_

Tibia

87.5

90.0

48.8

30.7

62.3

16

Humerus

82.5

93.3

69.0

15.3

43.9

11

Femur

85.0

93.3

70.2

24.3

80.2

10

Ulna

82.5

76.7

63.1

7.4

18.9

11

Radius

62.5

76.7

59.5

3.7

6.6

5

Pelvis

72.5

90.0

47.6

2.1

12.3

1

Calcaneum-astragalus

26.2

31.7

19.0

45.2

22.9

11

Incisors

86.2

86.7

100.0

48.2

68.6

48

Cheek teeth

59.1

71.0

78.9

14.4

19.6

39

Scapula

62.5

63.3

45.2

0.0

4.2

5

Mean value

73.9

78.6

64.8

17.2

35.9

12

Minimum number of
individuals present
in sample

20

15

42

99

106

137

* Taken from Wolff (1973: 94, Fig. 2).

CoPROcoENOSis Hypothesis

Mellett (1974) compared bone extracted from feces of Recent mam-
malian carnivores with fossil bone fragments from an Eocene deposit.
He noted a striking similarity between the bones from these two
sources and concluded that the Tertiary fossils were deposited as car-
nivore fecal material. Several authors have suggested that fossil re-
mains from specific Tertiary and Pleistocene localities were accumu-
lations of carnivore feces (McGrew, 1963) or regurgitated pellets of
owls (Brodkorb, 1959; Etheridge, 1965; James, 1963; Dawson, 1958)
or diurnal avian predators (May hew, 1977). Examples of fossil material
actually preserved in recognizable coprolites (Mellett, 1974) or owl
pellets (Gawne, 1975; Storch, 1969) are rare.

The following description of Recent fecal material and pellets of
carnivorous vertebrates establishes the parameters by which the co-
procoenosis hypothesis of fossil vertebrate accumulations may be
tested. Undigested remains of prey from two classes of predators,
birds and mammals, will be treated. Fecal material of fishes, amphib-
ians, and reptiles were not surveyed but this is not judged to be a

1979

Korth—Microvertebrate Taphonomy

239

Fig. 1.— A) Dorsal view of rodent skulls experimentally abraded -in a tumbling barrel
(left), and extracted from coyote feces (center) and a Great Horned Owl pellet. Skulls
from the owl pellet and tumbling experiment show sutural separation of cranial bones
while the skull from the coyote scat exhibits breakage of cranial bones. Scale x 1.6. B)
Ventral view of the same.

serious omission. Among the latter, only snakes are quantitatively im-
portant as predators on terrestrial vertebrates. Snakes digest all bone
of their prey and only keratinaceous substances such as hair, horns,
feathers, and epidermal scales of reptiles are unaffected by digestion
(Bellairs, 1970).

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Survey of Modern Predators
Owls

Condition of —Material from a total of 40 to 60 pellets of two
species of owls, Bubo virginianus (Great Horned Owl) and Tyto alba
(Barn Owl), was collected and examined in order to observe the con-
dition of individual bones. A census (Table 1) of the relative numbers
of skeletal elements was from a sufficient number of pellets to repre-
sent a minimum number of 15 (Great Horned Owl) and 20 (Barn Owl)
individuals.

In general, the condition of the bone was as described by previous
authors (Montgomery, 1899; Reed and Reed, 1928; Moon, 1940; Cza-
plewski, 1976; Mayhew, 1977), There was no evidence of acid erosion
of the bone from the pellets of either owl species (Figs. 1, 2, 3, 4, and
5). In a few instances, the ends of long bones of very young individuals
showed some discolo ation possibly caused by stomach acids, al-
though there was no evidence of corrosion. There were no broken
bone fragments in the pellets of the Barn Owl. Even very delicate
bones (scapulae, ribs, cranial bones) were not broken.

Great Horned Owl pellets showed some breakage in the bones of
larger prey {Sylvilagus , Dipodomys) but the bones of all smaller prey
were unbroken. Frequently jaws were found articulated with skulls,
and several articulated vertebrae were found in pellets. Once removed
from the matrix of the pellet, these elements did not remain in asso-
ciation with one another because the soft connective tissue had been
digested by the owl.

Jaws and skulls generally contained all teeth within them, although
the teeth were loose in their sockets and readily fell out upon release
from the pellets. This was true for cheek teeth as well as for incisors
of rodents.

Contrary to previous authors (Montgomery, 1899; Moon, 1940; Mel-
lett, 1974; Mayhew, 1977), no characteristic crushing was found on the
base of any skull. Only in specimens of Sylvilagus was any breakage
of skull material noted. However, it was common to find the occiput
and tympanic bullae separated from the remainder of the skull but near
their proper position. The occiput and bullae were separated along the
sutures of the skull and showed no breakage, suggesting that they were
not broken during capture or ingestion by the owl but became sepa-
rated in the stomach of the owl after the soft connective tissue which
bound them to the rest of the skull had been digested (see Fig. 1).
Skulls of very young individuals were generally separated into all of
the separate bones along the cranial sutures with no sign of breakage
on even the most delicate cranial bones.

Preferred prey.— The animals represented in the owl pellets were

1979

Korth— Microvertebrate Taphonomy

241

Fig. 2. — A) Lateral view of mandibles experimentally abraded in a tumbling barrel (left),
and extracted from coyote feces (center) and a Great Horned Owl pellet (right). Mandible
from the owl pellet shows no effect of digestion, that from the coyote scat shows irreg-
ular breaks, whereas the abraded mandible has characteristic wearing down of the angle,
coronoid, and condyle, as well as bone thinning and formation of holes at the base of
the sockets of the molars, at the pulp cavity for the incisor, and on the ventral surface
of the jaw around the incisor. Scale x2.5. B) Medial view of the same.

predominantly nocturnal or crepuscular mammals (generally rodents)
and some birds. Several authors have noted that nearly all animals
represented in owl pellets were nocturnal mammals. Diurnal forms and
lower vertebrates were rare or absent. The occurrence of the mammals

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Fig. 3.— -A) Tibia-fibula experimentally abraded in a tumbling barrel (left), and extracted
from coyote feces (center) and a Great Homed Owl pellet (right). Tumbled bone shows
abrasion on the ends and loss of the fibula. That from the coyote feces exhibits angular
breakage of the fibula and on the proximal end. The bone from the owl pellet is unaltered.
Scale x2.3. B) Humeri experimentally abraded in a tumbling barrel (left), and extracted
from coyote feces (center) and a Great Horned Owl pellet (right). Tumbled bone shows
abrasion of ends and processes, bone from coyote scat shows angular breakage, and
from the owl pellet shows no effect. Scale x2.4.

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Korth— Microvertebrate Taphonomy

243

Fig. 4.— A) Femora experimentally abraded in a tumbling barrel (left), and extracted
from coyote feces (center) and a Great Horned Owl pellet (right). Condition of bones
as described for humeri (Fig. 3B). Scale x2.4. B) Scapulae experimentally abraded in
a tumbling barrel (left), and extracted from coyote feces (center) and a Great Horned
Owl pellet (right). Condition of bones from predator scats as described for other limb
bones (Figs. 3 and 4A). Tumbled bone shows rounded medial edge and loss of coracoid
and acromion processes, as well as abrasion of proximal end. Scale x2.4.

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Fig. 5.— A) Isolated frontal bones of rodents extracted from a Great Horned Owl pellet
(left) and experimentally abraded in a tumbling barrel (right). The bone from the owl
pellet is unbroken and unaffected by digestion. The tumbled bone has rounded edges
and thinning of the central area of the bone resulting in breakage in that area (see Fig.
2 for similar breakage on the mandible). Scale x2.4. B) Pelves of small mammals ex-
perimentally abraded in a tumbling barrel (right), and extracted from coyote feces (cen-
ter), and a Great Horned Owl pellet (left). Condition of different bones is the same as
discussed for limb bones (see Figs. 3, 4). Scale x2.4.

1979

Korth— Microvertebrate Taphonomy

245

Fig. 6.-— Profile curves of percentage representation (number present/number expected)
of small mammal bones found in predator feces or pellets (data from Table 1). Horizontal
axis represents individual elements: M = mandible, T = tibia, X = maxilla, I = incisor,
F = femur, H = humerus, U = ulna, P = pelvis (innominate), R = radius, S = scap-
ula, CT = cheek teeth, C-A = calcaneum and astragalus.

in the pellets generally reflected the relative abundance of these forms
in the community (Evans and Emlen, 1947; Anderson and Long, 1961;
Epperson, 1976; Jones, 1952; Glue, 1970).

Glue (1970) established that predator preference is exhibited by the
fauna represented in owl pellets due to: 1) range of the predator versus
the range of the prey, 2) activity rhythm of the bird and the prey, 3)
method of capture, 4) size, and 5) color and mode of locomotion. All
of these biases were reflected in the animals represented in the pellets
of the two owls tested. The Barn Owl pellets yielded smaller mammals
and more meadowland forms, predominantly Peromyscus and Micro-
tus. The Great Horned Owl pellets contained larger mammals (lepor-
ids, geomyids) and more arid climate forms, Perognathus and Dipod-
omys.

Percentage representation. —An aspect of fecal pellet assemblage
not considered by Mellett (1974) in his description of the coprocoenosis
hypothesis is the percentage of representation of elements present in
the assemblage. Eor example, with a minimum number of individuals
of 10, there should be 20 femora, 20 humeri, and so on, in order to

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have a 100% representation. From census taken of individuals repre-
sented in Barn Owl and Great Horned Owl pellets (Table 1), it is
evident that all elements surveyed show a very high percentage rep-
resentation in both species of owls. Elements such as vertebrae and
phalanges were quite abundant in the owl pellets, however, these
bones were not counted. The very low value for the calcanea and
astragali may result from the method used in processing the pellets.
These small elements, along with smaller cheek teeth, may have been
overlooked. Alternatively, the lack of these tarsal elements could be
caused by the predators’ failing to ingest the feet of the prey which
would have little nutritive value.

The mean value for percentage of representation of the elements
surveyed of prey in the Barn Owl pellets is 73.9%. Great Horned Owl
pellets show an even higher mean representation of elements (78.9%).
The curve generated for individual percentages of representation of
selected elements for both owls is nearly identical (Fig. 6). They both
show extremely high representation for all elements. The lowest per-
centages are for the calcanea and astragali, below one half the value
of the mean.

Hawks

Condition of Diurnal avian predators regurgitate pellets of

undigested materials as do owls. However, the ingested bone is subject
to decomposition through the action of stomach acids. Bone ingested
by diurnal birds of prey (kestrels, buzzards, hawks) is frequently to-
tally digested, and partial digestion results in a characteristic etching
of bones and teeth of the prey (Moon, 1940; Mayhew, 1977). Other
diurnal predatory birds, such as herons, totally digest all bone and
leave no identifiable bone elements in either pellets or feces (Glue,
1970). Bones that are partially digested by diurnal predatory birds are
clearly distinguishable from those of owls (Moon, 1940; Glue, 1970;
Mayhew, 1977).

Long bones recovered from pellets of diurnal avian predators show
erosion of the epiphyses and ends of the shafts, which results in sharp
pointed ends that show a thinning of the bones in these areas (Mayhew,
1977) (molars and incisors) are reduced in size and exhibit etch-

ing on the entire exposed surface. Enamel is often reduced to a fine
powder on the surface of the tooth. Teeth that remain in the skull or
jaw through the digestive process have eroded crowns but show no
effect of digestion on the roots. Skulls of prey may be discarded before
ingestion (Mayhew, 1977).

Preferred The biases on prey selection of diurnal carnivorous

birds are similar to those outlined for nocturnal birds (Glue, 1970).

Percentage representation.— Ho census was taken of diurnal avian

1979

Korth-=— Microvertebrate Taphonomy

247

Fig. 7.— -Fragments of bone extracted from coyote feces (above) and fossil bone frag-
ments from locality Kx 110 (below). Bone ingested by the coyote has angular breaks
along the edges, the fossil bone shows marked rounding of the edges. The fossil bone
fragment on the left shows chipping and flaking of outer layers, attributed to drying
subaerially. Scale xl.6.

predator pellets. The representation would probably be unpredictable
and inconsistent in many ways because of the differential digestion of
bony elements in the stomachs of these birds (Moon, 1940; Mayhew,
1977).

Mammalian Predators

Condition of Remains from over 50 individual coyote scats

collected in Keya Paha and Brown counties of northcentral Nebraska
were examined by the author. Much of the bone contained in this
material consisted of sharply broken bone fragments as described by

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VOL. 48

Mellett (1974). Smaller mammals that were present in the carnivore
feces were represented by whole limb elements and jaws. In a few
instances an articulated pes or manus of a rodent was observed. Thin
walled bones (skulls and scapulae) were always broken. All ulnae,
astragali, and calcanea were unbroken. Other major limb bones were
generally whole (62% of the recognizable humeri and femora recovered
were unbroken, and 86% of the tibiae were whole). Jaws were gen-
erally broken except for those of very small rodents. Unlike remains
from owl pellets, jaws in mammal scats rarely contained the original
number of teeth (see Figs. 1, 2, 3, 4, 5, 7).

Preferred The animals represented in the coyote feces stud-

ied provided a good qualitative indication of the mammals present in
the area (Jones, 1964). No attempt was made to compare relative abun-
dance of the mammals present in the feces to the actual fauna of the
area, however, because no quantitative data on the latter are available.
Several authors have shown that animal remains in mammalian car-
nivore feces reflect the general abundance of the prey in the commu-
nity (Fichter et al., 1955; Korschgen, 1957), although some biases have
been noted for some carnivores (Pearson, 1964).

Factors affecting prey selection outlined by Glue (1970) for avian
predators are also applicable to mammalian predators (Pearson, 1964;
Mellett, 1974).

Percentage representation.— Tht representation curve for the coy-
ote scat material (Fig. 6) shows some variation from the owl pellet
curves. The lower representation of some long bones, notably the tibia,
in the carnivore curve is most likely due to bone breakage during
mastication, resulting in a series of small unidentifiable fragments.
Generally, the mammalian carnivore curve is quite similar to that of
the avian predators including the marked low percentage of tarsal ele-
ments. The mean value for the percentage of representation of ele-
ments for the coyote fecal material is 64.8%. This value is the lowest
of the predators tested.

Recognition of Fecal Accumulations in the Fossil Record

A scatological microvertebrate fossil accumulation should be rec-
ognizable by the following criteria: 1) a high percentage representation
of bones of the animals present (greater than 60%), 2) the majority of
limb bones are whole, 3) skulls common and frequently whole, 4) no
evidence of mechanical abrasion (stream abrasion).

Skull material in a coprocoenosis should be either whole or disar-
ticulated along cranial sutures as in owl pellets, crushed with angular
breaks as in carnivore feces, or etched by acid as in diurnal bird pellets.
Most or all limb elements of smaller animals should be whole. Bone

1979

Korth—Microvertebrate Taphonomy

249

of larger animals should be all fragmented and occasionally have tooth
marks from mastication by mammalian carnivores.

Mellett (1974) also suggested that such assemblages would be found
in depositional environments other than stream deposits. Hence, a
sedimentological study of the fossil deposit may also indicate the
source of origin for microvertebrate accumulations.

The paleoecological value of a coprocoenosis is questioned by Mel-
lett (1974) based on evidence presented by Pearson (1964) for predator
preference in mammalian carnivores. However, Pearson notes that the
least preferred prey of the mammalian carnivores are controlled by
other factors, such as avian predators. Any coprocoenosis, which has
input from both avian and mammalian carnivores, should give a rela-
tively close approximation to the living population of mammals. From
the descriptions of fecal and pellet material above, it should be possible
to determine if a single type of predator or a combination of predators
has contributed to an assemblage.

Fluvial Hypothesis

Several authors have described the characteristics of Tertiary and
Pleistocene microvertebrate assemblages from fluvial deposits (Wil-
son, 1960; Wolff, 1973). Mellett (1974) has argued, however, that many
assignments of fluvial origin to fossil assemblages are not based on
sedimentological evidence and may be incorrect. Some aspects of the
fluvial hypothesis, such as selective sorting (Wolff, 1973), orientation,
and necessary stream velocities required for transport (Dodson, 1973)
have been discussed for elements of small mammal skeletons. Other
aspects of a fluvial hypothesis for microvertebrate fossil accumulation,
however, have not been studied. A series of experiments on stream
abrasion and hydraulic equivalence of bone and on rates of decom-
position of fleshy tissues of small mammals have been conducted in
this study in order to establish a more complete model for fluvial de-
position of micromammalian bones.

The sedimentological interpretation of the fossiliferous deposit is
essential to the fluvial hypothesis of fossil accumulation. The environ-
ment of deposition of sediment can be determined on criteria of tex-
ture, fabric, degree of sorting, size of sedimentary particles, and the
presence of sedimentary structures (Allen, 1965, 1970). Fossil material
transported and deposited along with alluvially sorted sediment should
show effects of sorting. The fossils present should be hydraulically
equivalent to the sediment of the deposit. Behrensmeyer (1975) deter-
mined hydraulic equivalents of bones and teeth of large animals by
correlating settling velocities of these elements with the settling veloc-
ities of quartz grains of various sizes. No such experiments have been

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Table 2.-— Experimentally determined settling velocities (in cmisec) of skeletal elements
of seven genera of mammals, based on eight to 10 trials for all animals except Equus
and Ovis, for which four to five trials were run. (FM = area of foramen magnum in

cm^).

Genera

Element

Range

Mean

Standard

deviation

Sorex

skull

6.7-7. 1

6.9

0.1

FM = 0.05

mandible

6.5-7.5

7.0

0.3

femur

4.8-5. 2

5.0

0.1

astragalus

3.5-4.5

4.0

0.3

calcaneum

4.0-4.6

4.3

0.2

atlas

2.5-2.6

2.5

— ■

Peromyscus

skull

5.9-8.3

7.4

0.8

FM = 0.166

mandible

10.0-11.9

10.5

0.4

- pelvis

4. 1-5.8

5.1

0.5

tibia-fibula

10.9-14.3

13.1

1.0

tibia-fibula (p)

7.9-9.2

8.7

0.5

tibia-fibula (d)

7.7-7.8

7.7

—

femur

7.5-9.4

8.6

0.7

femur (p)

5.6~6.4

5.8

0.3

femur (d)

3.4~5.4

4.0

0.8

ulna

4.4-5. 5

5.0

0.6

radius

4.9-5.6

5.3

0.2

astragalus

6.7-7.0

6.8

0.1

calcaneum

6. 1-7.1

6.7

0.4

humerus

6.4-9. 1

8.2

0.8

humerus (p)

3. 8-8.9

5.6

1.7

humerus (d)

5. 2-8.3

6.4

1.0

atlas

4.3-5.4

4.9

0.3

lumbar vertebrae

4.8-6.0

5.4

0.4

metatarsal

4.5-5.2

4.8

0.2

phalanx

3.3-5. 1

4.0

0.6

scapula

3.0-4.2

3.5

0.5

rib

1. 9-2.7

2.2

0.3

upper incisor

10.0-11.6

10.7

0.5

lower incisor

9.1-10.0

9.5

0.3

maxilla -

8.9-10.0

9.3

0.4

maxilla+

6.7-7.0

6.8

0.1

molar (Mj)

9.4-10.4

10.0

0.4

molar (Mg)

10.0-10.2

10.1

0.1

molar (MT

8.9-11.4

9.5

0.5

Sciurus

skull

8.0-9.2

8.6

0.2

FM = 0.44

mandible

18.8-32.2

24.2

4.8

tibia

21.7-25.0

24.1

1.1

femur

15.2-20.8

18.6

2.0

astragalus

13.2-15.2

13.8

0.8

calcaneum

14.3-16.7

15.5

0.7

atlas

8.0-10.6

9.0

1.0

scapula

15.4-17.2

16.2

0.8

molar (M^)

16.7-17.8

17.5

0.4

1979

Korth — Microvertebrate Taphonomy

251

Table 2. — Continued.

Genera

Element

Range

Mean

Standard

deviation

Sylvilagus

skull

11.6-14.1

12.8

1.0

FM = 0.64

mandible

28.1-29.6

28.6

0.7

pelvis

8.8-9.2

9.1

—

femur

15.9-25.0

19.8

2.8

astragalus

14.5-16.4

15.3

0.8

atlas

10.6-11.4

11.1

0.2

scapula

15.4-20.8

19.0

2.0

molar (M2)

14.9-16.4

15.6

0.5

Procyon

skull

15.2-16.0

15.6

0.4

FM = 1.11

mandible

50.8-62.8

56.6

3.5

femur

26.6-32.3

29.0

2.3

astragalus

15.6-20.8

18.1

1.5

calcaneum

19.2-22.7

20.8

1.0

atlas

12.1-15.0

13.1

1.1

phalanx

16.2-19.8

18.0

1.1

rib

7.5-9.4

8.3

1.0

Ovis

skull

33.0-34.3

33.7

0.5

FM = 3.42

mandible

47.3-49.0

48.3

0.7

atlas

21.4-22.9

22.4

0.7

Equus

skull

47.3-66.9

53.4

6.8

FM = 9.08

mandible +

74.1-94.6

83.7

7.8

mandible -

88.5-91.4

90.5

1.4

femur

61.0-70.3

65.3

3.5

astragalus

61.0-68.6

64.1

2.8

atlas

28.3-30.5

29.3

0.9

(p) = proximal half of the bone; (d) = distal half of the bone; + (after maxilla of
Peromyscus) = major portion of the palate included on the specimen; - (after maxilla
of Peromyscus) = maxillary contribution to the palate is missing medial to the tooth
row in the specimen; + (after mandible of Equus) = symphysis was on the ramus; -
(after mandible of Equus) = ramus was separated behind the symphysis.

previously conducted on smaller animals. Experiments, discussed be-
low, were done to determine hydraulic equivalents of bone of micro-
mammals in order to establish an hypothesis for determining the size
of associated sediment in microvertebrate fossil deposits.

Hydraulic Equivalents of Bone

Experiments

Experiments employed in the determination of the settling velocities
of bones follow the procedure outlined by Behrensmeyer (1975:492).
Bones of seven animals were used— (shrew), Peromyscus
(mouse), Sciurus (squirrel), Sylvilagus (rabbit), Procyon (raccoon),
Ovis (sheep), and Equus (horse).

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VOL. 48

All elements of each form were from the same individual except
Equus for which bones of several individuals of equivalent size were
used. All tests were repeated eight to 10 times except for Ovis and
Equus for which, because of limited availability of the testing facility,
only four to five trials per element were made.

In addition to the whole-bone experiments, tests were run on prox-
imal and distal halves of the major limb bones and on maxillae of
Peromyscus. Test results for all forms are given in Table 2.

The results of the experiments on settling velocity allowed recog-
nition of five major groups of skeletal elements with different hydraulic
properties (called “Voorhies dispersal groups” by Behrensmeyer,
1975). Several elements remain in their assigned dispersal group
throughout the size range tested. In group I, the rib is the only element
consistently present. In group I /II, the atlas, pelvis, radius, and ulna
are consistently present although the latter two elements were tested
in Peromyscus only. Group II is characterized by the calcaneum, as-
tragalus, scapula, and the major limb bones except the tibia (= tibia-
fibula of some forms). Isolated molars of the tested animals belong to
this group as well, except in the case of the two smallest forms, Sorex
and Peromyscus. Only a few bones of a few taxa have settling veloc-
ities intermediate between those of groups II and III. No element of
all taxa is consistently found within group II /III. The jaw of nearly all
animals falls in group III. The tibia (= tibia-fibula) of the two rodents
tested fell in group III as well.

The most inconsistent results throughout the tested taxa are of the
skull. As can be seen in Table 3, the skull falls quite rapidly in Sorex
(group III) and moderately fast in Peromyscus, Ovis and Equus (group
II). In the intermediate-sized animals tested, Sciurus, Sylvilagus, and
Procyon, it falls as a member of group I /II, slower than in either the
extremely small or large animals.

The phalanges and metapodials are highly variable in relative size
and morphology depending on the weight and mode of locomotion of
the individual taxa. Such elements are often impossible to distinguish
taxonomically in microfossil assemblages; hence, no effort was made
to determine the settling velocities of these elements.

Comparison with Previous Work

Dodson (1973) and Voorhies (1969) determined the sequence of ele-
ment transport based on initial velocity required to transport the bone.
Dodson (1973) used a skeleton of Mus (mouse) in his flume experi-
ments. His results are quite similar to those of the generalized settling
sequence (Table 4). The only major differences are in the placement
of the radius and ulna. Dodson placed these bones in group II /III
(some of the last bones to be transported), whereas these elements are

1979

Korth— -Microvertebrate Taphonomy

253

Table 3 —Experimentally determined settling groups. Contents of each group based on
relative settling velocity, with group II representing the modal range of velocities for
test elements (data from Table 2).

Dispersal groups (see Voorhies, 1969; Behrensmeyer, 1975)

Genera

I(low)

I/II

II

ii/iii

iii(high)

Sorex

atlas

calcaneum

molar (Mg)

mandible

astragalus

femur

skull

Peromyscus

rib

metatarsal

maxilla +

molars

tibia-fibula

scapula

lumbar vert.

calcaneum

maxilla-

phalanx

atlas

astragalus

incisors

radius

skull

mandible

ulna

humerus

pelvis

femur

Sciurus

skull

astragalus

tibia

atlas

calcaneum
scapula
molar (M^)
femur

mandible

Sylvilagus

skull

astragalus

mandible

atlas

molar (Mg)

pelvis

scapula

femur

Procyon

rib

atlas

astragalus

femur

mandible

skull

molar (M*)
calcaneum
phalanx

Ovis

rib*

atlas

skull

mandible

scapula*

molar*

Equus

atlas

astragalus

femur

mandible

skull

molar*

* Data from Behrensmeyer, 1975:492.

Annals of Carnegie Museum

VOL. 48

254

O

Fig. 8.~Settling velocity of selected skeletal elements in a variety of terrestrial mam-
mals. Horizontal axis represents settling velocity of the element in cm/sec. Vertical axis
represents size of the individual taxa based on the area of the foramen magnum (A =
!4 X TT X w X h) in cm^. Data from Table 2. Both axes are logarithmic.

Fig. 8.— (Continued)

Table 4. — Comparison of dispersal groups determined by settling and flume experi-
ments.

I

i/ii

II

ii/iii

III

A. Generalized sequence based on settling velocity (this paper).

rib

atlas

calcaneum

molars

mandible

radius

astragalus

(small

tibia

ulna

humerus

mammals)

pelvis

scapula

femur

molars

maxilla

B. Transport sequence

of Mus (Dodson, 1973:19).

thoracic vert.

pelvis

skull

calcaneum

mandible

maxilla

vertebrae

tibia

radius

femur

humerus

ulna

C. Transport sequence

of Cams and

Ovis bones (Voorhies, 1969:69).

vertebrae

ulna

femur

mandible

skull

rib

scapula

tibia

(ramus)

mandible

sacrum

phalanges

humerus

(fused rami)

sternum

metapodia

pelvis

radius

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Annals of Carnegie Museum

VOL. 48

in group I /II in the settling model. Clearly, the differences here involve
the shape of the bones. The thin elongate bones are streamlined, and
it would probably be quite difficult for a current to initiate movement
in such bones. The gracility of these bones (low density and high sur-
face area) would not allow them to settle very quickly.

The closest representative in the present experiments to Dodson’s
specimen of Mus is Peromyscus (measurements given in Dodson,
1973:16, for the Mus specimen are within 10% of those for the Pero-
myscus used in the settling experiments). In both instances the skull
is in group II. In the flume experiments the occiput and nasal bones
were separated from the skull, and the skull used in the settling ex-
periments was intact. Boaz and Behrensmeyer (1976) have shown that
there is a marked difference in the hydraulic action of the skull ac-
cording to whether it is intact or broken, in their flume experiments
involving human bones. Thus, the skull shows no consistency in the
two experiments with mice. Differences between the flume and settling
experiments are few and seem related to the density of the bones. The
only difference in the results of the experiments occur in the bones
with a low density (skull, radius, and ulna).

The results of Voorhies’ (1969) flume experiments with skeletons of
Canis and Ovis are also remarkably similar to the settling sequences
observed in this study. Differences occur again in the placement of the
skull (group II in the flume and group I /II in the settling tests). The
tibia is intermediate in Voorhies’ results (group II), the settling results
place it in group III. However, it should be noted that the settling
experiments utilized rodents which have proportionately large tibiae.
No tests were made on the tibiae of the larger mammals, but it is
believed that this bone would most likely sort with the other major
limb bones.

A consistent sequence of settling velocity of bone elements exists
with minor variations for nearly the entire spectrum of mammalian
size (from Sorex to Equus). Behrensmeyer (1975) has shown that there
is a relationship between bone size and sediment size in deposits of
large mammalian fossils. The experiments described above allow ex-
tension of the predictive model to micromammalian assemblages.

Predicted Fluvial Assemblages

Graphs relating body size to settling velocity of bone elements pro-
duce linear relationships when plotted on logarithmic scales (Fig. 8).
The ordinate value for these graphs is equal to the area of the foramen
magnum (area = 14 x tt x width x height) of the individual tested.
This value is related to the body size (minus fat) of the individual
(Radinsky, 1967) and is appropriate in relating the sizes of the animals

1979

KoRTH — MiCROVERTEBRATE TaPHONOMY

257

Table 5. — Settling coefficients of isolated skeletal elements (Kfi of placental mammals

from shrew to horse size.

Element

(in sec^)

mandible

pelvis

tibia

femur

humerus

cheek teeth

astragalus

calcaneum

atlas

skull

rodent incisor*
distal tibia*
proximal femur*
distal humerus*
maxilla
ulna*

.0011

.0071

.0009

.0018

.0019

.0015

.0029

.0027

.0071

.0035

.0012

.0028

.0049

.0040

.0026

.0066

* Coefficient based on data for Peromyscus only, see Table 2.

in these experiments because it allows a quantitative assessment of
size independent of any specializations of the skeleton. It may be noted
that this measurement appears to be appropriate only for comparing
placental mammals. The writer conducted settling experiments on a
marsupial, Didelphis, of approximately equal size to the Procyon spec-
imen used in the tests. The opossum bones exhibited hydraulic behav-
ior very similar to that of the Procyon bones. The area of the foramen
magnum of Didelphis, however, is equal to that of Sciurus and thus
the Didelphis results would plot anomalously with the results of the
placental mammals. Therefore, in order to avoid complications and
definitions of size parameters, the results of the Didelphis tests are not
recorded in the tables.

Due to the regularity of the experimental results of settling velocities
of skeletal elements, equations can be constructed and used as a pre-
dictive tool in determining the hydraulic influences on the bones of
microvertebrates. The equation for determining equivalent particle
size of quartz grains to settling velocity of bone was determined by
Behrensmeyer (1975) to be dq = .000928 v^, where dq is equal to the
diameter of the equivalent quartz grain in centimeters, and v is the
settling velocity of the particular bone in centimeters per second (cm/
sec). The given coefficient (.000928) must have a dimension of seconds
squared per centimeter (sec^/cm) in order to be consistent with the
equation.

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VOL. 48

From the settling data, an equation for the settling velocity of a

particular bone can be expressed as v

/A

V K.

, if V is the predicted

settling velocity in cm/sec, Af is equal to the area of the foramen mag-
num of the individual in cm^, and Ke is the experimentally determined
coefficient of the settling velocity in sec^ for any given element in the
skeleton (see Table 5 for Kg values for the tested elements). Whole
elements were used in most of the determinations of Kg. Kg values for
partial elements are based solely on the Peromyscus results. Partial
elements give entirely different results from the whole elements.

Chi-square tests for consistency were run on all Kg determinations
of whole bones. Only with the atlas and skull (excluding the skull of
Sorex) did the value reach the 95% confidence level. The pelvis and
astragalus were at the 90% level, femur and calcaneum at the 85%
level, the mandible at the 70% level, cheek teeth at 50%, and the tibia
at only the 15% level of confidence.

If the value for settling velocity, based on the size of the animal and

particular element of the skeleton

is substituted for the

settling velocity in Behrensmeyer’s equation of quartz grain equiva-
lents (dq == . 000928 v^), an equation for the determination of the diam-
eter of quartz grains based on the size of an animal and a particular

element of that animal, can be derived

In any given microvertebrate assemblage, several different elements
of a single taxon are usually present. Utilization of the quartz grain
equivalent equation on all elements might, in such cases, yield several
different values for the expected diameter of the quartz associated with
the bones. The frequency of occurrence of each element should, there-
fore, be considered. By calculating dq only for the predominant ele-
ment or elements, a significant value should be attained. It is important
that determination of the abundance of elements be based on the per-
centage of representation of elements present and not on absolute num-
bers of elements.

The relative abundance of the taxa present in a fluvial accumulation
is also an important consideration. The value of dq for the most com-
mon elements of a taxon which makes up a large percentage of the
population (from calculated minimum number of individuals for all
taxa present in the accumulation) is more significant than a calculated
dq from elements of taxon only represented by a few individuals.

The calculation of dq for the most common skeletal elements of the
dominant taxa in a well-sorted assemblage should result in a narrow
range of values for dq. Once dq is determined, size analysis of the

1979

Korth— Microvertebrate Taphonomy

259

enclosing sediment should yield a value for the modal distribution for
the size of quartz particles present equal to or very near the predicted
dq value if the hypothesis is correct.

Condition of Bone Before Alteration by Alluvial Processes

The condition of bone deposited in alluvial sediment is dependent
on two main factors— 1) the effect of stream transport on the bone,
and 2) the condition of bone when introduced into the stream system.
The source of the bone is critical in dealing with the question of its
initial condition. Death of small mammals can be caused by disease,
parasites, unusual climatic conditions (such as prolonged drought), as
well as predation. The latter cause provides an extremely efficient
method of eliminating fleshy tissues and the disarticulation of skele-
tons. Animals that are not digested by carnivores would be subject to
disarticulation by insect activities. Tests were run to determine the
effectiveness of the latter process.

Effects of Subaerial Exposure

Previous wor/:.— Payne (1965) observed activities of insects on baby
pig {Sus scrofa) carcasses. He established six stages of decomposi-
tion—!) fresh stage^ — -no evidence of decomposition or vermin activity,
2) bloated stage— some activity around eyes and orifices, bloating, 3)
active decay stage— fully infested, skin broken in stomach and other
softer areas, 4) advanced decay stage— still infested, skin mostly re-
moved, little remaining fleshy tissue, 5) dry stage— bones isolated, only
small patches of dry skin present and 6) remains stage— only bones
remain and begin to decay.

The rate of decomposition of the fleshy parts of animals and the
durability of bone to subaerial exposure are important in determining
the preservation potential of mammalian bone in the fossil record.
Voorhies (1969) made observations on the decomposition and dis-
articulation of carcasses of calves, coyotes, rabbits, badgers, and rac-
coons in the semiarid environment of northern Nebraska. The calves
reached the decomposition stage 4-5 in 14 days and stage 6 in 90 days.
The smaller animals attained the fifth and sixth decomposition stage
in relatively shorter time, roughly proportional to their body size. The
smallest mammals (rabbits) took only three days to attain stage 4-5.
Clark and Guensburg (1970) observed rates of decay of muskrats,
skunks, rabbits, and domestic cats. The flesh was removed from the
carcasses of the animals in three weeks. Bones were destroyed after
a total of six weeks.

Payne (1965) in a more detailed experiment on foetal pigs, recorded
that the average time necessary to attain the fifth stage (dry stage) was
eight days. Johnson (1975) observed the decomposition of 39 carcasses

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Annals of Carnegie Museum

VOL. 48

Table 6.^ — Rates of decomposition of small mammals in Nebraska during summer 1978.
N = number of specimens observed, MS = average mass of individuals in grams. De-
cay stages as defined by Payne (1965).

Genera

N

MS

Days exposed

0

1

2

3

4

5

8

14

Sorex

4

1.85

1

3

4

5

5

5

6

_

Reithrodontomys

2

6.90

1

2

3

4

4

5

6

_

Blarina

2

16.50

1

2

3

3

4

4-5

5

6

Peromyscus

5

22.65

1

2

3

3

4

4-5

5

6

of squirrel" to opossum-sized mammals in an Illinois forest and showed
that the process is strongly temperature dependent, complete decom-
position taking an average of 48 days in the summer compared to 104
days in the spring and 88 days in the autumn.

Experiments .—Tho, carcasses of several small mammals were placed
outdoors to allow natural decomposition to occur. Most specimens
were placed beneath a weighted screen to prevent large scavengers
from disturbing the specimens. Specimens were observed twice a day
for six days and then once every few days for the remaining time. The
specimens were sheltered from rain. Taxa represented were Sorex,
Reithrodontomys , Blarina, and Peromyscus. Temperatures during the
study varied from a low around 60°F at night and highs up to 100°F
during the days.

The stages of Payne (1965) are used in the analysis of the decom-
position of the small mammals observed in this study. Table 6 relates
the stage of decomposition of each taxon to the day of observation.

Insects had begun to eat the Sorex bones after only eight days, after
which the Sorex and Reithrodontomys specimens were removed from
the test area to prevent any more loss of the skeleton.

A high level of importance has been put on the 4-5 stage of decom-
position in this discussion. Clearly, this stage is the minimum stage
necessary to allow disarticulation of the bone and transport of the
bones (alluvially) as separate elements. As seen in Table 6, the disar-
ticulation and transport stage (4-5) is quickly attained by smaller mam-
mals. Dodson (1973) observed decomposition of a mouse {Mus) car-
cass that was submerged in a tank of pond water. He found that
complete decomposition took 77 days. The average time for Peromys-
cus, subaerially, to decompose (stage 5) was five to six days. The
difference is almost certainly due to the efficiency of carrion-eating
insects compared with aquatic bacteria. Because most small mammals
die on land, the subaerial data on rates are probably more applicable.
Even if the early stages of decay took place in water, bloating would

1979

Korth— =Microvertebrate Taphonomy

261

allow the bodies to float and subsequently to be snagged or beached
where normal subaerial decay would proceed.

The time necessary to liberate bones from connective tissue is im-
portant because the great majority of all micromammal fossil remains
are disarticulated. Articulation would suggest that the connective tis-
sue was still present at the time of burial. It is important to appreciate
that the time necessary to eliminate fleshy tissues from skeletons of
small mammals is extremely short. The present study and those of the
earlier workers cited show that insect carrion activities are nearly as
effective a means of disarticulation as digestive processes of carnivo-
rous mammals and birds, which were offered as a means of disartic-
ulation in the coprocoenosis hypothesis (Mellett, 1974).

Durability of bone .—Tht durability of bone exposed subaerially has
been discussed by Clark et al. (1967), Behrensmeyer (1975, 1978), and
Voorhies (1969). Behrensmeyer (1978) concerned herself with bones
of larger animals and concluded that the time needed to destroy bones
varies with climate and that a maximum of 10 to 15 years is attainable
under the most favorable conditions. She made no attempt to study
the bones of smaller animals. Clark et al. (1967), in reference to bones
of smaller animals, suggested that they would last only a short time of
exposure due to their gracility. Voorhies (1969:31) made actual obser-
vations on bone left exposed for extended periods of time. After one
year of exposure, he observed the amount of decomposition of the
smaller animals tested and noted that they were soft and beginning to
disintegrate. Behrensmeyer (1978) noted that bones of small mammals
crack and splinter into fragments when exposed.

Bones of micromammals cannot endure long periods of time exposed
on the open ground surface. This implies that the bulk of the micro-
vertebrate fossil remains had to be exposed for a short period of time
to allow for the loss of soft tissue, but covered by sediment shortly
afterward in order to escape total destruction. The digestive processes
of carnivorous animals eliminates the soft tissue in a manner as effi-
cient as that of insect activities. In either case, the bone, once stripped
of its fleshy cover, must be buried relatively soon or the bone will be
completely destroyed.

Decomposition of Scatological Material

Tests were conducted on the durability of owl pellets in a stream to
determine whether bone would remain intact in the pellet and be de-
posited as such. If the pellets maintain their integrity in a stream, the
individual bones would not be subject to stream abrasion or sorting
which should be characteristic of fluvially deposited assemblages.

Owl pellets were released in a small stream, averaging 0.6 m deep
and 2 m wide, flowing roughly 30 cm /sec at the surface of the water.

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VOL. 48

Many large limestone blocks were present in the stream and created
minor eddies and areas of rapid flow.

After the pellets were placed in the stream, they became saturated
in the first 5 to 10 sec and settled until about 90% of the pellet was
submerged. They continued to float at this level until the completion
of the experiment. After 20 m of transport, the first bones began falling
from the outer surfaces of the pellet, and continued to do so throughout
the remainder of the test. The pellets became increasingly smaller as
bones and small bunches of fur became dissociated. At 85 m, the
pellets were broken into two or three smaller parts which allowed for
an even higher rate of bone loss. At roughly 200 m, no bones were left
in the remaining pellets, which then consisted only of loosely consob
idated fur.

The small bones once freed from the pellet fell to the bottom of the
stream and continued to be moved downstream. It was impossible to
follow the movements of the bones because they became essentially
invisible against the irregular bottom of the stream.

Because of the buoyancy of the pellets, they followed the main cur-
rent in the few areas where the stones restricted the flow. They rarely
were trapped against rocks or snags. The vertical sides of the stream
were artificially reinforced with large limestone blocks, and thus there
was no chance that the pellets could have been beached on the side
banks. If a pellet became snagged, bones would continue to fall from
it and they would be carried downstream from the snagged pellet.

When held under a faucet with low to moderate flow, owl pellets
break up very rapidly. The first isolated bones fall out after 10 to 15
sec. Collected samples of coyote fecal material when tested broke up
readily under the faucet.

It seems highly unlikely that fecal material would remain cohesive
enough in a stream to be deposited as a scat or pellet accumulation.
The stream would dissect the fecal material even if the scat or pellet
itself was not being transported, and move the bones as individual
particles.

The condition of bone released into a stream from fecal material
would be as described above.

Abrasive Effects of Stream Transport on Bone

Once bone is released into a stream system, it is subject to abrasion.
Most bones of micromammals would probably be whole or nearly
whole when introduced into a stream (see previous sections). In the
present study, tests were run to determine the effect of stream abrasion
on bone. Behrensmeyer (1975) mentions preliminary experiments on
stream abrasion of bone but does not give details.

1979

Korth — Microvertebrate Taphonomy

263

Experiments

An articulated skeleton of an adult Peromyscus, which had been
stripped of flesh by insects, and a disarticulated skeleton of an adult
Microtus, extracted from an owl pellet, were placed in separate tum-
bling barrels. The barrels were half filled with quartz grains averaging
2 to 4 mm in diameter and then filled with water. The barrels were
rotated at a velocity equal to a linear velocity of approximately 24 cm/
sec. Observations on the condition of the skeletons were made ap-
proximately every 10 h, and tumbled for a total of about 80 h.

The first effect of tumbling was the disarticulation of the Peromyscus
skeleton. This was probably partially due to the presence of the water
softening and weakening the remaining tendons and connective tissue.
All of the rootless teeth in the skull and jaws of the Microtus specimen
except the incisors separated after only a short period of time. The
rooted teeth of the Peromyscus specimen remained in the jaw and
skull throughout the experiment.

The skull of Microtus first lost the auditory bullae and nasal bones,
then the occiput and interparietal bone, and progressively disassociat-
ed the cranial bones, beginning with those at the base of the skull and
then those more anterior until all bones had been separated. The skull
of Peromyscus followed the same pattern but did not start to break up
until the Microtus skull was nearly completely disarticulated. The cra-
nial bones separated from the other components of the skull along the
sutures and did not fracture at the time of separation. Once the delicate
cranial bones had separated from the skull, they began to be rounded
along the edges and thinned in the central areas. Small holes appeared
in cranial bones in areas of thinning after considerable transport, once
they had been separated from the cranium (Figs. 1 , 5A). The only bones
of the skull which showed breakage before separation were the zy-
gomatic arches, which broke away quickly, the pre maxillae, which
showed breakage along the socket for the upper incisors, and the max-
illae, in which the sockets for the teeth were broken away (dorsally)
once the base of the skull was gone and the sockets were exposed to
abrasion. The breakage of the maxillae and premaxillae was due to the
thinning process described for the isolated cranial bones. The similar-
ity between skulls partially disarticulated by stream action and those
regurgitated by owls is striking. It would be virtually impossible to
distinguish by which mechanism a skull with the bones of the base
missing was produced. Only if there is evidence of the thinning of bone
and rounded holes in the thinned areas, can such material be assigned
to a particular source.

The mandibles of the two tested specimens were eroded at about
the same rate, although the Peromyscus retained all teeth and the

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Microtus did not. The first signs of erosion of the dentary were the
wearing away of the coronoid process and the angle of the jaw. Holes
were broken in the jaw laterally at the pulp cavity of the incisor and
at the base of the sockets for the molars (Fig. 2). The bone gradually
chipped away from around the incisor throughout its entire length. The
anterior tip of the incisor showed some attrition.

The isolated teeth of Microtus showed some attrition on their sharp
edges but no breakage. The single isolated incisor of Microtus was
eventually broken in half.

Pelves (innominates) showed signs of corrosion quite early in the
experiment. Evidence of abrasion on the iliac blade was present after
only a short period of tumbling. The pubic ramus was also quickly
broken away (Fig. 5B). The entire pubis and ischium were eventually
eliminated.

Of the long bones, the scapula was most easily affected. Almost
immediately, small flakes of bone from the dorsal border of the scapula
were broken off, leaving an irregular but smoothly rounded edge.
Shortly afterward, the dorsal spine and acromion and coracoid pro-
cesses were broken away. Only the glenoid fossa and proximal rem-
nants of the base of the spine and cranial and caudal borders remained
by the end of the tumbling experiment (Fig. 4B).

The first breakage on the major limb bones was the loss of the fibula
from the body of the tibia-fibula. The only other damage to the limb
bones produced during the experiments was the gradual erosion of the
ends and prominent ridges and crests such as the trochanters of the
femur, the medial epicondyle of the humerus, and the lateral malleolus
of the tibia-fibula (Figs. 3, 4A). No breakage of the diaphyses of the
limb bones occurred at this point. The breakage of the shaft of a limb
bone only occurred where a prominent crest on the diaphysis was
eroded away and the internal cavity was exposed. Eventually this pro-
cess would result in a major break of the diaphysis, but only after
extensive rounding of the epiphyses.

The major tarsal elements, calcaneum and astragalus, showed little
evidence of abrasion. The distal end of the calcaneum had a minor
amount of abrasion on it. The astragalus showed minor attrition on the
dorsal surface of the tibial facet and the ventral surface of the naviculo-
astragular facet.

Phalanges and metapodials were difficult to observe because of their
small size which made the periodic recovery of these elements nearly
impossible. However, by the end of the tumbling experiment, the re-
covered phalanges and metapodials were not noticeably abraded and
still intact.

Although no isolated rooted teeth were tested, it is most likely that
roots of these teeth would be lost as are all small processes on the

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other bone, such as the zygomatic arches and fibula. Small fragments
of larger bone derived from carnivore feces or some other source,
should show rounding of the edges and loss of any elongate fragments
that may have been produced by its initial crushing.

Interpretations

As previously noted, major limb bones are not broken until exten-
sive corrosion has affected the epiphysis. Breakage of limb bones with
little or no signs of attrition may occur during subaerial exposure due
to drying and splintering (Behrensmeyer, 1978) or may be due to im-
pact of relatively high velocity while in the stream flow. A bone that
has been subjected to subaerial conditions would be dried and would
become quite brittle and subject to fracture with very little pressure.
The low velocity generated in the tumbling experiments was not high
enough to produce velocities that would create a force of impact great
enough to break the limb bones. Natural stream velocities, which are
many times higher than that of the tumbling barrels in the present
experiment (Leopold et al., 1964), may be sufficient to cause breakage
of small bones.

Recognition of Fluvial Accumulations in the
Fossil Record

The percentage representation of the various skeletal elements in a
stream sorted assemblage would probably differ from that of a fecal
or pellet accumulation (Table 1). Wolff (1973) reported the percentage
representation of elements from a fluvially-deposited Pleistocene fossil
assemblage. The best represented elements in his collection were in-
cisors which had only a 48% level of expected representation. The
mean percentage representation for Wolffs collection was only 12%.
The percentage curves for fecal or pellet assemblages are several times
higher (Fig. 6), with mean percentage values of five to six times that
reported by Wolff (1973). The alluvial hypothesis of microvertebrate
accumulation would predict results similar to that presented by Wolff.

For each individual taxon in a fluvial assemblage, a particular ele-
ment or elements may be markedly abundant and all others nearly
absent. This is shown by Wolff (1973:95). The most abundant elements
for each species represented may be different depending on the size
of the animal and hydraulic properties of the element. The overall
curve of percentage representation would be very low with a low mean
percentage. Some elements may be much better represented than the
others depending on the size distribution of the animals represented
and the relative abundance of each taxon in the fauna.

In a sorted assemblage, the degree and amount of abrasion on each
bone should be variable. If shape and density are the major factors in

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determining the sequence of sorting (Behrensmeyer, 1975), those ele-
ments introduced into the stream near the area of deposition would be
less abraded than those that were subject to transport for a consider-
able distance.

A fluvial accumulation of microvertebrate bones should be recog-
nizable in the fossil record through the following characteristics: 1) the
sediment enclosing the fossils should generally show primary sedi-
mentary structures of alluvial origin; 2) the modal size of sand grains
in the deposit should be predictable from the size and hydraulic char-
acteristics of the skeletal elements present, 3) the bones should show
varying degrees of stream abrasion, and 4) the percentage represen-
tation of bones of the entire fauna should be low.

Fossil Examples

In order to test the reliability of the previously established criteria
for determining the origin of microvertebrate fossil assemblages, fossil
material from two localities was examined. Both localities produced
almost exclusively remains of small vertebrates, mostly mammals. The
sediment at both localities consisted predominantly of clean quartz
sand with clay clasts.

Localities Tested

University of Nebraska State Museum Locality Bw 110 (Achilles
Quarry) is located in Brown County, Nebraska (SWI4 SWV4 NE!4
SW14, section 31, T33N, R23W). It is situated on the north bank of
Fairfield Creek, 72 ft above the creek and 39 ft above the contact of
the Valentine Formation with the underlying Rosebud Formation. Uni-
versity of Nebraska State Museum Locality Kx 110 (Annie’s Geese
Cross = Locality “B” of Voorhies, 1971) is in Knox County, Ne-
braska (NWi/4, section 23, T33N, R3W).

Both localities are within the Crookston Bridge Member of the Val-
entine Formation (Skinner et al., 1968), of late Barstovian Age (Webb,
1969; Voorhies, 1971).

Condition of the Bone

The condition of the bone from both localities is nearly identical to
that of the bone experimentally abraded in the tumbling barrels. The
small processes and ridges of the long bones are characteristically
abraded as they are in the tumbled bones (Figs. 9, 10). The most
obvious examples of rounded broken edges are in the scapulae, where
the breakage of the fossils is nearly identical to that of the tumbled
Recent bones (Fig. lOB). The process of bone thinning and final break-
age described in the abrasion experiments is also evident in the ace-
tabula of the fossil pelves (Fig. IIB).

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Fig. 9.— A) Tibia-fibulae of small mammals experimentally abraded (left) and from fossil
localities Kx 1 10 (center) and Bw 1 10 (right). Fossil bones show similar abrasion of ends
and loss of the fibula as in the abraded bone (see Fig. 3A). Scale x3.1. B) Humerus of
a rodent experimentally abraded in a tumbling barrel (left) and fossil humeri from lo-
calities Kx 110 (center) and Bw 110 (right). Fossil bones show similar abrasion of ends
and processes seen in the tumbled bone (see Fig. 3B). Scale x4.1.

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Fig. 10. — A) Femur of a rodent experimentally abraded in a tumbling barrel (left) and
fossil femora from localities Kx 110 (center) and Bw 110 (right). Fossil bones show
similar abrasion of proximal end seen in the tumbled bone. (See text for explanation of
breakage of the shaft of the fossils.) Scale x3.5, B) Rodent scapula experimentally abrad-
ed in a tumbling barrel (left) and a fossil scapula from locality Kx 1 10. Fossil bone is
similarly abraded as is the tumbled bone (see Fig. 4B). Scale x4.

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Fig. 11. — A) Rodent maxillae experimentally abraded in a tumbling barrel (left) and
maxillae of fossil rodents from localities Kx 1 10 (center) and Bw 1 10 (right) in dorsal
view. Fossil maxillae show breakage of sockets for teeth dorsally, as in the tumbled
maxillae. The specimen on the right still contains the teeth, hence, breakage is not as
noticeable. Scale x3.3. B) Pelves experimentally abraded in a tumbling barrel (left) and
fossil pelves from localities Kx 1 10 (center) and Bw 1 10 (right) in dorsal view. Fossils
show similar abrasion of edges as in the tumbled bone. Breakage in the acetabulae of
the fossils is similar to the thinning and breakage seen in other tumbled bones (see Figs.
2 and 4A). Scale x3.6.

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Fig. 12. — A) Medial view of a mandible experimentally abraded in a tumbling barrel
(left) and two mandibles of fossil rodents from locality Kx 1 10 (center and right). Fossil
bones show characteristic breakage of abraded bone along the ventral surface of the
incisor and loss of the angle, coronoid process, and condyle. Scale x3.6. B) Lateral
view of the same. Fossil bones have breaks at the base of the sockets of the teeth and
at the pulp cavity for the incisor as in the abraded bone (see Fig. 2). Scale x3.6.

The most remarkable similarity to tumbled bones is in the condition
of the fossil mandibles. The breakage of the lateral surface of the
mandible at the base of the sockets for the teeth and at the pulp cavity
of the incisor is present in differing degrees in the fossil material (Fig.

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271

12). This is precisely the type of breakage seen in abraded bones but
not in the bones from the carnivore feces or owl pellets. Also, the
bone which surrounds the incisors of rodents on the ventral surface of
the mandible is reduced or absent on the fossils, as in the tumbled
mandibles. In some instances the entire incisor has been lost in the
fossil specimens because the ventral surface of the bone has been
totally eliminated. The holes broken through the wall of the mandible
are rounded on their edges as in the tumbled bone.

The only recovered cranial materials are maxillae. Virtually all max-
illae show breakage on the dorsal surface at the base of the sockets of
the teeth in much the same manner as the Microtus maxilla used in
the tumbling experiment (Fig. IIA). In one instance, the entire maxilla
of a plesiosoricid insect! vore is preserved including the rostral, orbital,
and zygomatic contributions of the bone. There is no evidence of
breakage on this specimen, but the edges are very well rounded and
a small hole has broken through the dorsal surface of the palate as in
the tumbled bone (Fig. 13).

Isolated teeth from the fossil accumulations commonly lack roots,
as was predicted for stream abraded teeth.

The only aspect of the condition of the fossil bone not found in the
experimentally abraded bone is the abundance of broken diaphyses of
the major limb bones. The majority of the major limb bones are rep-
resented by half of the bone or less. Bones from the Achilles Quarry
(Bw 110) are more frequently broken than are those from the Annie’s
Geese Cross Locality (Kx 110). Of 154 major limb bones recovered
from the former locality, only four bones are whole. A greater percent
of bones from Annie’s Geese Cross are unbroken but still show only
very low percentages of whole bones. Sixty-four percent of the radii
recovered from the latter locality are whole. This is the highest per-
centage of whole bones in the assemblage. Thirty-eight percent of the
humeri are whole. Only 15%, 10%, and 3% of the ulnae, femora, and
tibiae, respectively, are whole. Bone recovered from carnivore feces
are commonly broken in half. Two differences are evident between
the broken bones from the fossil sites and those in the coyote feces.
Although some bones from the coyote feces are broken, the percentage
of whole elements is much higher than in the fossil material. The ele-
ments that are most frequently broken in the carnivore feces are the
femur and humerus. Sixty-two percent of the identified humeri and
femora were whole. Tibiae were whole in 68% of the cases, and the
radii and ulnae were nearly always whole. Thus the percentage of
whole elements is markedly higher in the carnivore fecal material than
in the fossil samples.

A second difference between the material from the coyote feces and
the fossil bone is in the occurrence of a preferred end of the major limb

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Fig. 13.-— A) Fossil maxillary bone of a plesiosoricid from Kx 110 in lateral view. Note
rounded edges of bone especially of the zygomatic arch, and breakage along root sockets
for the teeth. Scale x3.2. B) Dorsal view of the same. Note breakage on dorsal surface
of the maxilla similar to that in experimentally abraded bones (see Fig. 1 lA). Scale x3.2.

bones. In the fossil material, 85% of the femora are proximal
ends and only 2% are distal. Over 80% of the tibiae are repre-
sented by distal ends, whereas only 5% of the specimens were prox-
imal. Seventy percent of the humeri were distal halves of the bone,
but only 3% were proximal halves. Similar biases in the preserved
ends of long bones were recorded by Wolff (1973) in his Pleistocene

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micromammal assemblage. He noted a similarity with the results of
Voorhies (1969). However, Voorhies dealt with larger animals and
attributed the relative abundance of selected ends of long bones to
activities of large scavangers, which would most likely destroy the end
of the bone that contained the most flesh. This selective scavenging is
probably not the same factor that alters the abundance in the micro-
mammals because these animals are quite small and if ingested, the
entire animal, or major portions of the animal are swallowed, and no
preference is shown for a particular end of a bone.

A bias in proximal versus distal ends of long bones is also seen in
bone from coyote feces. The most marked difference in selective pres-
ervation of ends is in the femora and humeri, where the preferred end
(proximal for the femora and distal for the humeri) is twice as abundant
as the other. This bias of preferred ends is much lower, however, than
the same ratio reported for the fossil localities. The only other bone
in the fecal material that showed a preference in the epiphysis present
was the tibia. In this bone, 9% of the specimens were proximal frag-
ments and 5% were distal. Distal tibiae are nearly twenty times more
abundant than the proximal fragments in the fossil assemblages.

The high frequency of partial limb bones in the two fossil assem-
blages tested may well be due to breakage caused by impact while
being transported in the stream, splintering or crushing during subae-
rial exposure, or even breakage due to ingestion by predators. The
preference of a particular end over another is probably due to selective
sorting because different ends of bones show differential hydraulic
behavior (see settling results for Peromyscus, Table 2),

There is a lot of fragmental bone of larger animals in the recovered
fossil material. Similar fragmented bone is present in remains from the
coyote feces. The rounding of the edges of the bone, however, indicate
that the bone has been transported in a stream environment and sub-
jected to abrasion. A few specimens of fossil bone fragments show
cracking and chipping of outer layers (Fig. 7), which may indicate
subaerial exposure of the bone for a considerable amount of time (Beh-
rensmeyer, 1978).

The occurrence of elements of larger taxa represented by whole
elements (notably the perissodactyls in Bw 110 and an antilocaprid
from Kx 110) show no signs of being ingested (tooth marks or sharp
angular breakage) but do show effects of limited stream abrasion. This
suggests that these elements were also transported into the area of
accumulation and deposited. Teleoceras (a rhinoceros) from Bw 110
is represented by a single navicular bone. An indeterminate equid from
this locality is represented by a single astragalus. A specimen of Mer-
ycodus (an antilocaprid) from Annie’s Geese Cross is a skull. It is
difficult to determine the condition of this skull because it was partially

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Fig. 14. — Profile curves of percentage representation of small mammal bones from two
fossil localities (data from Table 1). Horizontal axis represents individual elements (see
Fig. 6 for definition of symbols).

destroyed when it was discovered. The remaining half is little damaged
and hardly abraded, however, so it was not transported far.

The screen washing process of obtaining the fossils may have caused
some damage to the bone. It is difficult to determine how much dam-
age is done. However, the washing process takes only 30 sec of agi-
tation in water, and mineralized bone such as that at both sites is not
likely to be significantly affected.

In summary, the condition of the fossil material from the Nebraska
localities more closely resembles the experimentally abraded bones
than bone ingested by predators.

Percentage Representation

Although informative in determining the source of fossil micromam-
mal bone, the condition of isolated bones is not conclusive in all cases.
A second factor that can help determine the source of bone in a fossil
assemblage is the percentage representation of elements in the accu-
mulation.

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275

The percentage representation of elements from Recent predator
feces (or pellets) is quite high except for the tarsal elements (Fig. 6).
The mean values for percentage representation in the modern carni-
vore feces and owl pellets analyzed by the writer range from 65% to
78% (Table 1). It is postulated that a coprocoenosis, whether from
birds, mammals, or both, would have a mean percentage of represen-
tation within this range with the lowest values for the tarsal elements.

An assemblage of bone that has been subjected to alluvial processes,
on the other hand, should have a low mean value for percentage rep-
resentation and one or two elements in abundance. An example of
such an accumulation was described by Wolff (1973).

Effects of Screening

As noted in the discussion of bone condition, the screening pro-
cesses of recovering the fossils may have some effect on the relative
numbers of different skeletal elements in the sample. Undoubtedly,
very small bones such as phalanges or very small isolated cheek teeth,
were lost through the screens during processing. The loss of apprecia-
ble numbers of elements would lower the percentage representation
for these elements and, in turn, would lower the mean value for the
percentage of representation. The following observations, based on
examination of the percentage representation curves for the fossil lo-
calities, indicate that screening has little effect on the curve. Elements
such as the small tarsals are among the most commonly preserved
elements and larger elements that could not be lost through the screens
(pelves and scapulae) are among the lowest represented. Long thin
elements (radii and ulnae) may pass through the screens and thus lower
the percentages of these bones but because tarsal elements of equal or
smaller diameter are well represented, such loss appears unlikely. The
washing has little, if any, effect on the percentage preservation of the
studied elements. The relative percentages of elements represented are
thus probably very near the actual amounts preserved in the sediment
before processing.

Results

The percentage of representation curves for the fossil localities un-
der discussion (Fig. 14) are much closer to that of the stream sorted
assemblage than that of a coprocoenosis (Fig. 6). The mean value of
percentage of representation of the Achilles Quarry (Bw 110) is only
17%, or about one-fourth of the mean percentage of representation
value for bones taken from carnivore feces or owl pellets. The per-
centage preservation for Annie's Geese Cross (Kx 110) is twice that
of Achilles Quarry (36%) but still only about half of the predicted value
for a coprocoenosis (Table 1). Although the values for percentage of

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representation for Kx 110 are higher than those for Bw 110 for all
elements except one, the general shape of the curves are quite similar
(Fig. 14) and clearly distinct from the predator curves. The occurrence
of relatively high percentages for selected elements in the fossil locaF
ities (femora in Kx 110 and tarsals in Bw 110) shows selective sorting
and attests to the fact that the environments of deposition in both
localities were not identical.

Thus the percentage of representation, as well as the condition of
the fossil bone, indicates that the fossil accumulations under discussion
are alluvially deposited assemblages.

Sedimentological Evidence

If alluvial processes are the major factor in determining the com-
position of micromammalian fossil assemblage, the bones should be
hydraulically equivalent to the associated sedimentary particles. The
method of determining if the bones and the sediment are hydraulically
equivalent is to determine the value of the bone from the equation
presented for the calculation of dq , and compare the determined value
with the actual size of the quartz grains associated with the bones, as
determined by sieve analysis. This procedure was followed for both
fossil localities.

The quantitative value used in the predictive model of bone equiv-
alents is based on the most common elements of the most abundant
taxa(on). This value, or “abundance,” for each element is calculated
by multiplying the percentage of representation of that element by the
relative abundance of the taxon to which it belongs (expressed as a
percentage of the total number of individuals in the fauna). The per-
centage of representation used in this calculation is not the overall
percentage for the entire fauna, but the percentage for the particular
taxon to which the elements belong.

Only those animals that represented over 5% of the fauna were used.

Hydraulic Equivalents of Micromammal Fossils at
Kx 110 and Bw 110

Only two animals represented a significant portion of the fauna from
the Achilles Quarry, the zapodid rodent Megasminthus tiheni and a
heteromyid rodent Cupidinimus sp. A. The former made up 14% of
the total mammalian fauna, and the latter, 62% of the fauna. Only one
species, Cupidinimus sp. B, was significantly represented from An-
nie’s Geese Cross (Kx 110). It made up 63% of the total fauna. No
other species made up more than 5% of the fauna.

Because no cranial material was present in the fossil accumulations,
estimates for the area of the foramen magnum of the fossil taxa were
determined by the proportional size of this foramen in closely related

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277

Table 7,— Quartz equivalence values (dq) of most abundant skeletal elements of the
predominant mammals at Bw 110 and Kx 110 (see text for definition of '‘abundance”).

Taxon

Element

Abundance

d.

0

Locality Bw 110

Cupidinimus sp. A

calcaneum

.53

0.54

1.00

Cupidinimus sp. A

astragalus

.12

0.51

1.00

Cupidinimus sp. A

distal humerus

.06

0.36

1.50

Cupidinimus sp. A

distal tibia

.05

0.52

1.00

Cupidinimus sp. A

cheek teeth

.04

0.97

0.50

Megasminthus tiheni

mandible

.06

1.53

-0.50

Megasminthus tiheni

maxilla

.05

0.65

1.00

Megasminthus tiheni

cheek teeth

.06

1.12

0.00

Locality Kx 1 10

Cupidinimus sp. B

proximal femur

.47

0.24

2.50

Cupidinimus sp. B

distal tibia

.46

0.42

1.50

Cupidinimus sp. B

calcaneum

.33

0.44

1.50

Cupidinimus sp. B

distal humerus

.32

0.29

2.00

Cupidinimus sp. B

mandible

.28

1.07

0.00

Cupidinimus sp. B

femur

.16

0.65

1.00

Cupidinimus sp. B

maxilla

.12

0.45

1.50

Cupidinimus sp. B

ulna

.12

0.19

2.50

extant animals. Size estimates of the fossil taxa were based on mea-
surements of the dentition. The living species used to estimate the area
of the foramen magnum of the fossil heteromyids was Perognathus.
This species is quite similar in its dentition to the fossil species and
there is no evidence that the dentitions of the fossil species are dis-
proportionate to the skull (as in Dipodomys). The skull of Cupidinimus
is known (Wood, 1935) and shows no marked reduction or enlargement
of the dentition relative to the modern Perognathus. The calculated
area of the foramen magnum for Cupidinimus sp. A from Bw 110 was
0.158 cm^, and for Cupidinimus sp. B from Kx 110 was 0.127 cm^.

Similarly, the area of the foramen magnum for the fossil zapodid
was derived from measurements of the Recent zapodid Zapus hud-
sonius. The calculated area for the foramen magnum of Megasminthus
tiheni was 0.182 cm^.

The unassociated postcranial elements were identified and referred
to the fossil species on the basis of 1) comparison with Recent forms,
2) comparison with fossil forms for which postcranial elements have
been described, and 3) size.

The zapodid from the Achilles Quarry was well represented only by
its dentitions, so the mandibles, maxillae, and isolated teeth were the
only elements used for determinations of dq. The heteromyid utilized

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Bw 110

Fig. 15.- — Histograms of grain size of sediment associated with microvertebrate fossil
material (above) and predicted quartz grain diameter based on bone elements present
in the assemblage (below). Horizontal axis of the lower in terms of dq (see text for

definition of “abundance”). Data from locality Bw 110.

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279

Kx 110

Fig, 16.— Histograms of grain size of sediment associated with microvertebrate fossil
material (above) and predicted quartz grain diameter based on bone elements present
in fossil assemblage. Data from locality Kx 1 10 (Table 7). See Fig. 15 for further expla-
nation of histograms.

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in the determination was significantly represented by calcanea, as-
tragali, tibiae (distal halves), humeri (distal halves), and cheek teeth.
Incisors were not used though well represented because hydraulic ex-
periments were run on whole rodent incisors only and most of the
fossil isolated incisors were fragmentary. The Kg values of the distal
humeri and distal tibiae of the heteromyid and the maxillae of the
zapodid were determined from the settling results of the Peromyscus
specimen (Tables 2 and 5). The determinations of dq are listed in Table
7. The determined dq values were converted to the nearest 0.5 0-size
and plotted on a histogram (Figs. 15, 16). When the calculated 0-size
of several elements were identical, the abundances were added. This
entire procedure was repeated for the fossils from Annie’s Geese Cross
Quarry.

The histograms of dq for Bw 110 shows a marked abundance at 1.00
0, with some minor amounts of abundance for surrounding 0-sizes
(Fig. 15B). The graph for Kx 110 (Fig. 16B) shows an equally marked
abundance at 1.50 0 to 2.50 0, again with minor abundances in 0-
sizes slightly smaller and larger.

Sieve Analysis

Sediment samples directly associated with the fossils from both lo-
calities were processed in a set of sieving screens. There was no evi-
dence of diagenesis of the sediment (regrowth on quartz grains or
alteration of feldspars).

The grain sizes of the sediments are apparently bimodal (Figs. 15A,
16A). The obvious peak in the coarser fraction consists exclusively of
clay clasts. The clasts were evidently transported to the locality and
not formed in the fossiliferous deposit diagenetically.

The relatively high amount of clay-sized fraction in the Bw 1 10 sam-
ples is most likely due to the breaking up of some of the clay clasts
during the sieving process.

The peak in the medium to fine sand-size present in the sediment
from both localities, is composed of a clean sand. The grains are very
well rounded. This sand is almost entirely quartz. Some feldspar grains
and rock fragments are present but in a minor amount (less than 5%).

The modal size range for the quartz sand from Bw 110 is 1.00 0 to
2.00 0. The modal size from Kx 110 is 2.50 0. Because the equation
for dq is based on quartz only, the modal values of the quartz sand are
the applicable data.

Comparison of Sediment Size to dq

For the Achilles Quarry (Bw 110) the modal quartz size includes the
determined dq within its range. The range of the sediment is 1.00 0
to 2.00 0, and the determined dq is 1.00 0. The determined dq is not

1979

Koeth— Microvertebrate Taphonomy

281

ie the center of the modal range as may have been expected, but sieve
results can be somewhat misleading. Blatt et al. (1972:48) state, ‘7 . .
if the screen opening is equated with particle size, the modal inter-
mediate diameter of particles in each sieved fraction is frequently
underestimated by 10 to 20%7’ Thus, with the consideration of a minor
shift of the sieve data to a coarser size, the determined dq is almost
exactly equivalent to the actual sediment size.

The sediment from Annie's Geese Cross (Kx 110) has a modal size
restricted to 2.50 0. The determined dq has a modal range from
L50 0 to 2.50 0, Again, as in the previous example, the sediment
size calculated from the fossil data is extremely close to that of the
actual sediment. The modal sand size at Kx 110 is 0.5 0 smaller than
the calculated value for dq as at Bw 110, and the difference can be
explained by the same reasoning.

Clearly, from the sedimeetological evidence the fossil assemblages
have been sorted with the surrounding sediment as was predicted for
the alluvial hypothesis of micro vertebrate fossil accumulation.

Conclusions

From the evidence presented on the condition of the fossil bone, the
percentage preservation of elements and the hydraulic equivalence of
bones and associated sediment, it is clear that the University of Ne-
braska State Museum localities Bw 110 and Kx 110 are alluvially
sorted.

Summary and Conclusions
Application

The importance of this study is that repeatable procedures have been
established for the determination of the means of deposition of mi-
crovertebrate fossil accumulations. The observations on the condition
of the fossil material and the percentage of representation of the bones
present can be determined for any fossil accumulation, and conclu-
sions can be drawn from them. The determination of sediment equiv-
alence is a more complex operation but equally applicable.

Microvertebrate fossil assemblages are more likely a product of se-
lective sorting and deposition by alluvial processes than accumulations
of bone material from carnivore feces or owl pellets. This statement
is based on the fact that carnivore feces and owl pellets are easily
destroyed either ie a stream flow or subaerially (by rain showers) and
that only very slight currents are n^wkci to transport isolated skeletal
elements (Dodson, 1973). Currents of surface runoff would be suffi-
cient to transport many elements of small animals, and could result in
an accumulation of bone in a fine mud, and ie sediments not associated

282

Annals of Carnegie Museum

VOL. 48

with any channelized stream flow. Once again, the key to the inter-
pretation of the origin of a fossil accumulation is to deal with the entire
assemblage and not to deal with just a few isolated bones.

The original source of microvertebrate fossil accumulations may
well be as fecal material or regurgitated pellets of owls, but due to the
lack of alteration of most of the bones of smaller animals by the pred-
ators studied, and the strong effect of stream processes, the actual
origin of an assemblage is masked.

Paleoecological Interpretations

Shotwell (1955, 1958) proposed that membership in “proximal”
(nearby) and “distal” (more distant) communities could be interpreted
from transported fossil assemblages by counting the number of ele-
ments per individual per taxon. This theory has been discussed by
numerous authors (Voorhies, 1969; Wilson, 1960; Clark et al., 1967;
Clark and Guensburg, 1970; Wolff, 1973; Behrensmeyer, 1975; Gray-
son, 1978) who have shown that such determinations of proximity are
not wholly applicable in most situations. Wolff (1973) argues most con-
vincingly against its application to micromammal assemblages.

Dodson (1973) concluded that stream processes would obscure the
paleoecologic reconstruction of fossil communities, based on flume
experiments on the bones of a mouse. From the evidence presented
here from two fossil localities, it appears that stream processes are
responsible for the relative abundance of particular species over oth-
ers. The differing degrees of abrasion on the elements of the dominant
species indicates that some individuals were introduced into the stream
flow system nearer the point of deposition than others, and that per-
haps the entire drainage system above the point of deposition is being
sampled.

The indication is that relative population size in the paleocommunity
cannot be determined from relative abundance of fossil forms in a
fluvial assemblage. Qualitatively, however, the fauna recovered may
provide an indication of the taxa present in the communities upstream
even though no valid estimate of relative population size can be made.

Acknowledgments

I am deeply indebted to Dr. M. R. Voorhies for the great amount of time and effort
he spent on directing my research. I also wish to thank the following people who con-
tributed significantly to my research in various ways: R. Corner, M. Dawson, B. Evan-
der, R. Hunt, K. Kolster, J. Lynch, P. McCabe, T. Moeglin, C. Messenger, and J.
Sutton.

The University of Nebraska State Museum, Division of Vertebrate Paleontology sup-
plied the author with a salary during the summer of 1977 when the field research was
being conducted.

1979

Korth— Microvertebrate Taphonomy

283

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Korth— Microvertebrate Taphonomy

285

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; VOLUME 48 7 SEPTEMBER 1979 ARTICLE 16

i

I GEOMYOID RODENTS FROM THE VALENTINE FORMATION
OF KNOX COUNTY, NEBRASKA

William W. Korth

' Rea Fellow, Section of Vertebrate Fossils

I

j Abstract

'! Four heteromyids {Cupidinimus nebraskensis, Perognathus furlongi, Perognathus
I' trojectioansmm, new species, and ^"Diprionomys” agrarius) and a geomyid {Lignimus
is cf. L. hibbardi) are recognized from the Crookston Bridge Member of the Valentine
[i Formation, University of Nebraska State Museum locality Kx 110. The large sample of
Cupidinimus nebraskensis collected from this locality provides evidence for the syn-
onymy of Cupidinimus and Perognathoides. The subfamilial assignment of the geomyid
is questioned.

The geomyoids present in locality Kx 1 10 are a combination of Barstovian and Clar-
endonian species, thus, little can be said concerning the definite age of the Valentine
Formation from these forms alone.

Introduction

The age of the Valentine Formation of Nebraska (in terms of pro-
vincial ages of Wood et al. , 1941) has long been argued, with very little
understanding of the micromammals from the formation (see Skinner
et al., 1968; Webb, 1969; Schultz et al., 1970). In recent years extensive
j I collecting of fossil microvertebrates by means of screen washing by
i I field parties of the University of Nebraska State Museum has been
i I undertaken in the Crookston Bridge Member of this formation. One

I I of the most fossiliferous localities is Annie’s Geese Cross Quarry
I (UNSM locality Kx 110) in Knox County, Nebraska. Voorhies (1971)

I briefly described the location of this quarry (which he called locality

Submitted for publication 12 February 1979.

287

288

Annals of Carnegie Museum

VOL. 48

“B”) and its stratigraphic position. Korth (1979) discussed the ta-
phonomy of this quarry.

This paper will describe a part of the mammalian fauna from Kx
110, the geomyoid rodents. A greater knowledge of this fauna will
yield a better understanding of the age of the Valentine Formation in
relation to other North American faunas.

Abbreviations used: CM = Carnegie Museum of Natural History;
UMMP = University of Michigan Museum of Paletontology; UNSM =
University of Nebraska State Museum; N = number of specimens
measured; M = mean; SD = standard deviation; CV = coefficient of
variation; OR = observed size range. Cusp nomenclature follows
Rensberger (1971). Measurements equal the maximum width of the
protoloph or metalophid (tra), metaloph or hypolophid (trp), and
anteroposterior length (A-P) of measured teeth taken at the base of the
crown. A-P measurement on P'* taken parallel to occlusal surface ex-
cept for Lignimus cf. L. hibbardi, which was taken perpendicular to
the posterior wall of the tooth. All measurements in millimeters.

Systematic Paleontology

Order Rodentia
Superfamily Geomyoidea
Family Heteromyidae
Perognathus furlongi Gazin, 1930
(Figs. 1, 2 and Table 1)

Three specimens are referable to this species—UNSM 56300, a complete mandible
with all teeth present; UNSM 56301, a maxilla with P‘‘-M^; and UNSM 56314, a partial
right mandible with Mg-Mg.

The size of these specimens is comparable to that previously re-
corded for Perognathus furlongi (Wood, 1935; James, 1963; Lindsay,
1972). The lower dentition of UNSM 56300 falls into the lower part of
the size range of P. furlongi, but UNSM 56314 is slightly larger and
more nearly approaches the mean sizes of this species (Lindsay,
1972:43). P4 of UNSM 56300 is below the observed range of size of
this tooth in the California sample. This minor difference in size of P4
cannot be viewed by itself as a diagnostic feature of the Crookston
Bridge form in light of the amount of variation of many characters of
P. furlongi (James, 1963) and the small number of specimens referred
to this species (Wood, 1935; James, 1963; Lindsay, 1972).

The lower molars of the UNSM specimens agree in morphology with
the referred specimens of P. furlongi. P4, however, is much shorter
anteroposteriorly than in the previously described specimens. The
cusps of the hypolophid of P4 are set well apart as in other specimens
of P. furlongi. A minor swelling (hypoconulid) occurs between and

1979

Korth— Valentine Geomyoid Rodents

289

Fig. l.—Perognathus furlongi, stereoview, UNSM 56301, x5.25.

Fig. 2.—Perognathus furlongi, stereoview, UNSM 56300, RP4-M3. x5.25.

290

Annals of Carnegie Museum

VOL. 48

Table \.— Measurements of the dentition o/ Perognathus furlongi.

UNSM

no.

p4

M‘

M"

M3

alveo-

lar

length

A-P

tra

tip

A-P

tra

tip

A-P

tra

tip

A-P

tra

trp

56301

1.42

0.68

1.21

0.87

1.17

1.11

0.70

1.15

1.07

0.70

0.91

0.81

4.15

UNSM

no.

P4

M,

M3

M3

alveo-

lar

length

A-P

tra

tip

A-P

tra

trp

A-P

tra

tip

A-P

tra

trp

56300

56314

0.68

0.63

0.79

0.86

0.99

1.07

0.80

LOO

1.06

1.12

1.01

0.98

0.72

0.82

0.88

0.96

0.71

0.72

4.03

distinct from the entoconid and hypoconid. The two cusps of the meta-
lophid are much smaller than those of the hypolophid and positioned
closer to the center of the tooth. The anterior cusps are closely ap-
pressed against the hypolophid, making the outline of the tooth nearly
triangular with a slightly broadened anterior apex.

The upper premolar and molars of the UNSM specimen agree in
shape and crown morphology with those in other P. furlongi. ap-
pears to be nearly circular in occlusal outline, much more so than is
described in published material. This may be a minor variation in tooth
morphology and not a specifically distinct character.

Perognathus trojectioamrum, new species
(Figs. 3, 7 and Table 2)

Perognathus sp. Klingener, 1968:68.

//o/otypc— UNSM 56311, right mandible with P4-M3.

Horizon and locality Barstovian (Miocene) Crookston
Bridge Member, Valentine Formation, UNSM locality Kx 110 and Nor-
den Bridge Quarry (UNSM locality BW 106), Brown County, Nebras-
lc3.

T/ypoJi^m.^Holotype and UNSM 56304-56309, UNSM 56312, and
UMMP 53019 (partial or whole mandibles with lower teeth); UNSM
56302, 56303, 56310, 56313, and UMMP 53025 (partial maxillae with
upper teeth).

Etymology.— trojectio, Latin, crossing; ansrum, Latin, genitive plural of goose, meant
to indicate the locality where the type was recovered.

Diagnosis.— -SmsLllQSt known Tertiary species of Perognathus \ P4
three- or four-cusped and much smaller relative to Mj than in other
species.

Description. — The mandible of P. trojectioansrum is slender and does not differ from
other species of Perognathus. The lower incisor is narrow with a convex anterior sur-
face. The upper incisor is unknown.

1979

Korth — Valentine Geomyoid Rodents

291

Table 2. — Measurements of the dentition of Perognathus trojectioansrum.

UNSM

no.

p4

M‘

M*

M3

alveo-
■ lar
length

A-P

tra

trp

A-P

tra

trp

A-P

tra

trp

A-P

tra

trp

56302

0.93

0.53

1.06

0.72

0.99

1.02

56303

0.94

0.50

1.04

56310

—

—

0.76

0.95

0.87

56313

1.03

0.55

0.99

0.75

1.00

1.01

UNSM

no.

P4

M,

M2

M3

alveo-

lar

length

A-P

tra

trp

A-P

tra

trp

A-P

tra

trp

A-P

tra

trp

56304

0.64

0.47

0.58

0.71

0.81

0.76

0.65

0.81

0.75

—

_

—

3.04

56305

0.66

0.56

0.75

56306

—

—

—

0.83

0.87

0.84

0.68

0.84

0.81

, —

—

2.49

56307

0.63

0.47

0.58

56308

0.55

0.47

0.47

56309

0.57

0.47

0.61

0.73

0.79

0.82

56311

0.55

0.44

0.58

0.72

0.75

0.79

0.57

0.80

0.80

0.48

0.64

0.44

2.37

56312

0.67

0.39

0.62

2.86

UNSM

no.

I

1

A-P

width

56304

0.69

0.41

56307

0.77

0.38

Cheek teeth are low crowned and cuspate when unworn or little worn. The teeth
become lophate after moderate wear. P4 is quite variable in pattern. In the holotype,
the two cusps in the hypolophid (hypoconid and entoconid) are of equal size and widely
separated. The metaconid and protostylid of the metalophid are subequal, transversely
aligned and separated almost as much as the cusps of the hypolophid. A minute hypo-
stylid is present on two specimens.

Most of the variation of P4 is on the metalophid. The protostylid on P4 of the two
specimens is reduced and slightly posterior to the metaconid. P4 in UNSM 56307 is
quadricuspate but differs from all other specimens in having an elevated metalophid.
Two more specimens have single-cusped metalophids with the cusp centrally located on
the lophid. In one of these, the three-cusped P4 is roughly circular in outline with the
three cusps placed peripherally on the tooth surrounding a central basin.

The size of P4 is much reduced relative to Mj and Mg.

Ml and M2 are six-cusped and do not vary in morphology from any other species of
Perognathus. The only known specimen of M3 of P. trojectioansrum is moderately
worn, so no details of cusp morphology can be accurately determined. The hypolophid
appears bicuspate, with the hypostylid absent.

The upper dentition of P. trojectioansrum is represented only by and Mb These
teeth are also similar to all other species of the genus. P^ has the basic four-cusped
heteromyid pattern. M^ consists of two transverse rows of three cusps divided by a
central valley. Alveoli show that M® is much reduced in size.

Discussion.— Perognathus trojectioansrum is the smallest recorded
species of Perognathus. James (1963) described P. minutus from the
Cuyama Valley of California and stated that it was smaller than any

292

Annals of Carnegie Museum

VOL. 48

Fig. 3. — Perognathus trojectioansrum. A) UNSM 56313, RP‘‘-M*. B) UNSM 56311,
occlusal view, P4-M3. C) Lateral view of same. Scale = 1 mm.

1979

Korth— -Valentine Geomyoid Rodents

293

fossil or Recent species of the genus. In all dimensions of the lower
dentition, P. trojectioansrum is smaller than any recorded specimen
of P. minutus (Lindsay, 1972). The upper dentition of P. trojectioans-
rum, however, is within the observed size range for P. minutus, though
generally in the lower part of the range. It is possible that the upper
teeth referred to P. trojectioansrum may be P. minutus, and only the
lower dentitions are of the former. However, it seems highly improb-
able that one species would be represented by only upper and the other
only lower teeth. There is no indication that either the lower or the
upper dentitions assigned to P. trojectioansrum represent more than
a single species.

The pattern of the molars in Perognathus trojectioansrum does not
differ from that of any other species of the genus. The only tooth
showing some variation from the basic Perognathus pattern is P4,
although the nature and form of the variation is not unique to heter-
omyid species.

Galbreath (1953) described a large sample of Heliscomys from the

Oligocene of Colorado which showed much of the same variation in
number and pattern of cusps on P4 as in P. trojectioansrum. Similar
ranges of variation on this tooth have also been observed in Recent
populations of Perognathus (Sutton, manuscript).

In proportions of teeth, P. trojectioansrum is quite similar to He-
liscomys. Both forms are characterized by small size and P4 being
disproportionately small with variable morphology.

Cupidinimus Wood, 1935

Perognathoides Wood, 1935
Peridiomys (in part) Wood, 1935 A.

Cupidinomys nomen nudum Shotwell and Russell, 1963.

Prodipodomysl (in part) Shotwell, 1967.

Perognathus (in part) Storer, 1970.

Emended diagnosis .—GcnQY^Wy small heteromyids; mesodont and
lophate rooted teeth; and six-cusped; P^ with variable occur-
rences of accessory cuspules on the protoloph within species; Ii slen-
der and rounded anteriorly; upper incisor asulcate; P4 with central
union of lophids and variable occurrence of minor cusps on the meta-
lophid and hypolophid; cingula distinct on molars.

Pu/tge.— Barstovian and Clarendonian (Miocene) of western North
America.

Type species .—Cupidinimus nebraskensis Wood, 1935:118.

Cupidinimus nebraskensis Wood, 1935
(Figs. 4, 5, 6 and Table 3)

Emended diagnosis .—-Small size; P4 with variable occurrence of hy-
poconulid, anteroconid and protoconid; P^ with accessory cuspule on

Fig. 4. — -Cupidinimus nebraskensis, stereoview, CM 10193 holotype, LP4-M3. x5.25.
Fig. 5 .—-Cupidinimus nebraskensis, stereoview, UNSM 56127, RP4-M1. x5.25.

Fig. 6. — Cupidinimus nebraskensis, stereoview, UNSM 56269, LP'*. x5.25.

Fig. 7. — Perognathus trojectioansrum, stereoview, UNSM 56310, RM^ x5.25.

protoloph in some specimens; M2^ and reduced relative to M/;
teeth mesodont; no hypostylid on P4

Referred material. —UNSM 56100, 56103, 56105-56113, 56115-56136, 56143-56227,
56299, 56357, 56360 (all lower teeth and mandible fragments), and UNSM 56101, 56102,
56104, 56114, 56137-56142, 56228-56298, 56361 (upper teeth and whole or partial max-
illae).

Description. — Wood (1935) adequately described the molars and mandible of C. ne-
braskensis, based on skeletal material and dentitions consisting of one nearly complete

1979

Korth — Valentine Geomyoid Rodents

295

Table 3. — Measurements (alveolar length) of the dentition of Cupidinimus nebraskensis.

Teeth

Measure-

ments

N

M

SD

cv

OR

Paratype

(CM

10170)

Holotype

(CM

10193)

Maxillary toothrow

9

3.93

0.24

6.11

3.53-4.21

3.83

P

A^P

1

0.92

—

width

1

0.60

—

—

—

—

A-P

1

0.88

—

—

—

dP^

tra

1

0.77

—

___

trp

1

0.95

—

—

—

—

A-P

61

1.43

0.13

9.10

1.14-1.70

1.28

p4

tra

61

0.70

0.07

10.00

0.55-0.85

0.68

trp

61

1.23

0.12

9.76

0.96-1.41

1.17

A-P

28

0.85

0.07

8.25

0.74-1.03

0.76

Ml

tra

28

1.15

0.11

9.56

0.96-1.50

1.05

trp

28

1.07

0.07

6.54

0.92-1.21

1.02

A~P

4

0.66

0.04

6.06

0.60-0.71

0.69

M2

tra

4

0.96

0.07

7.29

0.92-1.06

0.96

trp

4

0.84

0.07

8.33

0.78-0.94

0.85

A-P

2

0.50

—

—

0.47-0.52

0.55

M^'

tra

2

0.64

—

—

0.62-0.65

0.69

trp

2

0.53

—

—

0.49-0.57

0.67

Mandibular toothrow

31

3.67

0.06

4.58

3.41-4.08

3.59

A-P

28

0.89

0.06

7.28

0.78-1.03

0.88

width

28

0.57

0.08

13.20

0.48-0.76

0.47

A-P

3

1.18

—

— ,

1.11-1.29

dP4

tra

3

0.69

—

—

0.60-0.77

—

trp

3

0.80

—

—

0.76-0.87

—

A-P

88

1.00

0.08

8.07

0.78-1.18

0.95

P4

tra

88

0.72

0.07

9.59

0.59-0.91

0.74

trp

88

0.90

0.08

8.56

0.75-1.11

0.90

A-P

74

0.89

0.07

7.86

0.76-1.07

0.82

M,

tra

74

1.06

0.09

8.39

0.88-1.27

1.00

trp

74

1.05

0.08

7.70

0.90-1.26

0.95

A-P

14

0.76

0.06

8.33

0.66-0.85

0.67

M2

tra

14

1.01

0.09

8.86

0.88-1.21

0.95

trp

14

0.95

0.07

7.62

0.85-1.11

0.89

A-P

2

0.66

0.65-0.67

0.70

M3

tra

2

0.81

—

__

0.70-0.92

0.70

trp

2

0.71

—

0.66-0.76

0.60

mandible with all teeth and a maxilla with two P'^s, and two partial mandibles

with dP4 (and Mj in one instance). The UNSM material referred to this species allows
for a more complete description and a better understanding of the variability in mor-
phology of the premolars.

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VOL. 48

According to Wood (1935), P4 of C. nebraskensis has a simple four-cusped pattern,
with the hypolophid wider than the metalophid. The holotype (CM 10193), which con-
tained the only known P4, does have a minor anteroconid in the center of the metalophid.
Due to the state of wear, this cuspule is preserved as an anterior bend in the enamel
surrounding the anterior wear facet of the tooth. This anterior extension gives the
metalophid a trilobed appearance with the central lobe being the smallest.

In the UNSM sample there is a great deal of variation in the morphology of P4. On
approximately 80% of the specimens a minute anteroconid occurs on the metalophid.
The anteroconid is usually central, but can appear on the internal slope of either the
metaconid or protostylid.

A protoconid is present on about half of the specimens. It is smaller than either of the
other two metalophid cusps and slightly posterior to them. The posterior slope of the
protoconid extends posteriorly, in many cases, to form a minor ridge which runs antero-
posteriorly. This minor ridge is similar to that reported on P4 of ""Perognathus” sas~
katchewanensis (Storer, 1970, 1975).

The hypolophid, as noted by Wood (1935), is broad and composed of two major widely
spaced cusps. Some specimens have a central hypoconulid. It is generally a swelling
between the major cusps and varies in size from minute to subequal with the other
hypolophid cusps.

Presence of any or all of these three additional cusps (anteroconid, protoconid, hy-
poconulid) on a P4 is variable and appears totally random.

There is no hypostylid on the metaloph of P4 of C. nebraskensis. On three of the 88
specimens, there is a minute anterior projection from the hypoconid that may be ho-
mologous to a hypostylid, but does not develop into a cusp.

The protoloph of P^ has an accessory cuspule in 15% of the sample. This cuspule can
occur either buccal or lingual to the protocone and. either at the base or near the tip of
the latter.

The posterior portion of the mandible of the holotype is damaged and little can be
said of the morphology of the bone in this area. Wood (1935:128) reconstructs the
probable shape of the coronoid, condyle, and angle of the mandible on the holotype.
Specimens in the present sample show that the mandibular condyle is perhaps a bit
higher than in Wood’s figure, but otherwise the restoration is quite accurate. The angle
of the mandible is inflected slightly. The medial foramen on the ascending ramus is about
halfway between M3 and the tip of the condyle. The coronoid process is short and
slender, and deflects externally away from the ramus.

The bulbous lateral expansion on the ascending ramus marking the pulp cavity of the
incisor is small and nearly circular. It rises only slightly above the wall of the ramus.

Discussion. — In Wood’s (1935) review of the Heteromyidae two new
genera, Perognathoides and Cupidinimus were described. Both genera
were described as having asulcate upper incisors, progressively hyp-
sodont rooted molars, and perognathine tooth pattern (P4 with union
of lophs central and H-pattern of lophid union on lower molars). Cu~
pidinimus was described as having progressive deciduous premolars,
calcaneal-navicular articulation in the tarsus, and a subricochetal mode
of locomotion. None of these traits was referred to in his description
of Perognathoides. Perognathoides was characterized as having a lin-
gual and buccal cuspule on the protoloph of P^, a character considered
unique to this genus.

Wood (1935) failed to compare these two forms directly, although

1979

Korth — Valentine Geomyoid Rodents

297

the great similarity of these two genera was recognized by a number
of later authors (Wilson, 1939; Downs, 1956; Lindsay, 1972). The cri-
teria that were used to separate these two genera appear inadequate

to warrant generic separation.

The premolars of Cupidinimus nebraskensis are more complex than
initially recognized by Wood (1935). The morphology of P/ is variable
and consists of characters that have been associated with other genera
of heteromyids (notably Perognathoides, Wood, 1935).

The variable presence of a protoconid on P4 , small size and relative
reduction of M2^ and Mg^ make C. nebraskensis distinct from similar
mesodont species of heteromyids. Lindsay (1972) referred a number
of specimens from the Barstow Formation of California to C. nebras-
kensis. However, the Barstow material lacks any of the accessory
cusps on P/, is smaller in size, possesses a hypostylid on P4 (not found
in the Nebraska sample), and does not have reduced or Mg^. This
California species is clearly not referable to C. nebraskensis, and
should be designated as a new species of Cupidinimus.

Although Wood (1935) diagnosed the genus as having progressively
hypsodont teeth, C. nebraskensis was defined as being low crowned.
Teeth of C. nebraskensis are evidently mesodont in height and equal
to or nearly equal to that of species of Perognathoides.

Wood (1935) inferred the mode of locomotion of Cupidinimus from
the proportions of limb bones and the structure of the tarsus. At the
time, no postcranials were known for Perognathoides. It was also
impossible to compare the deciduous premolars of Cupidinimus to
Perognathoides, for which the deciduous premolars were unknown.
Hence, the only character available for comparison at the time of de-
scription of the two genera was the presence of accessory cuspules on
the protoloph of P^. Although this trait was cited as diagnostic for
Perognathoides, there was evidence of accessory cuspules on only
one of the two species assigned to the genus. The only known P^ of
P. tertius (later referred to P. halli. Wood, 1936^) is well worn and
shows no evidence of a distinct cuspule on either side of the protocone.
Wood (1935) inferred the presence of two accessory cuspules from the
slight transverse elongation of the protoloph. Later, Wood (1937) de-
scribed a species of Perognathoides, P. cuyamensis, which possessed
only a single cuspule on the protoloph of P^. James (1963) found this
to be a variable trait on Perognathus and questioned its diagnostic
value for Perognathoides.

Lindsay (1972), studying the first extensive collection of Perogna-
thoides, found that accessory cuspules on P^ occurred on half of the
sample of 50 specimens. The variability of this character in Cupidini-
mus nebraskensis, Perognathoides halli (Lindsay, 1972), Perognathus

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Annals of Carnegie Museum

VOL. 48

furlongi (James, 1963), and "'Diprionomys'' agrarius (see below) in-
dicates that this character does not separate Cupidinimus from Pe-
rognathoides.

Wood (1935) thoroughly described the postcranial skeleton of C.
nebraskensis. It most closely resembles that of Perognathus with a
number of characters tending toward specializations for ricochetal lo-
comotion seen in Recent Dipodomys. The only postcranial elements
reported for Perognathoides are calcanea from California assigned to
P. cuyamensis (Wood, 1937). They are described as being more slen-
der than the calcaneum of Perognathus, a character cited for Recent
ricochetal forms but not specifically noted for the calcaneum of Cup-
idinimus (Wood, 1935).

No calcaneum or cuboid is present among the postcranial bones of
the holotype of C. nebraskensis (CM 10193) though measurements for
one are listed in Wood (1935, table III). In his reconstruction of the
pes of C. nebraskensis, Wood (1935:139) figures a calcaneum restored
from another specimen, not in the Carnegie Museum of Natural His-
tory collections. The navicular on the holotype is badly abraded with
cancellous bone exposed on one surface. No articular surfaces are
evident on the lateral side of the navicular making it impossible to
determine the nature of the calcaneal contact with other tarsal bones.
Korth (1979) assigned several isolated calcanea and astragali to C.
nebraskensis {=Cupidinimus sp. B). When these elements are articu-
lated, the distal margin of the calcaneum does not extend beyond that
of the astragalus. One calcaneum can be articulated with the tarsal
elements of the holotype, and does not extend distal to the astragalus,
or articulate with the navicular.

Wood (1935) used presence of a calcaneum-navicular contact to di-
agnose Cupidinimus and as the basis for assigning it to the Dipodo-
myinae (although it was also cited as occurring in Paramys). Because
this character cannot now be verified, it seems advisable to eliminate
it from the diagnosis of the genus.

Thus, based on dental morphologies, Cupidinimus is inseparable
from Perognathoides. The synonymy of these genera poses a minor
nomenclatural problem because both were named in the same paper.
More species have been assigned to Perognathoides than Cupidini-
mus. However, the genoholotype of Cupidinimus (C. nebraskensis) is
an excellent specimen consisting of a partial skull, nearly complete
mandible with all teeth present and little worn, and a great deal of the
postcranial skeleton. The genoholotype of Perognathoides {P. quar-
tus) is a partial skull with heavily worn incomplete dentition, and a
single referred specimen, a mandible, also has badly worn dentition.
For comparison, it is much more advantageous to refer these species

1979

Korth — -Valentine Geomyoid Rodents

299

to Cupidinimus rather than Perognathoides. An extensive collection
of C, nebraskensis is available for comparison, whereas P. quart us is
known from only two specimens. Thus, Cupidinimus seems a better
choice for the generic assignment, A formal revision is presently im-
practical because many species have been diagnosed on poorly pre-
served material and known from only limited hypodigms. Species that
are well represented show a great deal of variability and render the
original diagnoses inapplicable. Due to this wide variability and limited
number of specimens, systematic revisions are not dealt with specifi-
cally here, pending a formal revision of the species of the genus.

Because the geeoholotype of Perognathoides is in such poor con-
dition, it could be argued that it is generically indeterminate. Thus, the
referral of all species assigned to Perognathoides with known occlusal
pattern of the teeth to Cupidinimus would not mean the synonymy of
these two genera. However, in all observable characters P. quartus
is congeneric with Cupidinimus .

All species previously assigned to Perognathoides should be re-
ferred to Cupidinimus {P. quartus [Hall, 1930]; P. tertius [Hall, 1930];
P. halli Wood, 1936^; P. cuyamensis Wood, 1937; P. cf. tertius Wil-
son, 1939; P. madisonensis Dorr, 1956; P. continentalis, nomen nu-
dum Reeder, 1958; P, cf. P. cuyamensis Klingener, 1968; P. Mein-
felderi Storer, 1970; P, eurekensis Lindsay, 1972). A thorough study
of these species may indicate that specific systematic revisions may be
necessary. Lindsay (1972) suggested some synonymies but only dealt
with a few species of Perognathoides and not the entire genus along
with the species of Cupidinimus.

In addition to the species of Perognathoides, a number of other
forms are probably also referable to Cupidinimus . Storer (1970) de-
scribed Perognathus saskatchewanensis from the Wood Mountain
Formation of Saskatchewan. This species was diagnosed by posses-
sion of an anterocoeid on P4 and presence of a central longitudinal
ridge on the same tooth, features considered unique to species of Pe-
rognathus. Storer failed to discuss crown height of this species, or to
provide figures depicting this dimension (also see Storer, 1975). When
specimens of '"Perognathus'' saskatchewanensis are compared with
other species of Perognathus and Cupidinimus, is is apparent that
"P." saskatchewanensis is mesodont as is Cupidinimus, much higher
crowned than Perognathus. The occurrence of an anteroconid on most
specimens of P4 of "P." saskatchewanensis agrees with C. nebras-
kensis and other species referred here to the latter genus (Lindsay,
1972; Sutton, manuscript).

Cupidinumus sp. from the Mission local fauna (Clarendonian) of
South Dakota '(Green, 1971), Prodipodomys? mascallensis from the

300

Annals of Carnegie Museum

VOL. 48

Quartz Basin (Barstovian) of Oregon (Shotwell, 1967), and Cupidino-
mys sp. from the Juntura Basin (Clarendonian) of Oregon are probably
also referable to Cupidinimus.

The distinction between some of the species referred here to Cupi-
dinimus is quite subtle and often is based only on mean size of an
extensive sample as in C. halli and C. madisonensis (Sutton, manu-
script). These discrete differences are observed only when large sam-
ples of the species are known. Because of this, difficulties arise in the
allocation of specimens to any species of Cupidinimus unless an ad-
equate sample of fossils is known. More extensive collections of
known species discussed above may well suggest systematic revisions
of the species of Cupidinimus.

^'Diprionomys^^ agrarius Wood, 1935
(Figs. 8, 9 and Table 4)

Referred material.— VNSM 56353-56356 (P'‘s), UNSM 56349 (RP4), UNSM 56350
(LM, or M2), UNSM 56351 (LM3), UNSM 56352 (edentulous mandible), UNSM 56358
(LdP'*) and UNSM 56359 {hdFd-

Description. — A left edentulous mandible is referable to this species based on the
comparable size of known teeth and the presence of alveolae for roots on all teeth. The
mandible is broken away anterior to the alveolus for P4. The posterior part of the
mandible is nearly complete. The ramus is quite deep and virutally identical to that of
""Diprionomys" agrarius (Wood, 1935:180, i84).

All cheek teeth are low crowned. The crown height is perhaps slightly higher than in
Perognathus, but clearly lower than in mesodont heteromyids like Cupidinimus. A
single P4 is present in the collection. It is basically four-cusped. The hypolophid has a
distinct hypoconid and entoconid with no other cuspules. The metaconid shows mod-
erate wear and is reduced to a large circular facet, which extends labially and posteriorly
to another nearly circular facet in the center of the tooth. Posteriorly, this facet unites
with the hypoconid by means of a narrow connection. A smaller protostylid is distinct
from the metaconid facet and is unworn.

The lower molars referred to this species differ from the holotype of “Z).” agrarius
only in their slightly larger size.

P”* has a simple pattern. The metaloph consists of three distinct cusps, the centrally
placed hypocone being the largest. The metacone and entostyle are displaced slightly
anterior to the hypocone giving the metaloph a posteriorly convex appearance. The
protoloph contains a single protocone. On one specimen (UNSM 56356) there is a small
cuspule just labial to the apex of the protocone.

DP^ is lower crowned than P^ and slightly more transverse anteriorly. It is composed
of two main transverse lophs, each with two cusps. A third, anterior loph composed of
a crescentic anterocone is connected to the posterior transverse loph via a strong lingual
cingulum. The transverse valley between the main lophs is shallow. The anterocone is
united with the central transverse loph labially.

Discussion.— In the morphology of the molars and mandible, the
UNSM specimens are nearly identical with “D.” agrarius, known
from the Cap Rock Member of the Ash Hollow Formation in Nebraska
(Wood, 1935) and the Norden Bridge local fauna (Klingener, 1968) of
the Valentine Formation. Morphologically, P4 from locality Kx 110 is

1979

Korth — Valentine Geomyoid Rodents

301

Fig. %.— "Diprionomys" agrarius, stereoview, UNSM 56356, LP'*, x5.25.

Fig. 9.- — '‘"‘Diprionomys” agrarius, stereoview, UNSM 56349, RP4. x5.25.

quite different from P4 in the holotype of ‘‘D.” agrarius, which has a
distinct anteroconid, and the metaconid and protostylid separated by
a deep longitudinal valley. The only P4 assigned here to ''Dipriono-
mys'' agrarius shows a remnant of what may have been a protoconid
displaced posterior to the other metalophid cusps. The union of the
lophs is, however, not central and a minor central depression is pres-
ent.

Klingener (1968) described five P4S of “D.” agrarius from the Nor-
den Bridge local fauna. He noted the differing degree of complexity of
the metalophid of P4 and established a size range for this tooth that is
large enough to include the Kx 110 material.

302

Annals of Carnegie Museum

VOL. 48

Table 4.-

—Measurements of the dentition of

“Diprionomys”

agrarius.

Teeth

UNSM no.

A-P

tra

trp

dp4

56358

1.66

1.19

1.59

p4

56353

1.75

0.85

1.62

56354

—

0.90

1.78

56355

1.63

0.90

1.83

56356

1.84

0.96

1.87

dP4

56359

2.04

0.97

1.35

P4

56349

Ml or M2

1.44

1.12

1.36

56350

1.51

1.82

1.77

M3

56315

1.35

1.53

1.50

The two lower molars assigned to “D.” agrarius are slightly larger
than those of the holotype. The differences in size, however, are no i
greater than those recorded by Klingener (1968) for the premolar. The |
hypostylid on Mg of the UNSM material is slightly larger than that of
the holotype but not significantly so. No other differences in mor~ '
phology exist in the molars.

The variability in the metalophid of P4 is also reflected by the oc- '
casional occurrence of an accessory cuspule near the apex of the pro- ,
tocone on the protoloph of a character also noted in D. cf. parvus \
from Nevada (Clark et al., 1964). (

^^Diprionomys"' agrarius was initially described by Wood (1935) and ,
assigned to the same genus Diprionomys parvus (Kellogg, 1910) from |
the Clarendonian of Nevada. “D.” agrarius differs most markedly j
from D. parvus in size and crown height. The molars of D. parvus are I
quite high crowned as implied by the straight lateral walls of the teeth.
“D.” agrarius, however, is very low crowned. The crowns of the
latter are quite similar in height to those of Peridiomys. 1

Because of the difference in crown height between “D.” agrarius |'
and D. parvus, Reeder (1958) included “D.” agrarius in a new genus !j
{Halticosomys, nomen nudum). Two isolated teeth from the Fish Lake I
Valley fauna of Nevada were identified as D. cf. parvus, based on
size. From the figures (Clark et al., 1964), these specimens appear I
lower crowned than the holotype of D. parvus (Kellogg, 1910) and the |

1979

Korth — Valentine Geomyoid Rodents

303

possesses an accessory cuspule as in “D.” agrarius. “D.” cf.

parvus, therefore, may represent a distinct species assignable to a new
genus which would also include “D.” agrarius. At present, none of
these species {D. parvus, ‘"D.” agrarius, “D.” cf. parvus) are well
known. Recognition of a new genus separate from Diprionomys to
include “D.” agrarius and cf. parvus should await a formal

review of the entire Heteromyidae. For convenience, “D.” agrarius
is retained in the genus with question until it is better known and a
more detailed study is made.

Family Geomyidae
Subfamily uncertain
Lignimus cf. L. hibbardi Storer, 1973
(Figs. 10, 11, 12 and Table 5)

Referred material.— VmU 56316-56326 (P4S), UNSM 56327-56337 (P^s), UNSM
56339, 56340, 56346 (lower molars), UNSM 56338, 56341-56345, 56347, 56348 (upper
molars) and UNSM 56315 (an edentulous mandible).

Description. — A left mandible lacking cheek teeth is here assigned to Lignimus. This
allocation seems certain because the size is comparable with the isolated teeth and there
is no indicattion of sockets for any roots in the molar alveoli.

The angle and a portion of the lateral wall of the mandible is broken away. The
diastema is shallow and relatively short. The mental foramen is situated below the center
of the diastema and halfway between the ventral and dorsal margins of the bone. The
ridge marking the attachment of the masseter muscle terminates just dorsal and posterior
to the mental foramen. It runs posteriorly, directed toward the ventral margin of the
jaw at the level of Mg. The posterior extent of this ridge is uncertain because the bone
is broken in this area.

The alveolar margin forms a straight anteroposterior ridge. The ascending ramus
blocks only Mg in lateral view. It is broad and rises well above the molars. The condyle
is about as high as the coronoid, which is relatively low and blunt. There is a distinct
basin area just below the condyle that extends anteriorly to the coronoid process and
ventrally to the lateral expansion of the jaw marking the pulp cavity of the incisor.
Medially, there is a depression lateral to Mg extending onto the ascending ramus to just
below the coronoid. The medial foramen of the ascending ramus is nearly midway
between Mg and the condyle, but slightly closer to the latter.

The enamel of the lower incisor is thin. The anterior margin of this tooth is flattened
and broad. In cross section, the tooth gradually tapers posteriorly.

The cheek teeth are hypsodont. All premolars and a single specimen of M^ are rooted.
All other molars lack roots. The lower premolars are vitaully identical with the holotype
of L. hibbardi, possessing one or two anterostylids, with lateral and lingual dentine
tracts extending to 14 or Vs the height of the crown from the base of the enamel.

The lower molars are recurved and rootless. Dentine tracts extend dorsally to about
half the height of the crown. The occlusal pattern is that described for L. hibbardi
(Storer, 1973). THe cusps rapidly wear into two straight transverse lophs that eventually
unite centrally in an H-Pattern. The protoconids are not as distinctly isolated as in L.
montis. P^ has four cusps. The protoloph consists of the protocone that, in unworn
specimens, is ovoid. The protocone wears to a shortened transverse loph. The metaloph
of P‘‘ is simple. The entostyle, smallest of the three cusps, connects to the hypocone by
a cingulum that curves forward from the latter. The entostyle is positioned anteriorly
about halfway between the apices of the hypocone and protocone. The metaloph wears
to a J-pattern, with the entostyle as the hook of the J.

304

Annals of Carnegie Museum

VOL. 48

Table 5— Measurements of the dentition o/ Lignimus cf L. hibbardi.

Teeth

UNSM no

A-P

tra

trp

p4

56327

1.58

1.06

1.88

56328

1.75

1.08

1.93

56329

1.71

1.17

1.88

56330

1.64

1.00

1,79

56331

1.46

0.86

1.73

56332

1.71

1.20

1.83

56333

1.55

1.10

1.86

56334

1.70

1.27

1.91

56335

1.72

1.12

1.87

56336

1.56

1.06

1.73

56337

1.47

0.92

1.78

M'

56338

1.16

1.87

1.72

56344

1.01

1.68

1.56

56345

1.18

1.71

1.59

56341

1.08

1.52

1.47

56343

1.06

1.52

1.37

56348

—

1.55

1.46

56342

1.10

1.41

1.25

It

56347

0.93

1.47

1.43

56315

1.42

1.18 (width)

P4

56316

1.81

1.13

1.59

56317

1.85

1.34

1.54

56318

1.81

1.35

1.62

56319

1.82

1.35

1.66

56320

1.92

1.47

1.72

56321

1.89

1.30

1.64

56322

1.89

1.32

1.48

56323

2.04

1.27

1.58

56324

1.98

1.52

1.66

56325

1.90

1.36

1.66

56326

1.60

1.18

1.61

M,

56346

1.19

1.69

1.58

M2

56340

1.11

1.58

1.51

M3

56339

1.06

1.39

1.39

Korth— Valentine Geomyoid Rodents

305

1979

Fig. 10.— Occlusal and lateral view of lower cheek teeth of Lignimus cf. L. hibbardi.

A) UNSM 56339, RMg. B) UNSM 56340, LM^. C) UNSM 56346, RM^. D) UNSM 56322,

RP4. Scale = 1 mm.

The upper molars are also recurved. The extent of the lateral dentine tracts is about
half the height of the crown, and perhaps slightly more on M^. All upper molars share
the same occlusal pattern as in L. hibbardi, and decrease in size from to M^. The
molars have two distinct transverse lophs. The lophs are joined lingually by an internal
cingulum. There is evidence of roots on only one specimen of where they are short
stubs equal to less than 10% of the total height of the tooth. has three roots. The
anterior root is strong and the posterior roots are weak.

Discus sion.—SiOTQV (1973) identified a number of specimens from
the Norden Bridge local fauna and the Crookston Bridge Member of
the Valentine Formation (Cherry County, Nebraska) as Lignimus sp.
He noted similarities between these specimens and L. hibbardi from
Kansas. The only difference cited between Lignimus sp. and L. hib-
bardi was the relative extent of the dentine tracts on the molars, which
reached about % of the height of the crown in L. hibbardi and only Vi
the height in Lignimus sp.

Both of these species differ from L. mantis, the type species, in
occlusal pattern of the molars and premolars, the presence of dentine
tracts on the lateral walls of the cheek teeth, and lack of roots on the
molars. The lower molars of L. hibbardi and the Nebraska species do
not form the distinct X-pattern of L. mantis. In the former, the rows

306

Annals of Carnegie Museum

VOL. 48

uig. 11. — Medial and occlusal views of upper cheek teeth of Lignimus cf. L. hibbardi.
A) UNSM 56329, RP^ B) UNSM 56345, RMh C) UNSM 56343, RM^ D) UNSM 56342,
LM^. Scale = 1 mm.

of cusps are more nearly parallel and wear as two separate transverse
lophs, finally uniting centrally in a pattern more similar to the H-pat-
tern of heteromyids (see Wood, 1935). Also, in L. hibbardi and the
Nebraska species, the anterior cingulum is more closely appressed to
the protoconid than in L. montis and does not form the anterobuccal
enamel lake when worn as in the latter.

The lower premolars of the Kansas and Nebraska species are char-
acterized by a distinct central trigonid basin surrounded by multiple
cusps. The major cusps (protostylid and metaconid) are widely sepa-
rated and connected to an anteriorly curving loph composed of one or
two smaller cuspules. L. montis has no such basin. The metalophid
consists of two major cusps that are only separated by a narrow valley,
and one or two additional smaller cuspules, not arranged in the same
way as in L. hibbardi.

of L. montis has a large circular protocone and an additional
anterocone in some specimens. No known specimens of of Lignimus
from Kansas or Nebraska have an anterocone. The protoloph on P^ of
the latter is flattened anteriorly and is a transverse loph when worn.
On unworn specimens this cusp is simple and ovoid in shape.

In L. montis all teeth are rooted and there is no interruption in

1979

Korth — Valentine Geomyoid Rodents

307

, j

Fig. 12. — Lateral view of left mandible of Lignimus cf. L. hibbardi, UNSM 56315.
Scale = 5 mm.

enamel on any side of the tooth. L. hibbardi and Lignimus cf. L.
hibbardi exhibit some degree of enamel failure on the lateral and lin-
gual sides of the teeth, and the molars are rootless. The protocone on
the upper molars of L. montis is large and somewhat isolated (Storer,
1970). The upper molars of the Kansas and Nebraska species have
protocones which are subequal with the other three cusps of the teeth,
and the depression separating the protocone from the cingulum is quite
shallow and any trace of an enamel lake is rapidly eliminated.

The differences between the Kansas-Nebraska Lignimus and L.
montis may be sufficient to warrant generic separation. However,
these species are all poorly represented, known from isolated teeth
only. Establishment of a new genus, if necessary, for the Kansas-Ne-
braska material should await recovery of more extensive collections
and more complete material of these taxa.

Storer (1970, 1973, 1975) referred Lignimus to the Entoptychinae,
based on the similarities of the cheek teeth with Gregorymys. Many
of the similiarities between L. montis and Gregorymys, however, are
not present in the Kansas and Nebraska species. This indicates that
either the species referred to Lignimus represent more than one genus
or that the type species is unique and may be convergent with entop-
tychines.

The mandible here referred to Lignimus cf. L. hibbardi differs great-
ly from those of entoptychines (see Rensberger, 1971). It most closely
resembles mandibles of more primitive pleurolicines (see Rensberger,
1974), though this similarity may only be a reflection of the primitive
character of the mandible.

308

Annals of Carnegie Museum

VOL. 48

The cheek teeth of L. hibbardi and L. cf. L. hibbardi are quite
similar to Pliosaccomys (Wilson, 1936).

The Kansas and Nebraska forms of Lignimus are intermediate be-
tween Pliosaccomys and ParapUosaccomys in development of hyp-
sodonty and completeness of enamel failure on the molars, and have
a crown pattern similar to these forms. The Unique “X” pattern on
the molars of L. montis is also present on the molars of Dikkomys
(Wood, 1936/?; Black, 1961) and may indicate a close relationship of
these two forms.

Storer (1973) suggests that a specimen from the Clarendonian Ava-
watz fauna from California referred to “dipodomyine n. gen. and sp.”
by Wilson (1939) is actually Lignimus, However, from the figures
available, the outline of P4 of the California species is clearly distinct
from that of Lignimus, Also, the increased crown height and com- 1
pleteness of the enamel failure on the teeth of the Avawatz form are I
unequaled in any species of Lignimus.

Summary

The geomyoids from Annie’s Geese Cross locality are of little help
in solving the “Valentinian” age problem. Two of the heteromyids,
Cupidinimus nebraskensis and Perognathus trojectioansrum, are re-
stricted to the Crookston Bridge Member of the Valentine Formation.
""Diprionomys'' agrarius is known from both the Clarendonian Cap
Rock Member of the Ash Hollow Formation and the Barstovian
Crookston Bridge Member of the Valentine Formation in Nebraska
(Wood, 1935; Klingener, 1968) and from the Wood Mountain Forma-
tion (Barstovian) of Canada. Perognathus furlongi is known through-
out the Barstow Formation of California. Lignimus, outside of Ne-
braska, is known from the Barstovian Wood Mountain fauna (Storer,
1975) and Clarendonian Ogallala Formation of Kansas (Storer, 1973).

More than these five species must be considered before a conclusion i
can be reached as to the age of the lower Valentine Formation.

Acknowledgments

Specimens from the University of Nebraska State Museum were loaned by M. R.
Voorhies. Drs. M. Dawson, J. Sutton, and L. Krishtalka critically read parts of this
manuscript. The work was supported through the Rea Predoctoral Fellowship of the
Carnegie Museum of Natural History. ;

Literature Cited

Black, C. C. 1961. Rodents and lagomorphs from the Miocene Fort Logan and Deep
River Formations of Montana. Postilla, 48:1-20.

Clark, J. B., M. R. Dawson, and A. E. Wood. 1964. Fossil mammals from the
Lower Pliocene of Fish Lake Valley, Nevada. Bull. Mus. Comp. ZooL, 131:27-63.

1979

Korth— Valentine Geomyoid Rodents

309

Dorr, J. A. 1956. Anceney local mammal fauna, latest Miocene, Madison Valley For-
mation, Montana. J. PaleontoL, 30:62-74.

Downs, T. 1956. The Mascall fauna from the Miocene of Oregon. Univ. California
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Galbreath, E. C. 1953. A contribution to the Tertiary geology and paleontology of
northeastern Colorado. Univ. Kansas PaleontoL Contrib., Vert., 4:1-120.

Gazin, C. L, 1930. A Tertiary vertebrate fauna from the upper Cuyama drainage basin,
California. Carnegie Inst. Washington, Publ., 404:55-76.

Green, M, 1971. Additions to the Mission vertebrate fauna, Lower Pliocene of South
Dakota. J. PaleontoL, 45:486-490.

Hall, E. R. 1930. Rodents and lagomorphs from the later Tertiary of Fish Lake Valley,
Nevada. Univ. California Publ., GeoL Sci., 19:295-311.

James, G. T. 1963. Paleontology and nonmarine stratigraphy of the Cuyama Valley
badlands, California. Part 1. Geology, faunal interpretations, and systematic de-
scriptions of Chiroptera, Insectivora, and Rodentia. Univ. California Publ., Geol.
Sci., 45:1-154.

Kellogg, L. 1910. Rodent fauna of the late Tertiary beds at Virgin Valley and Thou-
sand Creek, Nevada. Univ. California Publ., Geol. Sci., 5:421-437.

Klingener, D. 1968. Rodents of the Mio-Pliocene Norden Bridge local fauna, Ne-
braska. Amer. Midland Nat., 80:65-74.

Korth, W. W. 1979. Taphonomy of microvertebrate fossil assemblages. Ann. Carnegie
Mus., in press.

Lindsay, E. H. 1972. Small mammal fossils from the Barstow Formation, California.

Univ. California Publ., Geol. Sci., 93:1-104.

Reeder, W. G. 1958. A review of Tertiary rodents of the family Heteromyidae. Diss.
Abstr., 18:1548.

Rensberger, j. M. 1971. Entoptychine pocket gophers (Mammalia, Geomyoidea) of
the early Miocene John Day Formation, Oregon. Univ. California Publ., Geol. Sci.,
90:1-209.

. 1974. Pleurolicine rodents (Geomyoidea) of the John Day Formation, Oregon,

and their relationships. to taxa from the early and middle Miocene, South Dakota.
Univ. California Publ., Geol. Sci., 102:1-95.

Schultz, C. B., M. R. Schultz, and L. D. Martin. 1970. A new tribe of saber-
toothed cats (Barbourofelini) from the Pliocene of North America. Bull. Univ. Ne-
braska State Mus., 9:1-31.

Shotwell, j. a. 1967. Late Tertiary geomyoid rodents of Oregon. Univ. Oregon Mus.
Nat. Hist. Bull., 9:1-51.

Shotwell, J. A., and D. E. Russell. 1963. Mammalian fauna of the Upper Juntura
Formation, the Black Butte local fauna. Pp. 42-76, in The Juntura Basin: studies
in earth history and paleoecology (J. A. Shotwell et al.), Trans. Amer. Phil. Soc.,

53:1-77.

Skinner, M. F., S. M. Skinner, and R. J. Gooris. 1968. Cenozoic rocks and faunas
of Turtle Butte, south-central South Dakota. Bull. Amer. Mus. Nat. Hist., 138:379-

436.

Storer, j. E. 1970. New rodents and lagomorphs from the Upper Miocene Wood
Mountain Formation of southern Saskatchewan. Canadian J. Earth Sci., 7:1125-

1129.

— . 1973. The entoptychine geomyid Lignimus (Mammalia: Rodentia) from Kansas

and Nebraska. Canadian J. Earth Sci., 10:72-83.

. 1975. Tertiary mammals of Saskatchewan. Part III. The Miocene fauna. Life

Sci. Contrib., Royal Ontario Mus., 103:1-134.

Sutton, J. F. 1977. Mammals of the Anceney local fauna (late Miocene) of Montana.
Unpublished Ph.D. dissert., University Microfilms, Ann Arbor.

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Annals of Carnegie Museum

VOL. 48

VooRHiES, M. R. 1971. Paleoclimatic significance of crocodilian remains from the
Ogallala Group (Upper Tertiary) in northeastern Nebraska. J. Paleontol., 45:119-121.

Webb, S. D. 1969. The Burge and Minnechaduza Clarendonian mammalian faunas of
north-central Nebraska. Univ. California Publ., Geol. Sci., 78:1-191.

Wilson, R. W. 1936. A Pliocene rodent fauna from Smiths Valley, Nevada. Carnegie
Inst. Washington Publ., 473:15-34.

-. 1939. Rodents and lagomorphs of the late Tertiary Avawatz fauna, California.

Carnegie Inst. Washington Publ., 514:31-38.

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Publ., 584:67-164.

Wood, A. E. 1935. Evolution and relationships of the heteromyid rodents with new
forms from the Tertiary of western North America. Ann. Carnegie Mus., 24:73-
262.

. 1936fl. Fossil heteromyid rodents in the collections of the University of Cali-
fornia. Amer. J. Sci., 32:112-119.

— . 19366. Geomyid rodents from the middle Tertiary. Amer. Mus. Novitates,

886:1-31.

-. 1937. Additional material from the Tertiary of the Cuyama Basin of California.

Amer. J. Sci., 33:29-43.

Wood, H. E., R. W. Chaney, J. Clark, E. H. Colbert, G. L. Jepsen, J. B. Reeside,
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■97 73

I (ISSN 0097-4463)

ANNALS'

<)/ CARNEGIE MUSEUM

: CARNEGIE MUSEUM OF NATURAL HISTORY

: 4400 FORBES AVENUE * PITTSBURGH, PENNSYLVANIA 15213

VOLUME 48 7 SEPTEMBER 1979 ARTICLE 17

i NOTES ON BATS (MAMMALIA: CHIROPTERA) FROM

BONAIRE AND CURASAO, DUTCH WEST INDIES

Hugh H. Genoways

Curator, Section of Mammals

' Stephen L. Williams

Collection Manager, Section of Mammals

Abstract

Five species of bats are reported from islands of Bonaire and Curasao, Dutch West
Indies. Natalus tumidirostris Miller is recorded from Bonaire for the first time. Signif-
icant differences are detected in three measurements for both sexes between populations
, of Glossophaga longirostris Miller from Curasao and Bonaire. Myotis nesopolus Miller
I is shown to be a distinct species and the senior synonym for the taxon M. larensis
I LaVal.

!

Introduction

i

' !

1

I

During the period 15 to 25 August 1977, a field party from the Car-
negie Museum of Natural History collected bats on the islands of Bon-
aire and Curasao in the Dutch West Indies. Five species are repre-
sented in the 435 specimens collected. Eight species of bats have been
reported previously from Curasao, and six have been taken on Bonaire
(Koopman, 1958; Husson, 1960; Smith and Genoways, 1974). We re-
port herein a seventh species, Natalus tumidirostris, from Bonaire.
Also discussed are an examination of variation in Glossophage lon-
girostris and the taxonomic relationships of the Myotis occurring on
the islands.

I This research and publication supported by the M. Graham Netting Research Fund
; through a grant from the Cordelia Scaife May Charitable Trust.

1 Submitted for publication 1 March 1979.

311

312

Annals of Carnegie Museum

VOL. 48

Methods and Materials

External and cranial measurements were taken by means of dial calipers. All mea-
surements are given in miOimeters. Measurements given are those of adults (phalangeal
epiphysis closed) except where noted, and were were made on preserved specimens.
Measurements were taken as follows: length of forearm, from the posteriormost pro-
jection of the elbow to the anteriormost portion of the wrist joint with the wing flexed;
length of third metacarpal, distance from the wrist to the distal end of the third meta-
carpal; length of tibia, length of lower leg from the knee to the ankle; greatest length
of skull, distance from the posteriormost projection of the cranium to the anteriormost
surface of the upper incisors; condylobasal length, distance from the posteriormost
projection of exoccipital condyles to the anteriormost projection of premaxillae; zygo-
matic breadth, greatest width across zygomatic arches at right angles to longitudinal
axis of cranium; mastoid breadth, greatest width across mastoid processes at right
angles to longitudinal axis of skull; postorbital breadth, least width across the postorbital
constriction at right angles to the longitudinal axis of the cranium; length of maxillary
toothrow, distance from posterior lip of alveolus of M^ to the anterior lip of alveolus of
canine; length of upper toothrow, distance from posterior lip of alveolus of M^ to the
anterior lip of alveolus of P; breadth across upper molars, greatest distance from labial j
margins of the upper molars at the widest point; postpalatal length, distance from the
posterior margin of the palate to the anteriormost portion of the foramen magnum; depth |
of braincase, distance from the line along the flat part of the braincase to a line on the
mid ventral part of the cranium touching the palate and the basioccipital.

Statistical procedures were performed on an IBM-360 computer at Camegie-Mellon
University and a DEC- 10 computer at the University of Pittsburgh. Univariate analyses
were performed using the program UNIVAR. This program yields standard statistics
(mean, range, standard deviations, standard error of the mean, variance, and coefficient
of variation), and employs a single-classification analysis of variance (F-test, significance
level 0.05) to test for significant differences between means (Sokal and Rohlf, 1969).

Stepwise discriminant analysis and canonical analysis (BMDP7M, Dixon and Brown,
1977) are techniqus that define and separate groups. The program performs a multiple
discriminant analysis in a stepwise manner, selecting the variable entered by finding the
variable with the greatest F value. The F value for inclusion was set at 0.01, and the F
value for deletion was was set at 0.05. Canonical coefficients were derived by multiplying
the coefficient of each discriminant function by the mean of each corresponding variable.
The program also classifies individuals, placing them with the group that they are nearest
to on the discriminant functions.

Most specimens examined are deposited in the Section of Mammals, Carnegie Mu-
seum of Natural History, and carry no institution designation in the lists of specimens
examined. Specimens from the American Museum of Natural History are marked
AMNH; those from the National Museum of Natural History, USNM.

Species Accounts

Mormoops megalophylla intermedia Miller

Specimen examined (1). — Bonaire: 8.5 km N, 2 km W Kralendijk, 1.

Our single specimen, an adult male with testes measuring 3 mm, was
taken on the night of 21 August 1977. The habitat in the area was
overgrazed pastureland with xeric mesquite^like vegetation. Also in
the area was a natural spring that had been modified with concrete
holding tanks. Nets were placed over the water where this individual
was taken, and among the trees in the pastures.

1979

Genoways and Williams — Bonaire and CuRAf ao Bats

313

This species has been reported previously from Bonaire (Husson,
1960; Smith, 1972). The length of forearm of our specimen (52.8) falls
between the values of two specimens (50.3, 53.4) reported by Smith
(1972) from Bonaire. We have followed Smith (1972) in the use of this
trinomial.

Glossophaga iongirostris elongata Miller

Specimens examined (404). — Bonaire: 8.5 km N, 2 km W Kralendijk, 147; 1.5 km
N, 6.5 km E Kralendijk, 52. Curasao: 2.8 km S, 4.5 km E Westpunt, 3; 6.8 km N, 15.6
km W Willemstad, 10; 4 km N, 3 km W Willemstad, 157; Willemstad, 6; 4 km S, 13.4
km E Willemstad, 29.

On Curasao, the large series of Glossophaga Iongirostris from 4 km
N, 3 km W Willemstad was taken in and around a small cave. The
habitat in the area was exceptionally dry with xeric forms of vegetation
(including cactus) scattered over a rock surface or shallow rocky soil.
Specimens from 4 km S, 13.4 km E Willemstad were taken at the
entrances of two small caves occurring in the caprock bordering the
coastline, whereas at 6.8 km N, 15.6 km W Willemstad bats were
netted around an abandoned plantation house. Near Westpunt, mist
nets were placed along the bed of a stream where the numerous large
manzanillo trees formed a canopy over the area. In Willemstad, bats
were taken in nets placed in a plant nursery possessing mesic types of
vegetation.

The situation at 8.5 km N, 2 km W Kralendijk on Bonaire is de-
scribed in detail in the account for Mormoops. At 7.5 km N, 6.5 km
E Kralendijk, Bonaire, bats were collected in caves opening in the
caprock along the coastline. The habitat in the area was dry, being
dominated by low, xeric^-adapted vegetation.

Represented in the specimens that we examined are 221 females and
183 males. Of the females taken between 16 and 25 August, 103 were
lactating (46.6%). One female taken on 19 August at 7.5 km N, 6.5 km
E Kralendijk was carrying a single embryo that measured 25 mm in
crown-rump length. Many of the specimens in our sample are nearly
I adult-sized young-of-the-year. Examination of a random sample of 100
! specimens from 4 km N, 3 km W Willemstad revealed the following
: age distribution: 15 adult males; 38 adult females (32 lactating); 17
subadult males (subadult pelage, but phalangeal epiphyses fused); 13
subadult females; eight young males (subadult pelage, phalangeal epiph-
yses unfused); nine young females. Average length of forearm in the
young males was 37.4 (36.8-38.1) N = 8; young females, 36.9 (36.3-
i 37.7) N = 10. All individuals were taken in mist nets, and were thus
capable of flight. Obviously, we sampled these populations just at the
end of a reproductive season; Wilson (1979) also reported this species
reproducing in August elsewhere in its geographic range. Mean length
of testes for 10 adult males was 3.6 (3-=4).

314

Annals of Carnegie Museum

VOL. 48

Table 1. — External and cranial measurements of male and female Glossophaga longi-
rostris eiongata from Bonaire and Curagao.

Locality

Sex

n

Mean

(Range) ± 2SE

cv

Length of forearm

Bonaire

(3

12

37.2

(36.1-38.1) ± 0.35

1.6

Curagao

d

18

37.2

(35.4-38.7) ± 0.38

2.1

Bonaire

9

14

37.7

(36.8-39.0) ± 0.34

1.7

Curagao

9

23

37.8

(36.7-38.9) ± 0.21

1.4

Greatest length of skull

Bonaire

(?

12

23.8

(23.1-24.6) ± 0.23

1.7

Curagao

d

17

23.5

(22.7-24.2) ± 0.21

1.8

Bonaire

9

13

23.7

(23.2-24.5) ± 0.20

1.5

Curagao

9

23

23.8

(23.3-24.5) ±0.11

1.1

Condylobasal length

Bonaire

d

12

22.2

(21.4-22.8) ± 0.24

1.9

Curagao

d

18

21.8

(20.8-22.5) ± 0.20

1.9

Bonaire

9

14

22.1

(21.7-22.6) ± 0.14

1.2

Curagao

9

23

22.0

(21.6-22.7) ± 0.12

1.3

Zygomatic breadth

Bonaire

d

11

9.3

(8.6-9.5) ± 0.16

2.9

Curagao

d

18

9.6

(9. 1-9.8) ± 0.09

1.9

Bonaire

9

10

9.2

(9.0-9.6) ±0.11

1.8

Curagao

9

16

9.6

(9.3-9.3) ± 0.08

1.6

Mastoid breadth

Bonaire

d

12

8.9

(8.3-9.2) ± 0.15

2.9

Curagao

d

18

9.1

(8.7-9.4) ± 0.10

2.2

Bonaire

9

14

8.9

(8.7-9.2) ± 0.09

2.0

Curagao

9

23

9.1

(8.9-9.4) ± 0.06

1.5

Postorbital breadth

Bonaire

d

12

4.6

(4.3-4.7) ± 0.08

3.0

Curagao

d

18

4.6

(4.4-4.8) ± 0.06

2.8

Bonaire

9

14

4.5

(4.2-4.9) ± 0.10

4.2

Curagao

9

23

4.6

(4.3-4.8) ± 0.06

2.9

Length of maxillary toothrow

Bonaire

d

12

8.0

(7.7-8.4) ± 0.12

2.5

Curagao

d

18

8.0

(7.5-8.3) ± 0.10

2.5

Bonaire

9

14

8.1

(7.3-8.5) ± 0.16

3.8

Curagao

9

23

8.1

(7.3-8.5) ±0.11

3.1

Breadth across upper molars

Bonaire

d

12

5.9

(5.3-6. 1) ± 0.14

4.0

Curagao

d

18

6.0

(5. 7-6.3) ± 0.08

2.8

Bonaire

9

13

5.8

(5.4-6. 1) ± 0.12

3.7

Curagao

9

22

6.0

(5.7-6.2) ± 0.06

2.3

1979

Genoways and Williams — Bonaire and CuRAf ao Bats

315

Because the morphometric relationship between the sexes of this
species is unknown (Swanepoel and Genoways, 1979), we tested sam-
pies of males and females for secondary sexual variation. Results of
these analyses (Table 1) revealed that females are significantly larger
in three (length of forearm, greatest length of skull, and condylobasal
length) of the eight measurements tested. In four of the remaining
measurements, the sexes averaged the same, and, in the fifth (length
of maxillary toothrow), females were 0.1 mm larger than males.

Miller’s (1900fl) Glossophaga elongata from Willemstad, Curasao,
was considered to be a distinct species until Koopman (1958) recog-
nized it as a subspecies of Glossophaga longirostris. We also com-
pared samples from Bonaire and Curasao by sex to explore the rela-
tionship between these island populations (Table 1). Males from
Curasao differed significantly from those on Bonaire in three mea-
surements. In two of these measurements (zygomatic breadth and mas-
toid breadth), the Curasao sample averaged larger, whereas in the
other measurement (condylobasal length) males from Bonaire were
larger. Females from Curasao averaged significantly larger than those
from Bonaire in three measurements (zygomatic breadth, mastoid
breadth, and breadth across upper molars). The exact meaning of the
differences between these populations is unclear at the present time
and must await comparisons with samples from Aruba and the adjacent
mainland. Also genic information would be useful in documenting re-
lationships among these insular populations. For the present, we have
assigned populations from Curasao and Bonaire to Glossophaga lon-
girostris elongata.

Natalus tumidirostris tumidirostris Miller

Specimen examined (1).-™~Bonaire: 8.5 km N, 2 km W Kralendijk, 1.

This is the first specimen of this species to be reported from Bonaire,
making a total of seven chiropteran species known from the island.
Our single example is an adult male (testes, measuring 1.5 mm in
length) taken on 18 August 1977 under the conditions described in the
account of Mormoops megalophylla.

This taxon, originally described from Curasao by Miller (1900/?),
was reviewed by Goodwin (1959). We have assigned our specimen to
N. t. tumidirostris based on its small size, which matches that of ma-
terial reported from Curasao by Goodwin. Differences in overall size
are the main characteristics used to distinguish subspecies of N. tu-
midirostris. External and cranial measurements of our specimen are
as follows: length of forearm, 35.0; greatest length of skull, 15.6; con-
dylobasal length, 14.2; zygomatic breadth, 7.7; mastoid breadth, 7.0;
breadth of braincase, 7.6; postorbital breadth, 3.0; length of maxillary
toothrow, 6.7; breadth across upper molars, 5.1.

316 Annals of Carnegie Museum vol. 48

species of Myotis. Groups on the right side include M. albescens (1), M. keaysi (2),
and M. nigricans (3). On the left side are clustered M. nesopolus from the mainland
(L), Bonaire (B), and Curasao (C).

Myotis nesopolus nesopolus Miller

Specimens examined (29).-— Bonaire: Boliva Dist., 1 (AMNH); 8.5 km N, 2 km W
Kralendijk, 23. CuRAf ao: Punda, 1 (USNM); 2.8 km S, 4.5 km E Westpunt, 4.

The conditions under which these specimens were taken are dis-
cussed in earlier accounts. Of the 27 specimens taken during our work,
only two were females; these individuals were non-pregnant when cap-
tured on Bonaire on 21 August 1977. Five males from Bonaire had
testes length of 3, 4, 4, 4, and 4 when netted on the same date. i

The Neotropical species of the genus Myotis were recently reviewed |
by LaVal (1973), but he did not include the taxon M. nesopolus de- ;
scribed by Miller (1900a) from the island of Curasao. This taxon has
been considered (Koopman, 1958; Husson, 1960) a subspecies of the
widespread M. nigricans since the work of Miller and Allen (1928). In
order to determine the status of M. nesopolus we subjected morpho-
metric data for it along with data for similar- sized Myotis from north-
ern South America (albescens, keaysi, larensis, and nigricans) to ca-
nonical analysis.

Canonical analysis provides a mechanism for graphically represent-
ing phenetic relationships among samples with the characters weighted
by variance-covariance analysis (Fig. 1). In Table 2, characters used
in this analysis are listed from the most useful to the least useful in

1979

Genoways and Williams — Bonaire and CuRAfAO Bats

317

Table 2.~Variables used in discriminant function analysis of South American Myotis,
Characters are listed in order of their usefulness in distinguishing groups, with the
character with the greatest between-groups variance and the least within-groups vari-
ance being selected first. Other traits are ranked using the same criteria. The statistics
are recalculated at each step.

Step

Character

F-value

U-stati§tic

1.

Length of tibia

86.22

0.1864

2.

Length of forearm

83.80

0.0352

3.

Breadth of braincase

26.98

0.0146

4.

Greatest length of skull

8.82

0.0100

5.

Breadth across upper molars

4.05

0.0082

6.

Zygomatic breadth

2.82

0.0071

1.

Postorbital breadth

3.93

0.0059

8.

Postpalatal length

2.99

0.0050

9.

Length of third metacarpal

2.37

0.0044

10.

Depth of braincase

1.90

0.0040

IL

Length of upper toothrow

0.96

0.0038

discriminating groups. Variate I accounts for 82.8% of the total dis-
persion, and Variate II accounts for 13.1%. Characters with high pos-
itive canonical coefficients for Variate I (values greater than 1.0) are,
in decreasing order of values, postorbital breadth, breadth of brain-
case, postpalatal length, and length of forearm. Those with high neg-
ative values include, ordered as above, length of tibia, zygomatic
breadth, and length of upper toothrow. In Variate II, positive values
of more than 1.0 were exhibited by postorbital breadth, breadth of
braincase, and greatest length of skull, in order of decreasing value.
The greatest negative value was possessed by breadth across upper
molars in Variate II followed by depth of braincase and zygomatic
breadth.

In Fig. 1, the specimens of Myotis larensis and those from Curasao

Table 3.- — Matrix of classification, based upon the discriminant functions of II mor-
phometric characters. Values indicate the number of individuals classified into each

group.

Samples

Classification groups

1

2

3

4

5

1. Myotis albescens

18

0

0

0

1

2. Myotis keaysi pilosatibialis

0

16

0

0

2

3. Myotis nesopolus larensis

0

0

15

1

0

4. Myotis nesopolus nesopolus

0

0

0

11

0

5. Myotis nigricans nigricans

1

3

0

0

16

318

Annals of Carnegie Museum

VOL. 48

Table 4. — External and cranial measurements of two subspecies of Myotis nesopolus.

See text for specimens examined. An asterick following the name of a measurement

indicates that the taxa differ at the 0.05 level.

Taxon

N

Mean

(Range)

± 2SE

cv

Length of forearm*

M.

n.

nesopolus

12

30.3

(29.5-31.2)

±0.28

1.6

M.

n.

larensis

30

31.9

(30.4-33.2)

±0.26

2.2

Length of third metacarpal*

M.

n.

nesoplus

12

29.3

(28.2-30.0)

±0.30

1.8

M.

n.

larensis

30

31.0

(29.5-32.1)

±0.25

2.2

Length of tibia*

M.

n.

nesopolus

12

14.5

(14.3-14.9)

±0.12

1.5

M.

n.

larensis

30

14.9

(14.2-15.8)

±0.12

2.2

Greatest length of skull*

M.

n.

nesopolus

12

13.1

(12.7-13.4)

±0.11

1.4

M.

n.

larensis

32

13.6

(13.0_14.4)

±0.10

2.2

Depth of braincase

M.

n.

nesopolus

12

4.8

(4.6-4.9)

±0.07

2.4

M.

n.

larensis

32

4.9

(4.5-5.2)

±0.06

3.6

Zygomatic breadth*

M.

n.

nesopolus

11

7.9

(7.5-8. 1)

±0.11

2.2

M.

n.

larensis

17

8.2

(8.0-8.3)

±0.05

1.3

Breadth of braincase

M.

n.

nesopolus

12

6.1

(6.0-6.2)

±0.04

1.0

M.

n.

larensis

31

6.1

(5.8-6.4)

±0.05

2.2

Postorbital breadth

M.

n.

nesopolus

12

3.2

(3.0-3.3)

±0.05

2.9

M.

n.

larensis

33

3.3

(3.0-3. 5)

±0.04

3.2

Length of upper toothrow* (P-M^)

M.

n.

nesopolus

12

5.7

(5.4-5.9)

±0.08

2.4

M.

n.

larensis

33

6.1

(5.8-6.5)

±0.05

2.5

Breadth across upper molars*

M.

n.

nesopolus

12

5.0

(4.9-5. 1)

±0.04

1.3

M.

n.

larensis

32

5.2

(4.8-5.6)

±0.05

2.9

Postpalatal length

M.

n.

nesopolus

12

4.5

(4.0-4.7)

±0.11

4.3

M.

n.

larensis

30

4.6

(4.2~5.0)

±0.09

5.4

and Bonaire are grouped on the left side of the plot. Another group,
composed of M. albescens, M. keaysi, and M. nigricans is found on
the right side of the plot. These two groups are widely separated on
Variate 1. Clearly, mainland M. larensis and the Myotis from the is-

1979

Genoways and Williams — Bonaire and CuRAf ao Bats

319

lands are distinct from the other species; however , their relationship
to each other is less clear. Examination of Fig. 1 indicates that these
samples can not be distinguished on either Variate I or Variate II. The
classification matrix (Table 3), on the other hand, shows only one
specimen of larensis misclassified with specimens from Curasao and
Bonaire. All specimens from Curasao and Bonaire are classified in
their own group.

This analysis clearly reveals that the material from Curasao and
Bonaire, to which the name M. nesopolus would apply, is most closely
related to M. larensis of the South American mainland. The question
remains as to whether these taxa are closely related species or sub^
species of a single species. A univariate comparison of the two taxa
(Table 4) indicates that M. nesopolus is significantly smaller than M.
larensis in seven of the 11 measurements tested and they average
smaller in three of the other measurements. LaVal (1973) characterized
M. larensis as having a very long third metacarpal, tibia, and skull in
relation to length of forearm (mean ratios 0.96, 0.48, and 0.43, respec-
tively). These ratios for M. nesopolus are 0.97, 0.48, and 0.43, re-
spectively. As is true for larensis, nesopolus can be distinguished on
absolute length of forearm (shorter) and tibia (longer) from nigricans,
albescens, and keaysi. Our specimens also generally seem to agree
with larensis in coloration and distribution of fur. We think that the
relationship between these taxa is best represented by considering
them to be distinct subspecies of a single species, with the island form
being characterized by overall smaller size. M. nesopolus is the senior
synonym for this species and should be treated as follows:

Myotis nesopolus nesopolus Miller

1900, Myotis nesopolus Miller, Proc. Biol. Soc. Washington, 13:123, April 6.

1928. Myotis nigricans nesopolus, Miller and Allen, Bull. U.S. Nat. Mus., 144:182, May
25.

Type locality.— Punda. area, Willemstad, Curasao.

Holotype. — Adult male in alcohol with skull removed, 101849
USNM, collected on 4 November 1899 by L. B. Smith.

Distribution.— Known from the islands of Curasao and Bonaire,
Dutch West Indies.

Remarks .—Provions reports of Myotis nigricans from these islands
by Hummelinck (1940), Koopman (1958), Husson (1960), and others
almost certainly pertain to M. nesopolus although we have not ex-
amined all reported specimens.

Myotis nesopolus larensis LaVal

1973. Myotis larensis LaVal, Sci. Bull. Nat. Hist, Mus. Los Angeles Co., 15:44, Feb-
ruary 14.

320

Annals of Carnegie Museum

VOL. 48 I

Type locality. — Rio Tocuyo, Lara, Venezuela. j

Holotype .—Adult female, skin and skull, 130709 AMNH, taken on 1
23 March 1938 by G. H. H, Tate.

Distribution. —Known from three localities around the Golfo de
Venezuela in northwestern Venezuela.

Remarks. — For additional information concerning this taxon see
LaVal (1973).

Specimens examined (33).- — Venezuela: Capatarida, Falcon, 20 (USNM); Ri'o To-
cuyo, Lara, 12 (AMNH); 110 km N, 25 km W Maracaibo, Zulia, 1 (USNM).

'i

Additional Specimens Examined .

j'

Myotis albescens (20).— Venezuela: Belen, T. F. Amazonas, 2 (USNM); 25 km SSE '
Puerto Ayacucho, T. F. Amazonas, 1 (USNM); San Juan, T. F. Amazonas, 8 (USNM); ,,
41 km NW Puerto Paez, Apure, 9 (USNM). i

Myotis keaysi pilosatibialis (18). — Venezuela: Rancho Grande Biological Station,
Aragua, 9 (USNM); 4 km NW Montalban, Carabobo, 1 (USNM); 9.4 km N Caracas, D.

F., 1 (USNM); 5 km N Caracas, 1 (USNM); 19 km E Caracas, Miranda, 4 (USNM); 8 [

km S Caracas, Miranda, 2 (USNM).

Myotis nigricans nigricans (20). — Venezuela: 9.4 km N Caracas, D. F., 3 (USNM); |
1 km N, 3 km W Caripe, Monagas, 7 (USNM); 19 km NW Urama, Yaracuy, 10 (USNM).

Molossus molossus pygmaeus Miller |

Specimen examined (1). — Bonaire: 8.5 km N, 2 km W Kralendijk, 1. :

Our specimen is a non-pregnant female taken on the night of 18 |
August 1977. Conditions under which the specimen was taken are de- j
scribed in the account for Mormoops megalophylla.

This species has been reported previously from Bonaire (Humme-
linck, 1940; Husson, 1960). There is considerable confusion over the
taxonomic status and relationships of small Neotropical Molossus. We
follow Jones et al. (1971) in considering “most, if not all, mainland
populations of small Molossus with pale^based hairs pertain to the
species Molossus molossus, originally described from the Lesser An- i
tilles.” However, this view is not held by all recent authors (see for
example, Handley, 1976). The taxon pygmaeus was originally de=
scribed from Curasao by Miller (19001?) and this name has been applied
to specimens from Bonaire by earlier authors (see Hummelinck, 1940; !
Husson, 1960). We consider pygmaeus to be a subspecies of M. mo-
lossus here but its status must be determined in the future when this
entire complex is studied.

Acknowledgments

Fieldwork resulting in specimens reported herein was supported by the M. Graham
Netting Research Fund through a grant from the Cordelia Scaife May Charitable Trust.
Permits to collect on Bonaire and Curasao were granted by Mr. R. Gorsira, Head of
Veterinary Service, Abattoir-Parera, Willemstad, Curasao.

1979

Genoways and Williams — Bonaire and CuRAf ao Bats

321

We would like to thank Daniel F. Williams for assistance with the statistical programs,
M. de la Fuente for field assistance, and Karl F. Koopman, (American Museum of
Natural History) and Don E. Wilson, (National Fish and Wildlife Laboratory) for allow-
ing us to examine material in their care.

Literature Cited

Dixon, W. J., and M. B. Brown (eds,). 1977. BMD P-77: Biomedical Computer
Programs, P-series. Univ. California Press, Berkeley, xiii + 880 pp.

Goodwin, G. G. 1959. Bats of the subgenus Natalus. Amer. Mus. Novitates, 1977:1-

22.

Handley, C. O., Jr. 1976. Mammals of the Smithsonian Venezuelan Project. Brigham
Young Univ. Sci. Bull., Biol. Ser., 20:1-89.

Hummelinck, P. W. 1940. A survey of the mammals, lizards, and mollusks. Studies
on the Fauna of Curasao, Aruba, Bonaire, and the Venezuelan Islands, 1:59-108.
Husson, a. M. 1960. De zoogdieren van de Nederlandse Antillen. Mammals of the
Netherlands Antilles. Fauna Ned. Antillen, ’s- Gravenhage and Willemstad, viii
+ 170 pp.

Jones, J. K., Jr., J. D. Smith, and R. W. Turner. 1971. Noteworthy records of bats
from Nicaragua, with a checklist of the chiropteran fauna of the country. Occas.
Papers Mus. Nat. Hist., Univ. Kansas, 2:1-35.

Koopman, K. F. 1958. Land bridges and ecology in bat distribution on islands off the
northern coast of South America. Evolution, 12:429-439.

LaVal, R. K. 1973. A revision of the Neotropical bats of the genus Myotis. Sci. Bull.
Nat. Hist. Mus. Los Angeles Co., 15:1-54.

Miller, G. S., Jr. 1900^. Three new bats from the state of Curasao. Proc. Biol. Soc.
Washington, 13:123-127.

— . 1900^. A second collection of bats from the island of Curasao. Proc. Biol. Soc.

Washington, 13:159-162.

Miller, G. S., Jr., and G. M. Allen. 1928. The American bats of the genera Myotis
and Pizonyx. Bull. U.S. Nat. Mus., 144: viii + 1-218.

Smith, J. D. 1972. Systematics of the chiropteran family Mormoopidae. Misc. Publ.
Mus. Nat. Hist., Univ. Kansas, 56:1-132.

Smith, J. D., and H. H. Genoways. 1974, Bats of Margarita Island, Venezuela, with
zoogeographic comments. Bull. Southern California Acad. Sci,, 73:64-79.

SoKAL, R. R., AND F, J. Rohlf. 1969. Biometry: the principles and practice of statistics
in biological research. W. H. Freeman and Co., San Francisco, xii -l- 776 pp.
SwANEPOEL, P., AND H. H. Genoways. 1979. Morphometries. Pp. 13-106, in Biology
of bats of the New World family Phyllostomatidae, Part III (R. J. Baker, J. K.
Jones, Jr., and D. C. Carter, eds.), Spec, Publ. Mus., Texas Tech Univ., 16:1-441.
Wilson, D. E. 1979. Reproductive patterns. Pp. 317-378, Biology of bats of the New
World family Phyllostomatidae, Part III (R. J. Baker, J. K, Jones, Jr., and D. C.
Carter, eds.). Spec. Publ. Mus., Texas Tech Univ., 16:1-441.

Back issues of many Annals of Carnegie Museum articles are
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at half price. Orders and inquiries should be addressed to:
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ANNALS

0/ CARNEGIE MUSEUM

CARNEGIE MUSEUM OF NATURAL HISTORY
4400 FORBES AVENUE ® PITTSBURGH, PENNSYLVANIA 15213
VOLUME 48 7 SEPTEMBER 1979 ARTICLE 18

RECORDS OF BATS (MAMMALIA: CHIROPTERA)

FROM SURINAME

' Hugh H. Genoways

Curator, Section of Mammals

I Stephen L. Williams

■ Collection Manager, Section of Mammals

!

S Abstract

Ten species are added to the 60 species of bats already known to occur in Suriname.
The species added include Micronycteris minuta, M. nicefori, Phyiioderma stenops,
Tonatia bidens, T. brasiliense, CarolUa brevicauda, Chiroderma trinitatum, Vampyressa
I bidens, Promops centralis, and P. nasutus. Additional information is presented on five
I species previously recorded from Suriname, including Pteronotus parnellii, Mimon
I crenulatum, Artibeus concolor, Chiroderma villosum, and Sturnira tildae.

\

Introduction

“The Bats of Suriname” by Husson (1962) stands not only as a
- classical work on mammals for the country of Suriname but for all of
South America. In this work, Husson (1962) reported 58 species of
bats as being known to occur in the country. Husson (1973, 1978) listed
I four additional species {Micronycteris brachyotis, Macrophyllum ma-
crophyilum, Sturnira tildae, and Molossus sinaloae) for the country.
Other authors have changed the status of two of these species-Am^-
trida minor {-Ametrida centurio, Peterson, 1965) and Myotis surina-
mensis (=unknown species not from South America, LaVal, 1973).

This research and publication supported by the M. Graham Netting Research Fund
through a grant from the Cordelia Scaife May Charitable Trust.

Submitted for publication 23 December 1978.

323

324

Annals of Carnegie Museum

VOL. 48

Eger (1977) placed Eumops geijkesi as a synonym of E. maurus.
Therefore, when our work began in Suriname, the chiropteran fauna
was known to consist of 60 species.

The major portion of our material resulted from a field party con-
sisting of S. L. Williams and M. de la Fuente, who collected mammals
in Suriname from 1 July to 15 August 1977. The Carnegie Museum of
Natural History also houses small collections of mammals made by M.
de la Fuente in October 1966 and February 1975. Included in these
collections were 1270 specimens of bats representing approximately
44 species. Ten of these species have not been recorded from Suriname
previously and are discussed below. Also discussed below are speci-
mens of five other species for which our data adds significantly to that
reported by Husson (1962, 1978). The chiropteran fauna of Suriname
is now known to consist of 70 species.

Methods and Materials

Our specimens were taken primarily by mist-netting and were preserved as skin /skull
or fluid preparations. The specimens are deposited in the Section of Mammals, Carnegie
Museum of Natural History.

Measurements of forearm and cranial dimensions were taken with dial calipers ac-
curate to 0. 1 mm. Only adult specimens (phalangeal epiphyses completely fused) were
measured in this study. Our measurements were taken as follows:

Length of Forearm. —Taken from the posteriormost projection of the elbow (olecra-
non process) to the anteriormost projection of the wrist joint with the wing in a flexed
position.

Greatest Length of Skull. — Distance from the posteriormost projection of the cranium
to the anterior edge of the upper incisors.

Condylobasal Length. — Distance from the posteriormost projection of the exoccipal
condyles to the anteriormost projection of the premaxillae.

Zygomatic Breadth. — Greatest distance across the zygomatic arches at right angles
to the longitudinal axis of cranium.

Mastoid Breadth. — Greatest distance across the mastoid processes at right angles to
the longitudinal axis of cranium.

Postorbital Breadth. — Least distance across the postorbital constriction at right angles
to the longitudinal axis of cranium.

Length of Maxillary Toothrow. — Distance from posterior lip of alveolus of M^ to the
anterior lip of alveolus of C*.

Breadth Across Upper Mo/ar.s.-— Greatest distance across upper molars at right angles
to the longitudinal axis of cranium; measured at the lateralmost projections of the labial
edges of the upper molars.

A few field weights were taken with a pan balance. Reproductive condition of the
skin /skull specimens was determined by gross dissection in the field, whereas the fluid
preserved specimens were dissected in the laboratory.

Acknowledgments

We are most grateful to Dr. Joop P. Schulz, Henry A. Reichart, Ferdinand L. J. Baal,
and K. Mohadin, members of the Nature Conservation Division of the Forestry Service,
Government of Suriname, and Stichting Natuurbehoud Suriname (STINASU), for is-
suing our scientific collecting permits and assisting us in all phases of our work while

1979

Genoways and Williams—Suriname Bats

325

in Suriname. We also wish to thank Dr. Karl F. Koopman, American Museum of Natural
History, and Dr. Patricia Freeman, Field Museum of Natural History, for allowing us
to examine specimens housed in their collections. K. F. Koopman, C. O. Handley, Jr.,
and A. L. Gardner confirmed the identifications of some species. Mr. M. de la Fuente
assisted with field work and his parents accommodated the field party while in Para-
maribo.

Financial support for field work in Suriname was received from the M. Graham Net-
ting Research Fund established by a grant from the Cordelia Scaife May Charitable
Trust.

Species Accounts

Pteronotus parneliii rubiginosus (Wagner)

Specimens Examined (4). — Brokopondo: Brownsberg Nature Park, 3 km S, 20 km

W Afobakka, 4.

Remarks .—Oni specimens are from the northeastern part of Suri-
name; the only previous record (Husson, 1962, 1978) of the species
was based on a single specimen from Tafelberg in the central part of
the country. Our specimens are three adult females and one adult male
collected on 7 July 1977. Two of the females were carrying embryos
that measured 30 and 31 mm in crown-rump length; the other evinced
no reproductive activity. The male had testes that were 5 mm long.
The specimens were taken in mist nets set across roads in an area of
mature tropical forest with large trees and moderate ground vegetation.
Other species taken at this place were Saccopteryx bilineata, Micro-
nycteris megalotis, Tonatia bidens, Anoura caudifer, Rhinophylla
pumilio, Sturnira tildae, and Eptesicus brasiliensis .

Smith (1972) revised this genus and applied the name parneliii to
this species. The subspecies rubiginosus was characterized as being
the largest of the species both cranially and externally. Length of fore-
arm of these specimens measured as follows (male followed by three
females): 62.3; 66.1; 65.9; 64.1.

Micronycteris minuta (Gervais)

Specimens Examined (2).— -Marowijne: 10 km N, 24 km W Moengo, 1. Suriname:
Powaka, 1.

Remarks Out specimens of M. minuta are the first to be recorded
from Suriname. At the place 10 km N, 24 km W Moengo, nets were
placed in and along the edge of a tropical forest bordering a river. At
Powaka, the vegetation again was primarily tropical forest and nets
were placed across a pond, roads, trails, and a creek. The specimen
from Powaka was a female that was carrying an embryo that was 13
mm in crown-rump length when taken on 30 July 1977. The other
individual was also a female but it did not evince reproductive activity
when obtained on 6 August 1977.

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This species is currently considered to be monotypic (Jones and
Carter, 1976). Our specimens from Suriname when compared with
individuals from Trinidad and Brazil, deposited in the American Mu-
seum of Natural History, revealed no distinguishing characteristics
(see also Goodwin and Greenhall, 1961: 228-229). External and cranial
measurements of our specimens (as listed above) are as follows: length
of forearm, 34.3, 35.2; greatest length of skull, 18.7, 18.8; condylobasal
length, 16.2, 16.2; zygomatic breadth, 8.8, 8.8; mastoid breadth, 8.7,
8.7; postorbital breadth, 4.2, 4.2; length of maxillary toothrow, 6.8,
6.6; breadth across upper molars, 6.0, 5.9.

Micronycteris nicefori Sanborn

Specimens Examined (2). — -Marowijne: 10 km N, 24 km W Moengo, 1. Suriname:
Powaka, I.

Remarks.— Our specimens of M. nicefori are the first to be recorded
from Suriname. These specimens were taken at the same places dis-
cussed above for M. minuta. The specimen from Moengo was an adult
male that had testes measuring 5 mm on 6 August 1977, whereas the
specimen from Powaka was a non-pregnant female taken on 30 July
1977.

This species is currently considered to be monotypic. We could
detect no differences in either size or morphology of the skulls and
teeth between our specimens and those from Trinidad and Colombia
deposited in the American Museum of Natural History. Both of our
specimens possess a faint, but distinct, mid-dorsal, white stripe. Ex-
ternal and cranial measurements of the specimens are as follows (male
followed by female): length of forearm, 36.1, 38.0; greatest length of
skull, 20.1, 20.3; condylobasal length, 18.0, 18.4; zygomatic breadth,
9.5,—; mastoid breadth, 8.7, 8.5; postorbital breadth, 4.3, 4.0; length
of maxillary toothrow, 7.3, 7.5; breadth across upper molars, 6.1, 6.0.
The male weighed 8 g and the female 7 g.

Mimon crenulatum crenulatum (E. Geoffroy St. Hilaire)

Specimens Examined (3).— Suriname: Powaka, 3.

Remarks .—Husson (1978) reported single specimens of M. crenu-
latum from southwestern Suriname and north of Paramaribo and two
from unknown localities. Two of our specimens were adult males that
had testes measuring 7 mm (30 July 1977) and 8 mm (9 August 1977).
The female was pregnant with an embryo that measured 18 mm in
crown-rump length when captured on 30 July 1977.

Handley (1960) discussed geographic variation in M. crenulatum; he
applied the subspecific name crenulatum to populations in northeast-
ern South America. Husson (1962, 1978), as we do here, followed this

1979

Genoways and Williams — Suriname Bats

327

arrangement. We have followed Handley (1960; see also Gardner and

Patton, 1972) in use of the generic name Mimon for this species in
place of Anthorhina, which was used by Husson (1962, 1978). External
and cranial measurements of our three specimens (two males and one
female, respectively) are as follows: length of forearm, 47.6, 48.4, 49.0;
greatest length of skull, 21.7, 22.2, 21.7; condylobasal length, 19.1,
19.2, 19.1; zygomatic breadth, 11.8, 12.1, 11.9; mastoid breadth, 11.0,
11.5, 11.4; postorbital breadth, 4.0, 4.1, 3.6; length of maxillary tooth-
row, 7.7, 7.8, 7.8; breadth across upper molars, 8.2, 8.4, 8.3. The
males weighed 7 g and 8 g.

Phylloderma stenops stenops Peters

Specimen Examined (1). — Coronie: Totness, 1.

Remarks .—Although Husson (1962) presents an extensive discus-
sion concerning this species, he did not have a specimen from Suri-
name of this rare species. Our specimen was taken on the night of 29
July 1977 in a net placed across a trail through a coconut grove. Other
species taken in the same area included Glossophaga soricina, Car-
oliia perspicillata, Artibeus cinereus, and a large-sized species of Ar-
tibeus. The specimen is a male that had testes that were 8 mm long.

Handley (1966) recognized two subspecies of this bat, with the name
P. s. stenops being applied to all material from South America. Husson
(1962) presented an extensive discussion, including measurements of
the holotype of this taxon from Cayenne, French Guiana. External and
cranial measurements of our specimen are as follows: length of fore-
arm, 71.8; greatest length of skull, 30.8; condylobasal length, 26.5;
zygomatic breadth, 15.0; mastoid breadth, 13.7; postorbital breadth,
9.1; length of maxillary toothrow, 10.2; breadth across upper molars,
9.7. These measurements are similar to those of the holotype (Husson,
1962; Carter and Dolan, 1978) and specimens from Guyana and north-
eastern Brazil (Hill, 1964).

j Tonatia bidens hidens (Spix)

! Specimens Examined (2),— Brokopondo: Brownsberg Nature Park, 3 km S, 20 km
I W Afobakka, 1, Saramacca: Bigi Poika, 1,

Remarks.- — Our specimens are the first of T. bidens to be reported
! from Suriname. Husson (1962) has shown that the specimen that
formed the basis of Jentink’s (1887) earlier report of this species from
Suriname is actually a Mimon bennettii. The condition under which
the specimen from Brownsberg Nature Park was taken is discussed in
the account of Pternotus parnellii. At Bigi Poika, the specimen of T.
bidens was taken in a net placed along a dock that passed through a
I swampy area to the edge of a river. The specimens are both adult
I

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VOL. 48

males having testes measuring 2 (7 July 1977) and 6 (14 July 1977) mm
long.

Also represented in our collection from Suriname is the closely re-
lated and morphologically similar species T. silvicola (also reported
previously by Husson, 1962, 1978; see Davis and Carter, 1978, for this
spelling of specific name). Based upon our material and specimens
from Brazil and Guyana housed in the American Museum of Natural
History, these species may be distinguished as follows: the two lower
incisors of bidens are large as compared with the almost spicule-like
incisors of silvicola, posterior to which in some cases the canines
meet; the skull of bidens is smaller and more delicate in appearance;
although a sagittal crest is clearly present in specimens of bidens, it
is never as greatly enlarged (high) as in silvicola ; in bidens the outline
of the skull slopes gently from the rostrum to the braincase, whereas
in silvicola the frontal region of the skull is inflated so that the brain-
case arises abruptly from the rostral region.

All Recent specimens of T. bidens are assigned to the nominate
subspecies (see Jones and Carter, 1976; Koopman, 1976). External and
cranial measurements of our two males (as listed above) are as follows:
length of forearm, 56.5, 55.1; greatest length of skull, 26.8, 27.7; con-
dylobasal length, 23.3, 23.1; zygomatic breadth, 13.2, 13.8; mastoid
breadth, 12.5, 12.3; postorbital breadth, 5.3, 5.6; length of maxillary
toothrow, 9.2, 9.5; breadth across upper molars, 8.3, 8.5. These mea-
surements are slightly smaller than those of a female from Guyana
(Hill, 1964).

Tonatia brasiliense (Peters)

Specimen Examined (I).--Brokopondo: Brownsberg Nature Park, 7 km S, 18.5 km
W Afobakka, 1.

Remarks specimen represents the first recorded occurrence

of T. brasiliense in Suriname. This specimen was taken in nets along
the forest edge near the headquarters for the Brownsberg Nature Park.
Other species taken in this area included Tonatia silvicola, Anoura
caudifer, Carollia perspicillata, Rhinophylla pumilio, Sturnira lilium,
Vampyrops helleri, Artibeus cinereus, and two large-sized species of
Artibeus. This adult male had testes measuring 4 mm long when cap-
tured on 9 July 1977.

There are four named taxa of small Tonatia occurring in the Nqo-
tropics— hras Hie ns e , minuta, nicaraguae, and venezuelae. Gardner
(1976) suggested that these taxa are synonymous, with T. brasiliense
the senior synonym. Other recent authors (Handley, 1976; Koopman,
1978) have followed this arrangement, as we do here. The infraspecific
variation of this species needs review before any subspecific names

1979

Genoways and Williams — Suriname Bats

329

are applied. External and cranial measurements of our specimen are
as follows: length of forearm, 35.5; greatest length of skull, 19.6; com
dylobasal length, 16.6; zygomatic breadth, 9.2; mastoid breadth, 8.8;
postorbital breadth, 3.0; length of maxillary toothrow, 7.9; breadth
across upper molars, 6.1.

Carollia brevicauda (Schinz)

Specimen Examined (1).— Brokopondo: Brownsberg Nature Park, 6 km S, 20 km W
Afobakka, 1.

Remarks .-—Om specimen of C. brevicauda is the first of the species
to be recorded from Suriname. However, the exact identity of speci-
mens reported as Carollia castanea castanea by Husson (1962, 1978)
must await examination of the specimens; the measurements of the
two specimens are quite similar to those of our specimen. The speci-
men was taken, along with individuals of Glossophaga soricina and
Carollia perspicillata, within 3 meters of the entrance of a small mine
dug into a hillside. The vegetation in the surrounding area was dense
tropical forest. The specimen is an adult male that had testes measuring
7 mm long when captured on 9 July 1977,

Pine (1972) considered this species to be monotypic. In several mea-
surements, our specimen was smaller than any individual reported by
Pine (1972) or Swanepol and Genoways (1978). Carollia brevicauda
is difficult to distinguish from the sympatric C. perspicillata. Our spec-
imen was smaller than individuals of C. perspicillata from Suriname.
Also, cranial characters as outlined by Pine (1972) were generally use-
ful in distinguishing the specimen, particularly the more curved and
posteriorly divergent upper toothrows, more bowed-out rami of the
lower jaw, and the outer lower incisors not being obscured by the
canines when viewed from above. External and cranial measurements
of our specimens are as follows: length of forearm, 37.3; greatest length
of skull, 20.9; condylobasal length, 18.4; mastoid breadth, 10.2; post-
orbital breadth, 5.4; length of maxillary toothrow, 6.6; breadth across
upper molars, 7.7.

Sturnira tildae de la Torre

Specimens Examined (5). — Brokopondo: Brownsberg Nature Park, 3 km S, 20 km
W Afobakka, 5.

Remarks .—Husson (1978) reported a single specimen of S. tildae
from Tempati, Commewijne District, and Hill (1964) has reported the
species from Guyana. Our five specimens further confirm the presence
of the species, at least in northern portions of Suriname. These spec-
imens were all taken in an area of mature tropical forest. One of the
specimens was an adult male with testes 7 mm long, whereas the other

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VOL. 48

four specimens were non-pregnant females. All were taken on 7 July
1977.

This species was originally described from Trinidad by de la Torre
(1959). The pelage of our specimens agrees with the original descrip-
tion; each hair reveals four bands of color. The upper inner incisors
of our specimens are bilobed with each lobe of equal size; lower in-
cisors are trilobed. We were able to appreciate the differences between
S. tildae and S. lilium in the size and position of the small posterio-
medial cusp on and the size of the lingual cusps on the lower
molars, but were unable to discern the supposed differences between
the two species in the height of the lingual cusps of the upper molars.

Marinkelle and Cadena (1971) gave measurements of individuals
from Colombia, Guyana, and Trinidad. Our specimens agree closely
with those data. External and cranial measurements of our one male
and two females, respectively, are as follows: length of forearm, 46.5,
44.9, 47.0; greatest length of skull, 23.9, 22.6, 23.1; condylobasal
length, 21.2, 20.2, 21.0; zygomatic breadth, 14.7, 14.2, 14.1; mastoid
breadth, 12.8, 11.9, 12.2; postorbital breadth, 6.0, 6.4, 6.3; length of
maxillary toothrow, 6.8, 6.5, 6.6; breadth across upper molars, 8.2,
7.7, 7.7.

Artibeus concolor Peters

Specimens Examined (32).— Saramacca: 5 km S, 2 km W Bigi Poika, 9. Suriname:
Powaka, 4; 1 km S, 2 km E Powaka, 19.

Remarks. — Evidently, the only specimen of this species previously
reported from Suriname (Husson, 1962, 1978) is the holotype from
Paramaribo (Peters, 1865). However, we found this species to be rel-
atively abundant in several savannah-forest edge situations. At the two
places where our large samples were taken, we released several indi-
viduals of this species. All of the 18 females that we obtained of this
species were not pregnant on the following dates: 4 January (1 indi-
vidual); 18 July (5); 30 July (2); 10-11 August (10). Three males taken
on 10-11 August had testes measuring 3.5, 5, and 5 mm.

Artibeus concolor is a monotypic species (Jones and Carter, 1976).
External and cranial measurements of three males and three females,
respectively, from 1 km S, 2 km E Powaka are as follows: length of
forearm, 45.1, 46.7, 47.0, 48.2, 48.3, 49.6; greatest length of skull, 21.2,
21.6, 21.2, 22.2, 22.0, 22.1; condylobasal length, 18.3, 18.6, 18.8, 19.6,

19.4, 19.1; zygomatic breadth, 13.1, 12.8, 12.9, 13.0, 14.0, 13.3; mas-
toid breadth, 11.3, 11.3, 11.1, 11.5, 12.0, 11.6; postorbital breadth, 5.4,

5.4, 5.7, 5.3, 6.1, 5.6; length of maxillary toothrow, 7.0, 6.7, 6.8, 7.4,
7.0, 6.9; breadth across upper molars, 8.9, 9.2, 9.2, 9.1, 9.6, 9.0. These
measurements are within the range of series from Guyana (Hill, 1964;

1979

Genoways and Williams— Suriname Bats

331

Swaeepoel and Genoways, 1978), Venezuela (Linares, 1969), and Co-
lombia (Barriga-Bonilla, 1965).

Chiroderma trinitatum trinitatum Goodwin

Specimens Examined (4). — Marowijne: 10 km N, 24 km W Moengo, 2. Nickerie;

26 km S, 55 km E Apoera, 1. Suriname: 1 km S, 2 km E Powaka, 1.

Remarks .—Out specimens are the first of this species to be recorded
from Suriname. The habitat northwest of Moengo was discussed in the
account for Micronycteris minuta. At the place 26 km S, 55 km E
Apoera, mist nets were set over a bridge crossing a river and an ad-
jacent clearing. Other species taken at this place were Carollia per-
spicillata, Artibeus cinereus, Chiroderma villosum, Sturnira lilium,
Vampyrops helleri, and two large-sized species of Artibeus. Near Po-
waka, nets were placed in a savannah and along the edge of a tropical
forest. More than 250 bats of the following species were collected:
Carollia perspicillata; Artibeus cinereus; A. concolor; two large-sized
species of Artibeus \ Chiroderma villosum; Vampyressa bidens; Uro-
derma bilobatum ; Vampyrops helleri ; Molossops planirostris . One of
the specimens from northwest of Moengo was a non-pregnant female
taken on 6 August 1977. The other three individuals were adult males
that possessed testes that measured 3 (22 July), 4 (6 August), and 4
mm (10 August) in length.

According to Jones and Carter (1976), material from Suriname
should represent the subspecies C. t. trinitatum, which they recorded
as occurring on Trinidad and in Amazonian South America south to
Peru and Brazil. However, our specimens from Suriname are much
smaller than typical trinitatum (Goodwin, 1958; Swanepoel and Gen-
oways, 1978) and seem to match specimens assigned to C. t. gorgasi
from Venezuela (Ojasti and Linares, 1971), Colombia (Barriga-Bonilia,
1965), and Panama (Handley, 1960). We have assigned our specimens
solely on geographic ground pending the completion of our ongoing
study of variation throughout the geographic range of this species.
External and cranial measurements of the three males (localities in
order listed above) and female, respectively, are as follows: length of
forearm, 38.4, 38.5, 38.8, 39.2; greatest length of skull, 21.2, 20.3, 20.8,
21.3; condylobasal length, 18.5, 17.7, 18.3, 18.8; zygomatic breadth,

12.8, 12.2, 13.1, 12.0; mastoid breadth, 10.6, 10.2, 10.5, 10.6; postor-
bital breadth, 5.1, 4.7, 4.9, 5.4; length of maxillary toothrow, 6.6, 6.6,

6.8, 6.7; breadth across upper molars, 9.3, 8.4, 9.5, 8.8.

Chiroderma villosum villosum Peters

Specimens Examined (2).- — Nickerie: 26 km S, 5.5 knn E Apoera, 1. Suriname: 1

km S, 2 km E Powaka, 1.

332

Annals of Carnegie Museum

VOL. 48

Remarks .—}lu?>son (1978) reported two specimens of this species
from Suriname. The conditions under which our two specimens were
taken are discussed in the account of C. trinitatum. One specimen is
an adult male with testes 4 mm long when obtained on 22 July 1977
and the other specimen is a non-pregnant female taken on 10 August
1977 (in order listed above).

According to Handley (1960) and Husson (1962, 1978) there are two
recognized subspecies within this species, with the nominate subspe-
cies, C. V. villosum, occupying South America except for northern
Colombia. External and cranial measurements of our male and female,
respectively, are as follows: length of forearm, 45.0, 47.3; greatest
length of skull, 24.4, 25.2; condylobasal length, 22.0, 22.7; zygomatic
breadth, 15.7, 16.2; mastoid breadth, 11.9, 12.5; postorbital breadth,
5.9, 5.7; length of maxillary toothrow, 8.6, 8.7; breadth across upper
molars, 11.2, 1 1.2. Weight of the female was 23 g. These measurements
are very similar to those given by Husson for his female specimen
from Suriname.

Vampyressa bidens (Dobson) '

Specimens Examined (2).— Suriname: 1 km S, 2 km E Powaka, 2. ;

Remarks. — These specimens represent the first recorded occurrence
of V. bidens in Suriname. The conditions under which these specimens ^
were collected are discussed in the account for Chiroderma trinitatum. i
Both specimens are adult females taken on 11 August 1977. One in-
dividual was carrying a single embryo that measured 16 mm in crown- i'
rump length; the other was not pregnant.

Our specimens show the dental formula (i 2/1, c 1/1, p 2/2, m 2/3)
typical of this species (Peterson, 1968). This species is monotypic '
(Jones and Carter, 1976). External and cranial measurements of our
two females are as follows: length of forearm, 37.1, 37,6; greatest
length of skull, 20.0, 20.6; condylobasal length, 17.4, 18.4; zygomatic '
breadth, 11.6, 12.0; mastoid breadth, 10.0, 10.2; postorbital breadth, j
5.0, 5.3; length of maxillary toothrow, 6.3, 6.5; breadth across upper
molars, 8.6, 8.3. These measurements agree well with material from ■
Guyana (Hill, 1964) and a large series from Peru (Davis, 1975).

Promops centralis centralis Thomas

Specimen Examined (1).— Marowijne: 10 km N, 24 km W Moengo, 1.

Remarks .—Thh is the first specimen of this species to be reported
from Suriname. The specimen was taken in a mist net set across the
edge of a river at the above locality. The individual is an adult female
that carried no embryo, but was lactating, when taken on 5 August |l
1977.

1979

Genoways and Williams— “Suriname Bats

333

Several recent authors, Handley (1966), Ojasti and Linares (1971),
and Koopmae (1978), have considered F, davisoni and occultus as
conspecific with F. centralis, which originally was described from
Yucatan. We agree with the assignment of occultus but examination
of specimens of davisoni from Ecuador and Peru in the American
Museum of Natural History and Field Museum of Natural History has
led us to question its placement with centralis. These specimens of
davisoni are relatively small and appear to us to approach the smaller
species P. nasuius; however, final judgment as to placement of this
taxon must await completion of our ongoing review of this genus.

Measurements of our specimen are similar to those of specimens of
F. c. centralis from Trinidad (Goodwin and Greenhall, 1961) and Ven-
ezuela (Ojasti and Linares, 1971). External and cranial measurements
of our specimen are as follows: length of forearm, 53.7; greatest length
of skull, 20.6; coedylobasal length, 18.8; zygomatic breadth, 12.7; maS“
toid breadth, 11.7; postorbital breadth, 3.7; length of maxillary tooth-
row, 7.7; breadth across upper molars, 8.8.

Promops nasutus Spix

Specimen Examined (1). — -Paramaribo: Paramaribo, 1.

Remarks .—Thh specimen represents the first recorded occurrence
of F. nasutus in Suriname. This adult female was collected by some
children from the roof of a house on 20 October 1966.

There appears to be at least five, and possibly six, subspecific names
available for bats in the F. nasutus complex in South America—
sutus, east-central Brazil; pamana, western Brazil; /os/cri, Paraguay;
ancilla, northern Argentina; downsi, Trinidad (Goodwin and Green-
hall, 1962). We believe that F. davisoni from Peru may pertain to this
group as well. Our specimen is geographically closest to F. n. downsi,
but our specimen has a much longer forearm than does the holotype
from Trinidad. The two specimens are quite close in cranial measure-
ments. Our specimen also approaches the size, both externally and
cranially, of specimens of the geographically distant F. n.fosteri. We
have not made a subspecific assignment of this specimen; this must
await an investigation of geographic variation throughout the species.
External and cranial measurements of our specimen are as follows:
length of forearm, 48.0; greatest length of skull, 18.6; condylobasal
length, 16.9; zygomatic breadth, 11.2; mastoid breadth, 10.8; postor-
bital breadth, 3.9; length of maxillary toothrow, 6.6; breadth across
upper molars, 8.0.

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VOL. 48

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LaVal, R. K. 1973. A revision of the Neotropical bats of the genus Myotis. Sci. Bull.
Nat. Hist. Mus. Los Angeles Co., 15:1-54.

Linares, O. J. 1969. Nuevos murcielagos para la fauna de Venezuela en el Museo de
Historia Natural La Salle. Mem. Soc. Cien. Nat. La Salle, 29:37-42.

Marinkelle, C. j., and A. Cadena. 1971. Remarks on Sturnira iildae from Colom-
bia. J. Mamm., 52:235-237.

1979

Genoways and Williams— Suriname Bats

335

Ojasti, J., and O. J. Linares. 1971. Adiciones a la fauna de murcielagos de Venezuela
con notas sobre las especies del genero DicUdurus (Chiroptera). Acta BioL Venez.,
7:421--441.

Peters, W. 1865. Uber Flederthiere (Vespertilio soricinus Pallus, Choeronycteris Lich-
tenst., Rhinophylla pumiiio nov. gen., Artibeus fallax nov. sp., A. concolor nov.
sp., Dermanum quadrivitratum nov. sp., Nycteris grandis nov. sp.). Monatsber.
Kon. Preuss. Akad. Wiss., Berlin, pp. 351-359.

Peterson, R. L. 1965. A review of the bats of the genus Ametrida, family Phyllo=
stomidae. Life Sci. Contrib., Royal Ontario Mus., 65:1-13.

. 1968. A new bat of the genus Vampyressa from Guyana, South America, with
a brief systematic review of the genus. Life Sci. Contrib., Royal Ontario Mus.,
73:1-17.

Pine, R. H. 1972. The bats of the genus Carollia. Tech. Monogr., Texas A&M Univ.,

8:1-125.

Smith, J. D. 1972. Systematics of the chiropteran family Mormoopidae. Misc. Publ.

Mus. Nat. Hist., Univ. Kansas, 56:1-132.

SwANEPOEL, P., and H. H. Genoways. 1978. Morphometries. Pp. 13-106, in Biology
of bats of the New World family Phyllostomatidae, Part III (R. J. Baker, J. K.

Jones, Jr., and D. C. Carter, eds.). Spec. Publ. Mus., Texas Tech Univ., 16:1-441.

Back issues of many Annals of Carnegie Museum articles are
available, and a few early complete volumes and parts are listed
at half price. Orders and inquiries should be addressed to:
Publications Secretary, Carnegie Museum, 4400 Forbes Avenue,
Pittsburgh, Pa. 15213.

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ANNALS

o( CARNEGIE MUSEUM

|| CARNEGIE MUSEUM OF NATURAL HISTORY
4400 FORBES AVENUE « PITTSBURGH, PENNSYLVANIA 15213
VOLUME 48 30 NOVEMBER 1979 ARTICLE 19

ALTO SALAVERRY: A PERUVIAN
COASTAL PRECERAMIC SITE

Shelia Pozorski

Research Associate, Section of Man

Thomas Pozorski

Assistant Curator, Section of Man

!

I Abstract

Alto Salaverry is one of two known Cotton Preceramic (2,500-1,800 B.C.) sites on
the coast of the Moche Valley, Peru. The site consists of both domestic and nondomestic
architecture, burials, and dense concentrations of refuse. Each of these site components
was studied in detail, resulting in much information about all features of the sitejt The
I architecture is highly variable and provides functional information as well as evidence
i of an incipient ranked society. Burial patterns are examined on the basis of interments
i both within and outside the confines of the site. Subsistence remains point to a predom-
' inantly marine economy supplemented by a great variety of plants.

The Site and Its Setting

Alto Salaverry, named for the pampa on which it lies, is located in
the extreme southeastern part of the Moche Valley on a bluff over-
i looking the Pacific Ocean (Fig. 1). Currently, the site lies some 1.5 km
I : inland from the sand beach immediately in front of the site and 3 km
I from the Salaverry Point rock mass further south. The Moche River
! mouth is fully 6 km away and to the north of the site.

I Coastal Uplift

1 The bluff upon which Alto Salaverry rests has been identified as a
I Pleistocene marine terrace about 120 m above sea level (Fred Nials,

I Submitted for publication 30 April 1979.

337

338

Annals of Carnegie Museum

VOL. 48

personal communication). About 400 m in front of the site and toward
the ocean, this terrace drops off sharply to a point where the land is
only about 4 to 6 m above sea level. Ongoing investigations within the
Moche Valley have revealed that the formation of much of the land
between Alto Salaverry and the ocean was a relatively recent phenom-
enon resulting from generalized coastal uplift correlated with tectonic
activity. When uplift occurred, the existing beach line was raised 4 to
6 m higher and substantial areas of previously submerged land were
no longer covered by water.

On the basis of the archaeological record, it appears that the major
documented time of tectonic activity was near the beginning of the
Moche portion of the Early Intermediate Period (A.D. 0-200). This
chronological placement is based on two lines of evidence— settlement
pattern data and subsistence changes. Survey by members of the Chan
ChamMoche Valley Project within the Moche Valley revealed that all
sites of a Salinar date or earlier, including both early ceramic and
Cotton Preceramic examples, are located back from the modern beach

1979

POZORSKI AND POZORSKI — AlTO SaLAVERRY

339

line and usually upon the largest now extant marine terrace. Later
sites, including a few middle to late Moche (A.D. 200-600) and many
Chimu (A.D. 1,000-1,470) examples, are present both upon the bluff
which existed prior to uplift and, more importantly, below the terrace
near the modern active beach in areas once under water. Of such sites,
the most notable are a Moche III-IV site near Huanchaco (Donnan
and Mackey, 1978) and scattered Chimu settlements associated with
sunken garden cultivation in areas of high water table.

A substantial change in the subsistence inventory at selected Moche
Valley sites has also been documented early in the Moche Period (Po-
zorski, 1976). At this time, several species of marine mollusks which
had been widely consumed earlier virtually disappear from the faunal
inventory. As these species were mainly taken from Huanchaco Bay
on the Moche Valley coast, it seems likely that their sudden near-
extinction is correlated with the coastline habitat changes resulting
from uplift.

Looking outside of the Moche Valley, comparable evidence in the
form of early site locations associated with the upper Pleistocene ter-
race suggests that coastal uplift may have been a widespread phenom-
enon—perhaps extending along much of the Peruvian coast. North of
the Moche Valley (Fig. 2), the Cotton Preceramic sites of Pulpar and
Huaca Prieta (Bird, 1948<2, 1948/?) are upon a comparable marine ter-
race, and in the vicinity of Huaca Prieta scattered ceramic cemeteries,
probably Late Intermediate Period (A.D. 1,000-1,470), lie along the
modern beach below the bluff. Within the Moche Valley, the precer-
amic site of Padre Aban, the Initial Period site of Gramalote, and a
Salinar Early Intermediate Period settlement— all in the vicinity of
Huanchaco Bay— are atop the marine terrace; and a Salinar site be-
tween Trujillo and the ocean is just above the 4 to 6 m contour ele-
vation and would have also been on the coast prior to uplift (Fig. 1).
To the south of the Moche Valley, the pattern would seem to continue
with marine terraces present though variable in height, and early sites
consistently back from the modern beach and upon the upper terrace.
Examples visited by the authors include preceramic and Initial Period
sites in the area of Guaeape in the Viru Valley (Willey, 1953; Strong
and Evans, 1952); Salinas, a preceramic site just south of the Chao
Valley (Alva, 1978); the preceramic sites of Huaynuma and Tortugas
(Engel, 1957«, 1957/?) between the Nepeha and Casma Valleys; the
Initial Period site of San Diego (Collier, 1962; Thompson, 1964) near
Puerto Casma in the Casma Valley; the preceramic and Initial Period
site of Las Haldas (Fung, 1969; Engel, 1970; Grieder, 1975; Matsu-
zawa, 1978); the preceramic site of Culebras (Fanning, 1967) at the
mouth of the Culebras Valley; the Initial Period site of Bermejo just
north of the Pativilca Valley (Silva, 1975, 1978); the preceramic site of

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Annals of Carnegie Museum

VOL. 48

Fig. 2. — Map of the north and central Peruvian coast showing the distribution of pre-
ceramic and early ceramic sites mentioned in the text.

Aspero in the Supe Valley (Willey and Corbett, 1954; Moseley and
Willey, 1973; Moseley, 1975; Feldman, 1977); the preceramic site of
Bandurria immediately south of the Huaura Valley, and the preceramic
site of Rio Seco between the Huaura and Chancay Valleys (Wendt,
1964). Additional references to tectonic uplift as far south as Otuma

1979

POZORSKI AND POZORSKI-— -AlTO SaLAVERRY

341

(Craig and Psuty, 1971) and as far north as Talara (Richards and
Broecker, 1963; Richardson, 1974) suggest that the phenomenon may
have affected the entire Peruvian coast. In view of the location and
dating of the early sites mentioned above and the settlement pattern
seen in the Moche Valley, it seems highly likely that this documented
tectonic activity occurred at one time during the early Early Inter-
mediate Period as a major earthquake.

Narrowing our assessment of uplift to the Moche Valley, it is evident
that the associated tectonic movements occurred well after the pre-
ceramic site of Alto Salaverry had been abandoned. An elevation
change of 4 to 6 m or more, exposing areas of submerged land, would
have increased the distance of preexisting land areas from the active
ocean. The earlier beach line would become considerably recessed,
surviving as a terrace. In view of this evidence, it is apparent that the
site of Alto Salaverry, when in use, was only about .4 km from the
sand beach in front of the site and 3 km from the Salaverry Point rock
mass (Fig. 1).

Immediate Vicinity of the Site

The local habitat of Alto Salaverry is a barren rock-strewn sand
desert continuously crossed by active barchan dunes. Slightly more
vegetation may have been present when the site was occupied— plants
such as algarroba {Prosopis chilensis) and achupalla {Tillandsia
species) were once much more common in such marginal areas. Nearer
the river vegetation was increasingly abundant. Such vegetation would
have been sufficient to support a very few small wild animals and
perhaps an occasional deer.

The site lies upon a series of irregular stabilized sand dunes. The
surface is scattered with angular to subangular pebbles and cobbles
except in areas of much dom.estic refuse where it is also littered with
bone, shell, fire-cracked rocks, and an occasional worked stone. Two
of the three complex structures are located atop low rises, probably
old dunes, which have been artificially altered to render the surfaces
suitable for construction (Fig. 3), Exclusively domestic architecture
and the associated refuse are concentrated in lower areas of the south-
central portion of the site. Surface indications of all components are
slight; therefore, extensive excavation was necessary to explore the
architecture and sample the rich midden.

Architecture

As elements of the architecture were cleared, it was evident that the
structures present vary greatly in layout, construction techniques, and
function. The layout of noedomestic structures ranges from a complex
network of rectilinear rooms and platforms to a simple circular pattern,

342

Annals of Carnegie Museum

VOL. 48

Fig. 3. — Site map of Alto Salaverry showing the distribution of structures, burials,
stratigraphic cuts, and midden concentrations.

whereas domestic architecture units are consistently small and simple
isolated or contiguous rooms. Boulders, cobbles, and rectangular
adobes were used in construction. An evaluation of layout, construc-
tion methods and materials, as well as other evidence such as position
within the site and association with refuse form the bases for functional
distinctions between domestic and nondomestic architecture.

Domestic Architecture

Numerous semisubterranean domestic structures are concentrated
in the southwestern portion of the site where refuse is deepest (Fig.
3). A total of nine domestic structures were excavated. The two deeply
buried examples, which were constructed on near-sterile sand, have
walls preserved to a height of over 1 m, whereas the remaining more
superficial structures, built on accumulations of refuse, are often con-
siderably weathered.

Units are generally composed of two or more contiguous rectangular
or semirectangular rooms, each 1.5 to 2 m across. Walls are very thin
(10 to 25 cm), constructed mainly of a single thickness of subangular

1979

POZORSKI AND POZORSKI — AlTO SaLAVERRY

343

Hearths

STRUCTURE A

/

m

Fig. 4.— Plan of Structure A showing two circular hearths set in two small rooms.

cobbles and boulders of hard dark basalt and crumbly gray granite set
in fine sand and silt mortar. In five structures, there are some large
handmade adobes of fine sand and silt, ranging in size from 53 by 25
by 18 cm to 37 by 21 by 18 cm. Some display impressions of finger-
marks as the result of their manufacture. They occur both intermixed
with cobbles and boulders or in rows (up to 7 in a line) laid lengthwise
but never stacked one upon another. Occasionally interspersed among
adobes and stones are chunks of colonial polychaete worm concretions
commonly called piedra poma. Though preservation of walls is some-
times as high as 1 m, the walls still most likely served as footings for
perishable superstructures. Interior room faces and floors are plastered
and well smoothed while outer faces are unfinished.

Structures A and B (Figs. 4 and 5) are the two best examples of
domestic architecture. Structure A (Fig. 4) consists of two complete
rooms and the partial remains of a third. In contrast to the surrounding
midden area, near-sterile clean sand filled the rooms. Each contains
a small circular hearth (10 cm deep and 15 cm in diameter) depressed
into the floor and lined with fine sand and silt now reddened from use.
The rim of each hearth is slightly raised above the floor. The hearth
in the west room lies near the center, whereas the one in the east room
is near the northeast corner. Entrance into the west room is from the
north. The east room presumably had its entrance at the northeast
corner where the wall is only 15 cm high, though, curiously, such an
arrangement would have made it difficult to avoid stepping on the

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Annals of Carnegie Museum

VOL. 48

Fig. 5. — View of Structure B from the north depicting two rooms and the principal step-
through entrance.

hearth. The wall dividing the rooms is 60 cm high measured from the
floor of the east room while it is only 35 cm high measured from the
floor of the west room, revealing a difference of 25 cm between the
floor levels of the two rooms. Scattered on the floor of the east room
were a few shells and sea lion bones; on the floor of the west room
near the hearth was half of a grinding stone or chungo measuring 23
cm wide, 18 cm long (if whole it would have been about 36 cm long)
and 9 cm thick.

Structure B (Figs. 3 and 5) consists of two small rooms connected

on the north by a thick wall or bench to two or three other rooms. The
southernmost two rooms were completely excavated. The east and
larger room was, like the two rooms of Structure A, filled with almost
pure clean windblown sand. The walls are extremely well-preserved
and average 90 cm high with a maximum of 105 cm in the northeast
corner. The interior wall surfaces are plastered with 1 to 2 cm of fine
sand and silt. On the south, east, and north walls, especially near the
northeast corner, are two distinctive silty clay layers, apparently ap-
plied as a type of wall decoration. A solid mass of bright yellow silty
clay once covered the rock and mortar wall interior of the first 67 cm

1979

POZORSKI AND POZORSKI— AlTO SaLAVERRY

345

above the floor. The remaining upper face of the wall, averaging 30 cm
or more, was covered by a solid gray silty clay layer. Although parts
of the colored silty clays were not preserved, it is evident that they
formed two solid masses of color— a narrow gray band situated above

a wider yellow area.

The silty clay prepared floor of the larger room is 2 to 5 cm thick.
A small hearth 10 cm deep and 16 to 22 cm in diameter is located near
the center of the room. Within this hearth were found one Choromy-
tilus chorus shell and 2 limpets {Fissurella species). On the floor be-
tween the hearth and the east wall, in an area 50 cm in diameter,
excavations uncovered 11 flat, round cobbles cracked by heat and one
piece of colonial polychaete worm. The entrance to the room (Fig. 5)
is from the north through a finely plastered step-through doorway 42
cm wide and at least 65 cm high. The base of the door is 33 cm above
the floor of the room. Just to the north or in front of this entrance were
situated at least 10 boulders, some up to 70 cm in diameter, and four
pieces of wood mixed with dark midden. Three of the wood pieces are
flat and unworked, about 70 cm long and from 5 to 9 cm wide, and
still have bark adhering to them. The fourth worked piece is described
below.

The west or smaller room of Structure B is different from the other
well-preserved rooms in both this structure and Structure A, for it was
filled with dark ashy midden. Like the east room, only its interior wall
surfaces are plastered. Originally the room connected with the larger
east room by means of a step-through doorway measuring 46 cm wide
and 39 cm tall, with its bottom 31 cm from the floor of the east room.
At some point, this door was sealed with cobbles and mortar and
plastered to conform with the rest of the wall.

Two more hearths reddened from heat were found during excava-
tions. One is next to Structure C (Fig. 3), a single line of boulders that
was once part of a larger construction. The other hearth was found
isolated 6.5 m W15°S of Structure A. It also was probably situated
within a more perishable structure now destroyed. Both hearths mea-
sure 15 cm in diameter and 10 cm deep. The isolated hearth and the
one in the east room of Structure A were taken to England by the
Archaeomagnetic Expedition 1975 for dating purposes. Results have
not yet been made available.

Semidomestic Architecture

Two standing rectangular complexes (Fig. 3, D and E) are present

on low rises at the east and west edges of the site. The term “semi-
domestic” is applied to these structures because architectural evidence
indicates their function was not exclusively domestic.

Structure D (Figs. 3 and 6) measures 10 m by 10 m and consists of

346

Annals of Carnegie Museum

VOL. 48

0 STRUCTURE D

^ Burial

Fig. 6. — Plan of Structure D which contains three rooms, three platforms, and a burial
within platform 4. Construction of room 3 and platform 4 predates that of platform 5
which in turn predates platform 6. Rooms 1 and 2 postdate room 3 and platform 4, but I
their relationship to platforms 5 and 6 is ambiguous.

three rooms and three platforms. The three rooms are situated in a
row along the east side. Room 1 (Fig. 6) is defined by five large boul-
ders and four cobbles on its north, west, and east sides, none of which
are connected by mortar or fill. The south wall consists of six cobbles
aligned to face north and which are not joined by mortar, but instead,
loose sand. There are two salitrified floor levels, each made of plaster
consisting of fine sand and silt with salt grass. Both are about 10 cm i

thick and separated by a deposit 40 cm thick~=-the upper 15 cm of ]

which is nonashy midden and the rest a mixture of sterile sand and salt J

1979

POZORSKI AND POZORSKI — AlTO SaLAVERRY

347

grass. Both of these floors were cut through by a large pit in the center
of the room which was filled with clean eolian sand.

Room 2 has three sides (north, east, and south) defined by aligned
small boulders and cobbles. The north wall is parallel to the south wall
of room 1, and sand fills the intervening space. The east wall is filled
with fine sand and silt mortar bearing impressions of adobes that were
once present. The south wall is filled with fine sand to silt and salt
grass. An apparent west wall off platform 5 (Fig. 6) is probably actually
a remnant of another platform, for its aligned and plastered face is
toward the west and not east into the room. The floor of the room is
2 cm thick and made of silt.

The west and north walls of room 3 consist of a double line of
boulders and cobbles filled with fine sand to silt and salt grass. The
south wall is also a double line of stones and adobes, filled with dark
midden as a mortar. The east wall is a single line of large cobbles. All
four walls have finely plastered interior faces, similar to the domestic
structures A and B. The northwest corner is curved, whereas the other
corners are square. Most of the material filling the room was clean
eolian sand mixed with some midden material. The floor is similar to
room 2, and like rooms 1 and 2, the walls are usually preserved to a
height of 20 to 30 cm.

The three platforms of Structure D (Fig. 6; 4, 5, and 6) are preserved
up to a height of 1 m. Platforms 4 and 5 are filled with a mixture of
find sand to silt and salt grass and contain no midden. The west and
north retaining walls of 5 and north wall of 4 consist of a single line of
boulders and cobbles aligned and plastered on the exterior. The south
retaining wall of 5 and the south and east retaining walls of 4 consist
of a double row of aligned stones, but are not plastered. Next to the
east wall of platform 4 a burial had been incorporated into the platform
fill during construction.

Platform 6 is of a different nature, for its fill is entirely of dark ashy
midden resembling the refuse that surrounds Structure D. The north,
west, and south retaining walls consist of boulders and cobbles, some
of which are up to 80 cm in diameter, held together by fine sand and
silt mortar. The exterior surfaces of these walls are formed of well-
aligned stones and bear evidence of fine plaster.

Examination of the architectural details of Structure D reveals its
construction sequence. Platform 4 and room 3, built on nearly sterile
sand, are bonded together and represent the first phase of construction.
Platform 5, also built on sand, was then added and plastered. Finally,
platform 6 was constructed on a small amount of refuse which had
accumulated outside platform 5, and its outer faces were also plas-
tered. Rooms 1 and 2 were also added subsequent to the construction

348

Annals of Carnegie Museum

VOL. 48

STRUCTURE E

Fig. 7. — Plan of Structure E containing twenty rooms and two platforms. Stone caches
were found in areas a and b and two whale bones were found in area c. Six construction
phases are present with 1) rooms 1 to 5 built first followed by 2) room 6 and platform
7; then 3) platform 8 and room 9 were constructed followed by 4) rooms 10 to 13, 5) rooms
14 to 17, and finally 6) rooms 18 to 22.

of platform 4 and room 3, and probably predate platform 6 because
they lie on near-sterile sand.

Structure E (Figs. 3 and 7) is larger than Structure D, measuring 30
m by 20 m. Most of the walls are primarily of a single line of boulders
and large cobbles preserved up to 70 cm high, but single rows of rect-
angular adobes (52 by 11 by 12 cm to 35 by 15 by 11 cm) are also
occasionally present. Construction techniques are similar to those of
Structure D with virtually all walls having a single prepared face rec-
ognizable by the alignment of stones and a thin plaster coat. Structure
E consists of 22 units (Fig. 7), all of which are rooms except 7 and 8.
Most floors are merely hard-packed sand, but rooms 4 and 9 have hard
silty clay floors 5 to 8 cm thick. Platforms 7 and 8 contain fill of fine
sand and silt mixed with salt grass and cobbles. Similar salt grass and
cobble fill is evident in the south wall of room 11. Double-faced wall

1979

POZORSKI AND POZORSKI— AlTO SaLAVERRY

349

coEstructioe is present in platform 7 and the north and south walls of
room 1 1 . Platform 8 is not a double-faced construction but rather the
result of expansion of Structure E. Both of its south and west retaining
walls are plastered on their exterior faces.

The distribution of dark midden forms an interesting pattern. Rooms
1 through 5 and most of room 6 and platforms 7 and 8 are built on
clean sand and contain almost no midden. The remainder of Structure
E, the northwest comer of room 6 and rooms 9 through 22, is built on
midden up to 50 cm deep and has refuse in the fill of the rooms and
occasionally even as part of the architecture.

Careful examination of architectural details such as plastered faces
and wall abutments and bonding reveals a minimum of at least six
construction phases: 1) rooms 1 through 5 were built, most likely as
an integrated unit with the main entrance on the south side of room 1 ;

2) the wails forming room 6 and platform 7 were constructed and their
exteriors plastered, with the principal entrance to room 6 on the north;

3) the exterior walls and fill of platform 8 and the south wall of room
9 were added; 4) rooms 10 through 13 were then constructed; followed
by 5) rooms 14 through 17; and finally 6) rooms 18 through 22, all with
an emphasis on plastering of exterior walls.

The presence of some refuse associated with Structures D and E is
evidence of the domestic aspect of their functions. However, the size
and complexity, especially of Structure E, suggest complementary
functions related to social rank and status. Both complexes represent
a compromise between the limitations of construction techniques in
use at Alto Salaverry and the desire for an iiiipressive structure above
ground. As long as wet-laid structures were essentially subterranean,
the fact that only one face was finely finished was unimportant because
only one face, the interior, was ever visible. The initial construction
of architecture on the ground surface apparently necessitated a choice
as to whether the inner or outer wall face should be the finely finished
surface. The examination of both Structures D and E revealed multiple
coostractioii phases, yet with very few exceptions, only the outermost
faces were finely finished. This suggests that the outward appearance
of the structures, especially Structure E, was of special importance
and probably related to the public aspect of their function. Thus, the
inhabitants of the structures, presumably of considerable social status,
would seem to be sacrificing interior refinements in order to present
a more impressive facade to others at the site.

Nondomestic Architecture

The most clearly nondomestic architectural unit is the circular sub-
terranean Structure F in the northern part of the site (Figs. 3 and 8).
There is no refuse in the immediate vicinitv of Structure F, and vir-

350

Annals of Carnegie Museum

VOL. 48

/

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STRUCTURE F

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Fig. 8. — Plan of Structure F showing a stone-lined circular construction built within an
old sand dune. Two step-like entrances are situated opposite each other leading down
into the structure. The center of the floor contains a stone-lined hole which probably

provided footing for a perishable roof support.

tually no artifacts or refuse were encountered during testing within the

structure.

Structure F began as a hole about 9 m across excavated about 1.8

m into the summit of a stabilized sand dune. The hole was lined with

1979

POZORSKI AND POZORSKI — AlTO SaLAVERRY

351

irregular rows of angular boulders held in place by silt and sand mortar
and carefully fitted to obtain a flat interior face. Regularity was sac-
rificed due to size variations in boulders and because the main em-
phasis was on creating a smooth inner face, A pit on the outside of
the base of the hole lining provided evidence for this construction
procedure. Seen from the outside, the lining wall slopes outward, and
the external face is both unfinished and extremely irregular. The floor
was paved with small cobbles covered with up to 5 cm of fine sand
and silt plaster. Traces of a final coat of this plaster still adhere to
portions of the inner wall.

Due to rapid filling by eolian sand. Structure F is preserved almost
intact. On opposite sides from each other are two step-like openings
leading down into the structure. The lowest level of both steps is still
approximately 90 cm above the floor, therefore access was probably
aided by portable steps. A stone-lined hole 40 cm in diameter in the
center of the floor may have provided footing for a perishable roof
support.

On the southeastern side of Structure F is a low wall, one to two
boulders high which once completely encircled the dune summit. Two
ascending steps near the dune top are part of this outer wall. Such an
enclosure may have served to stabilize the artificially altered dune
surface while also facilitating access.

Mortuary Practices

Only two burials were encountered within the site limits, but an
associated preceramic cemetery was discovered only 300 m from the
site. Most evidence of local preceramic mortuary practices comes from
the two burials found within the site which were carefully excavated
and recorded. However, extensive looter action in the nearby ceme-
tery exposed numerous bodies and meager trappings which were also
examined.

Burials within the Site

The first burial encountered was in platform 4 of Structure D, an
area artificially filled with yellow silt and plant fiber (Figs. 3, 6, and 9).
Except for soil discoloration in the immediate vicinity of the body,
there was no evidence of the excavation of a burial pit, indicating the
inhumation must have taken place during platform construction.

Traces of body wrappings were present. The outer covering was of
loosely twined junt o {Cyperus species) matting tied around the body
with thick cotton cord. Inner wrappings consisted of several layers of
twined cotton textiles which were badly decayed from direct contact
with the body. In the vicinity of the skull were pieces of a finely knit
cotton textile, the possible remnants of a cap. Within the bundle, the

Fig. 10.— -Flexed burial of a young child found within a stratigraphic excavation of
midden material. Two boulders were found on top of the body.

1979

POZORSKI AND POZORSKI — AlTO SaLAVERRY

353

Fig. 11. — View from the west of the burial of a young child covered by two boulders,
found within midden material.

body was tightly flexed, lying on its back and left side. The body axis
was oriented north-south with the pelvis toward the north and the head
south. The legs were crossed and drawn up, and both arms were bent
and crossed, placing the hands near the chin. A close examination of
the skeleton indicated a male of medium stature and advanced age.
His teeth were badly worn and decayed, and his spinal column was
severely arthritic.

A second interment was discovered beneath 115 cm of refuse in the
vicinity of domestic architecture, but not associated with any structure
(Figs. 3, 10, and 11). The burial lay at the bottom of a stratigraphic
excavation to investigate the midden, and using profiles drawn to record
refuse strata, the burial pit was easily traced. Although a total of 115
cm of refuse-bearing soil covered the body, the burial pit was dug
much earlier, when only 35 cm of refuse had accumulated. From this
earlier surface, the grave was dug through refuse until sterile sand was
reached.

As sand was brushed away to expose the burial, two small boulders
came into view (Fig. 11), lying directly on the central portion of the
body. They had been placed on the complete burial bundle just before
the grave was closed. An outer twined junco mat, inner layers of

354

Annals of Carnegie Museum

VOL. 48

Fig. 12. — Five pieces of worked wood found during excavations: a-c are from Structure
C, d was found just north of the main entrance to Structure B, and e was discovered
just outside of the east wall of Structure B. These sticks could have been used for prying
shellfish from rocks or as digging tools for plant cultivation.

twined cotton cloth, and a finely knit headpiece also made up the body
wrappings of the second burial.

The body was tightly flexed and lay on its right side (Fig. 10). The
spinal column was oriented north-south with the head south and feet
north. Both legs were tightly drawn up and parallel while the arms
were bent and crossed, bringing the hands to just beneath the chin. On
the basis of dental eruption, the skeleton was determined to be that of
a young child about 10 or 11 years old. Subsequent examination of the
bones revealed that the individual exhibited bone pathologies associ-
ated with anemia, a common condition among prehistoric Peruvian
populations (Trinkaus, 1977).

The Preceramic Cemetery

About 300 m northeast of Alto Salaverry is a Cotton Preceramic
cemetery which contains many burials probably associated with the
occupation of the site. As defined by looting, the cemetery is 8-shaped,
with clusters of burials on top of and at the foot of a small knoll. Within
an area approximately 20 by 40 m, scattered bone fragments are visi-

1979

POZORSKI AND POZORSKI— AlTO SaLAVERRY

355

Fig. 13. — Twenty-seven flat round cobbles from a cache of 37 found next to platform

8 of Structure E. These stones were probably used as hammerstones for pigment pro-
cessing or opening mollusks.

ble, representing 20 to 25 adult individuals. Occasional associations of
single bodies with looters’ pits suggest a general pattern of individual

interments.

The very few grave goods exposed were in a badly weathered con-
dition. These include twined cotton textiles, twined reed matting, and
cotton fishnet. The cotton textiles were especially decayed and were
therefore probably once directly upon the body whereas the mats likely
formed the outer body covering. Finally, the net was probably a burial
accompaniment.

Features common to both burials within the site plus scanty evi-
dence from the nearby cemetery indicate a generalized burial pattern.
Special features, such as burial locations within the site and even with-
in architecture as well as the placement of cobbles over the child’s
body, reflect varied social practices and perhaps status.

Artifacts

Four basic kinds of artifacts were found at the site- — -worked wood,

worked and unworked flat round cobbles, grinding stones, and textiles.

356

Annals of Carnegie Museum

VOL. 48

All of these were found either in the main midden area and nearby
structures or associated with Structure E.

Wooden Artifacts

Five pieces of worked wood were encountered during excavations
(Fig. 12). One (Fig. 12d), found just outside the main entrance to Struc-
ture B, has the shape of a paddle with a tubular shaft and one broad
flat end. The overall length is 45 cm, the shaft diameter 4 cm, and the
flat end 3 cm thick and 6 cm wide. The end opposite the flat ' ‘paddle”
comes to a rounded point. One similar, though smaller, wooden paddle
(Fig. 12e) was found outside of the east wall of Structure B. The re-
maining three worked sticks (Fig. 12a~c) were found next to the small
hearth associated with Structure C. These sticks could have functioned
as prying tools for gathering rock-perching shellfish and/or as digging
implements for plant cultivation.

Stone Artifacts

Unworked flat round cobbles were found scattered throughout much
of the domestic midden and architecture. The majority were cracked
by heat due to exposure to fire, such as those found in Structure B.
However, a cache of 22 unburned flat round cobbles was uncovered
along the north wall of Structure E (Fig. 7a). Each cobble is roughly
the same size (11 cm in diameter), weight (600 g), and color (greenish
gray). Deliberately gathered, they could also have been intended for
use as heating stones for cooking or as hammerstones.

Another cache of flat round cobbles was encountered in Structure
E, next to the east end of platform 8 (Fig. 7b). Thirty-seven were found
together, of which 32 bear marks of use as hammerstones on their
circumferences (27 are illustrated in Fig. 13). All are gray to black in
color and approximately the same size (6 cm in diameter) and weight
(100 g). The pattern of wear varies from a few nicks at opposite ends
to heavy battering along the whole circumference (Fig. 13). Several
have red pigment (probably hematite) on or near their worked surfaces,
which indicates their possible use in pigment processing. Some may
have been used for opening mollusks.

A few other examples of worked cobbles were found in several
places within the western portion of the site. These are roughly the
same size as the previously mentioned unworked cobbles. The only
sign of alteration on each of the stones is a narrow indented groove
that extends partially or fully around the circumference of the center
of the stone. It seems likely that these worked cobbles functioned as
net weights.

A few grinding implements were discovered. In addition to the chun-
go found in Structure B, three more chungos were encountered. One

1979

POZORSKI AND POZORSKI — AlTO SaLAVERRY

357

was along the south side of room 14 of Structure E and measures 43
by 23 by 16 cm. Another, measuring 23 by 18 by 9 cm, was found
within room 19 of Structure E. A third measures 15 by 15 by 8 cm and
was associated with a batan measuring 40 by 20 by 15 cm. Both were
located just outside the northeast corner of room 19. A second batan
(20 by 20 by 12 cm) was encountered on the surface about 10 m south-
east of Structure D.

Textiles

Several types of textiles were encountered during excavations at
Alto Salaverry, the most common type being cotton twined textiles.
Of the 22 examples of twined textiles, ranging in size from a few square
centimeters to 19.5 cm by 20.3 cm, all are weft-twined and all wefts
are 2-ply, Z-spun, and S-twist, twined in a Z manner. All warps on the
other hand are 2-ply, S-spun, and Z-twist. There are six to 12 warps
per centimeter and .4 to 1 cm spacings between the wefts. Where
selvages were preserved, each weft pair terminated in a simple square
or granny knot.

Thread diameters within twined textiles vary from .2 to 1.2 mm.
Individual threads found in excavations reflect the same frequencies
of spinning and twining characteristics as do the warps and wefts with-
in the more complete textiles. Of 67 threads recovered, 56 are 2-ply,
S-spun, and Z-twist, corresponding to warp thread characteristics,
whereas the remaining 11 examples are 2-ply, Z-spun, and S-twist,
corresponding to weft thread characteristics.

Twenty textile examples exhibit weft-twining using exclusively split-
paired warps; of the remaining two specimens, one has nothing but
straight-paired warps, whereas the other has both. This predominance
of split-paired warps suggests a late Cotton Preceramic date according
to the sequence established for the central coast by Moseley and Bar-
rett (1969).

Cotton netting fragments were also found during excavations at Alto
Salaverry. Nine small fragments were collected, all having small mesh
ranging from 5 to 20 mm on a side. The threads are usually 2-ply, S-
spun, and Z-twist. Six netting examples are knotted using 2-element
square knots. The three other examples are composed of successive
threads joined by simple cow hitches.

The third type of cotton textile present at Alto Salaverry is knitting.
Four examples were recovered, all done in a simple looping manner,
only much tighter than that seen in netting.

Four specimens of 4-ply cotton string were excavated at the site.
Two consist of two 2-ply, S-spun, Z-twist threads replied in an S.
manner. The other two are just the opposite: two 2-ply, Z-spun, S-
twist threads replied in a Z manner.

358

Annals of Carnegie Museum

VOL. 48

The only type of noncotton textile is junco iCyperus species) mat-
ting, principally associated with the two burials within the site. In each
case, the matting is constructed of flat whole junco stems used as
warps with split junco fibers as wefts. The junco wefts consist of two
strands of 2-ply, S-spue, Z-twist fibers twined in an S manner, and the
space between the wefts is 6 to 9.5 cm. Nine junco fiber fragments
were found in other areas within the site— five are 2-ply, S-spun, and
Z-twist; three are 2-ply, Z-spue, and S-twist; and one is 3-ply, S-spue,
and Z-twist.

Additional Artifacts

A small sparrow-sized bird enclosed in a beaten plant fiber wrapping
tied with a small 2-ply, S-spue, Z-twist cord was found while clearing
room 1 of Structure D. A small amount of red hematite pigment
wrapped in a twined textile was discovered just outside the southeast
corner of room 10 of Structure E.

The find of two adjacent whale bones in room 6 of Structure E (Fig.

7c and 14) also deserves mention. The two bones are an atlas mea-
suring 87 cm long and an upturned portion of a skull 35 cm across,
possibly part of the occipital. No other artifacts were found near these ^
bones, but their apparent nonalimentary presence in a large clean room [
within the largest structure at the site suggests these bones served in [
a ceremonial capacity.

Subsistence Activities

Ash darkened soil, rich in plant and animal remains, was coecen- i'
trated in the central and southern sectors of the site near the small i
domestic structures. A total of 30 test pits were made within this area [
of refuse to discover its extent, depth, and contents, and to select ■
locations for controlled stratigraphic excavations to secure quaetita- ;■
live subsistence data. The subsistence inventories of the test pits were ^
consistent; therefore, the two locations that were selected for detailed i,
excavation were areas where refuse was deep and varied and where ;
there was no indication of buried architecture (Fig. 3).

A 1 by 2 rn cut was made near the center of the main body of refuse [
(Fig. 3). Subsistence remains were present to a depth of 115 cm, and
the child burial was discovered in the sterile sand beneath this refuse,

A second cut, 1 by 1.75 m, was made into refuse banked against the !
south wall of Structure D. During both cuts, a working face was com- i
pletely exposed to facilitate subsequent excavation by natural levels, j:
Resultant material was passed through a 6 mm mesh screen and all
plant and anim^al remains as well as artifacts were collected. .

Quantitative information about the Alto Salaverry plant and animal
remains is presented in Tables 1 through 4. Columns 1, 2, and 3 are |

1979

POZORSKI AND POZORSKI — AlTO SaLAVERRY

359

Fig. 14. — Two whale bones found in room 6 of Structure E.

simple quantitative assessments of each species. The first column rec-
ords the number of levels in which each species occurred; column 2
records a count for the species; and column 3 gives the weight of the
plant or animal residue collected. The fourth and fifth columns, on the
other hand, present the reconstructed dietary contribution of each
species, first in terms of an absolute volume, and then as a percentage,
by volume, of the total diet. Such reconstructions are always difficult
due to the large number of variables to be considered. The sampling
and analysis of subsistence material, and ultimately the reconstructed
dietary proportions, were based on the assumption that plant and an-
imal remains within an excavated volume occur in frequencies which
are indicative of their importance to the occupants of the site. In keep-
ing with this, a methodology was designed which served to evaluate
as nearly as possible only those remains within the sample volume in
terms of their dietary contribution. This methodology has been de-
scribed in detail elsewhere (S. Pozorski, 1976).

The plant and animal species recovered from Alto Salaverry are
unusually numerous and varied. Predictably, most animal species re-
flect the marine orientation suggested by the site’s proximity to the
ancient beach line. However, the range of plant cultigens is also con-

360

Annals of Carnegie Museum

VOL. 48

Table 1. — Alto Salaverry animal remains from controlled Cut 1.

Species

Levels

where

present

MNI

Weight
in g

Meat
volume
in cm®

Percent of
meat
diet

Mollusks

Scutalus sp.

(land snail)

5

2

+ 1

4.0

+

Drymaleus verillium
(land snail)

1

1

+

+

+

Choromytilus chorus
(purple mussel, choro)

14

54

942.5

2,700.0

17.8

Semimytilus algosus
(thin shelled mussel)

14

1,120

945.0

1,119.5

7.4

Brachidontes purpuratus
(small striated mussel)

13

232

152.5

231.5

1.5

Protothaca thaca
(large clam)

9

1

5.0

10.0

+

Petricola rugosa
(borer)

4

1

2.5

+

+

Mesodesma donacium
(clam)

11

12

35.0

30.0

+

Donax peruvianas
(tide zone clam)

14

394

477.5

197.0

1.3

Fissure Ha sp.

(limpet, barquillo)

14

30

162.5

300.0

2.0

Tegula atra
(gastropod)

14

58

255.0

58.0

+

Turbo niger
(gastropod)

14

71

102.5

71.0

Crepidula dilatata
(slipper shell)

10

16

5.0

20.0

Polinices sp.

(gastropod)

4

3

+

1.5

+

Thais chocolata
(gastropod)

13

10

45.0

20.0

+

Thais delessertiana
(gastropod)

13

27

32.5

40.5

+

Cantharus sp.

(gastropod)

10

14

10.0

+

Nassarius gayi
(gastropod)

8

14

+

=2

_

Concholepas concholepas
(abalone-like gastropod)

10

2

77.5

100.0

Chiton (2 species)

14

14

140.0

140.0

+

Unidentified shell

14

-

217.5

-

-

Crustaceans

Platyanthus orbignii
(purple crab, congrejo)

14

52

215.0

1,025.0

6.7

Balanus tintinnabulum
(barnacle)

14

217

142.5

-

-

1979

POZORSKI AND POZORSKI — AlTO SaLAVERRY

361

Table 1. — Continued.

Species

Levels

where

present MNI

Weight
in g

Meat

volume

in cm®

Percent of
meat

diet

Echinoderms

Tegrapygus niger
(sea urchin, erizo)

4

+

+

+

Fish

Mustelus sp.

(shark, tollo)

11 1

2.5

704.0

4.6

Rhinobatos planiceps
(guitarfish, guitarra)

3 1

12.5

225.0

1.5

Myliobatis peruvianus
(ray, ray a)

5 1

2.5

248.5

1.6

Paralonchurus peruanus
(croaker, roncador)

12 2

7.5

728.5

4.8

Sciaena deliciosa
(croaker, lorna)

14 21

45.0

3,564.0

23.4

Sarda chilensis
(bonito)

1 1

+

106.5

Lepisoma philippi

{trambollo)

4 1

+

90.0

+

Mugil cephalus
(mullet, lisa)

1 1

97.5

+

Unidentified fish

14

90.0

2,250.0

14.8

Birds

Unidentified bird

5

2.5

35.0

+

Mammals

Otaria byronia
(sea lion, lobo del mar)

2 1

65.0

1,086.8

7.1

Unidentified mammal

3

+

+

+

Total

Combined values for percent-
ages less than 1%

15,204.8

94.5

5.5

100.0

* + designates weights less than 2.5 g (Column 3), unreconstructed volumes (Col-
umn 4), and percentages less than \% (Column 5).

^ - is used to indicate ‘ no value."

siderable, especially for a permanent settlement so far removed from
arable land.

Animal Remains

The people of Alto Salaverry were almost entirely dependent on the
nearby ocean for animal protein (Tables 1 and 3). The location of the
site reflects the importance of coastal resources in their diet, and the

362

Annals of Carnegie Museum

VOL. 48

Table 2. — Alto Salaverry plant remains from controlled Cut /.

Species

Levels

where

present

Seed

count

Weight
in g

Food

volume
in cm®

Percent of
plant
diet

Cultivated

Phaseolus lunatus

5

6

+

7.3

+

(lima bean, pallar)
Phaseolus vulgaris (?)

4

3

2.5

1.0

+

(common bean, frijol)
Gossypium barbadense

13

22

50.0

(cotton, algodon)

Capsicum sp.

3

1 stem

+

20.0

+

(pepper, afi)

Cucurbita sp.

11

80

9,000.0

96.1

(squash, calabaza)
Lagenaria siceraria

10

9 stems

27

45.0

(gourd, mate)

Persea americana

2

1

2.5

62.5

+

(avocado, palta)

Bunchosia armeniaca

3

3

-h

30.0

(cansaboca)

Psidium guajava

4

2

+

59.4

+

(guava, guayaba)

Lucuma obovata

3

2

-t-

187.5

2.0

ilucuma)

Wild

Cenchrus echinatus

5

8

+

(burr)

Panicum sp.

10

32.5

(grass, grama)

Gynerium sagittatum

5

.

5.0

_

(cane, cana brava)
Tillandsia sp.

11

25.0

(achupalla)

Prosopis chilensis

5

11

+

ialgorroba)

Mixed fibrous species*

14

=.

312.5

_

_

Unidentified plants

12

-

30.0

-

-

Total

Combined values for per-
centages less than 1%

9,367.7

98.1

1.9

100.0

^ Category includes a mixture of such species as Panicum sp., Gynerium sagittatum,
and Scirpiis tat or a.

species inventory for the site details specific animals upon which ex-
ploitation focussed. Both a sand beach immediately in front of the site
and the rock promontory of Salaverry Point south of the site were
easily accessible.

1979

POZORSKI AND POZORSKI — AlTO SaLAVERRY

363

Table 3. — Alto Salaverry animal remains from controlled Cut 2.

Species

Levels

where

present

MNI

Weight
in g

Meat
volume
in cm®

Percent of
meat
diet

Mollusks

Scutalus sp.

(land snail)

5

1

+

2.0

+

Choromytilus chorus
(purple mussel, choro)

19

81

1,860.0

4,050.0

18.1

Semimytilus algosus
(thin shelled mussel)

18

1,093

1,205.0

1,093.0

4.9

Brachidontes purpuratus

(small striated mussel)

19

340

162.5

340.0

1.5

Protothaca thaca
(large clam)

13

1

2.5

+

+

Eurhomalea rufa
(large clam)

5

1

25.0

+

+

Petricola rugosa
(borer)

9

1

+

+

+

Mesodesma donacium

(clam)

9

5

30.0

22.5

+

Donax peruvianas
(tide zone clam)

19

288

312.5

144.0

Phola chiloensis
(angel wing)

5

1

+

+

+

Fissurella sp.

(limpet, barquillo)

15

40

277.5

400.0

1.8

Tegula atra

(gastropod)

18

27

125.0

27.0

+

Turbo niger
(gastropod)

18

38

77.5

38.0

Crepidula dilatata
(slipper shell)

10

31

32.5

38.8

Polinices sp.

(gastropod)

3

3

+

1.5

+

Thais chocolata
(gastropod)

12

5

47.5

10.0

+

Thais delessertiana
(gastropod)

13

14

15.0

21.0

+

Cantharus sp.

(gastropod)

6

4

7.5

+

+

Nassarius gayi

(gastropod)

7

13

2.5

Mitra orientalis
(gastropod)

2

1

+

0.5

+

Concholepas concholepas
(abalone-like gastropod)

9

4

325.0

200.0

Chiton (2 species)

18

14

100.0

140.0

-h

Unidentified shell

18

-

1,092.5

-

364

Annals of Carnegie Museum

VOL. 48

Table 3). --Continued.

Species

Levels

where

present

MNI

Weight
in g

Meat
volume
in cm®

Percent of
meat
diet

Crustaceans

Piatyanthus orbignii
(purple crab, congrejo)

19

75

260.0

1,500.0

6.7

Balanus tintinnabulum
(barnacle)

17

216

185.0

_

_

Echinoderms

Tetrapygus niger
(sea urchin, erizo)

4

+

+

+

Fish

Must el us sp.

(shark, tolio)

7

1

+

792.0

3.5

Myliobatis peruvianus
(ray, ray a)

3

1

20.0

710.0

3.2

Paralonchurus peruanus

(croaker, roncador)

12

2

10.0

744.0

3.3

Sciaena deliciosa
(croaker, iorna)

16

23

32.5

3,069.0

13.7

Xenoscarus denticulatus
{pococho)

1

1

2.5

120.0

Sarda chiiensis
(bonito)

2

1

+

71.0

+

Lepisoma philippi
{trambollo)

10

1

+

345.0

1.5

Genypterus maculatus
(eel, congrio)

1

1

-i-

43.0

+

Mugil cephalus
(mullet, lisa)

11

2

5.0

1,950.0

8.7

Unidentified fish

19

-

135.0

3,375.0

15.1

Birds

Unidentified bird

3

12.5

175.0

+

Mammals

Otaria byronia

(sea lion, lobo del mar)

4

1

252.5

2,898.0

12.9

Unidentified mammal

3

-

2.5

88.8

+

Total

Combined values for per-
centages less than \%

22,409.1

94.9

5.1

100.0

Shellfish were second only to fish as an animal protein source (Ta-
bles 1 and 3). Species which figure prominently in the refuse include
mussels {Choromytilus chorus, Semimytilus algosus, and Brachi-
dontes purpuratus), gastropods {Teguia atra. Turbo niger, Thais de-

1979

POZORSKI AND POZORSKI — AlTO SaLAVERRY

365

Table 4. — Alto Salaverry plant remains from controlled Cut 2.

Species

Levels

where

present

Seed

count

Weight
in g

Food
volume
in cm^

Percent of
plant
diet

Cultivated

Phaseolus lunatus

1

4

+

5.3

+

(lima bean, pallar)
Phaseolus vulgaris (?)

2

2

+

0.8

+

(common bean, frijol)
Gossypium barbadense

16

44

17.5

(cotton, algodon)

Capsicum sp.

3

3

+

30.0

+

(pepper, afi)

Cucurbita sp.

15

61

2.5

4,500.0

91.9

(squash, calabaza)
Lagenaria siceraria

17

5 stems

15

47.5

(gourd, mate)

Persea americana

5

2

2.5

250.0

5.1

(avocado, palta)

Inga feuillei

1

1

+

2.5

+

ipacae)

Bunchosia armeniaca

5

11

2.5

110.0

2.2

(cansaboca)

Wild

Cenchrus echinatus

5

7

+

(burr)

Panicum sp.

2

52.5

(grass, grama)

Tillandsia sp.

13

37.5

{achupalla)

Prosopis chilensis

6

11

+

{algorroba)

Mixed fibrous species

17

_

197.5

_

_

Unidentified plants

13

-

17.5

-

-

Total

Combined values for

percentages less than 1%

4,898.6

99.2

0.8

100.0

lessertiana , Thais chocolata, and Concholepas concholepas), limpets
{Fissurella honduranensis , F. crassa, and F. peruviana), and chitons
{Mesotomura echinata and Enoplochiton niger). All are types which
require the rocky littoral habitat available at Salaverry Point (Keen,
1971; Olsson, 1961). Clearly, the inhabitants of Alto Salaverry chose
to exploit the more distant, but rich, biomass of the rocky point instead
of the much nearer sand beach.

Evidence in the form of shell breakage patterns reveals systematic
procedures for shell procurement and processing. Mussels and most

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VOL. 48

Fig. 15. — Examples of nicked limpet shells which show evidence of the prying process
used to lift them off their rocky habitat.

gastropods are easily gathered at low tide, but chitons, limpets, and
C. concholepas offer more physical resistance. Regularly occurring
nicks were noted in the same position on whole shells of several
species. On limpets, nicks were found along the narrow end of the
oval shell (Fig. 15). On shells of the abalone-like C. concholepas , nicks !"
were consistently at the point where a hinge-like bulge in the shell
provided additional leverage for prying (Fig. 16). These data reflect a ’
simple but systematic procurement procedure involving the use of a
simple prying tool, such as a pointed stick, to detach certain species. '

Larger shells, especially the large purple mussel (C. chorus), exhibit
a consistent fracture pattern of right angle breaks near the hinge.
Chunks of whorl sections had been battered away from gastropod
shells. Both procedures suggest standard shellfish processing activities
and reveal that the meat was extracted from the live shell by bashing
and not steamed out. Finally, a number of whole or near- whole C. |
chorus shells were worn along the posterior margin, a characteristic ,
which suggests they had been used as simple scrapers.

Over half the total animal protein consumed at Alto Salaverry was
supplied by fish (Tables 1 and 3), and material remains of fishing gear I

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POZORSKI AND POZORSKI— AlTO SaLAVERRY

367

Fig. 16.™ Examples of nicked Concholepas concholepas shells which were damaged
when pried off rocks.

are prominent among the artifacts. Two members of the croaker or
drum family {Paralonchurus pemanus and Sciaena deliciosa) plus
sharks {Mustelus species), rays {Myliobatis peruvianas), and mullet
{Mugil cephalus) supplied most of the fish meat. Small-mesh net frag-
ments and grooved stone net sinkers suggest the use of haul seines to
trap the sharks, rays, guitarfish, mullet, and some croakers as they
fed near shore. Some of the larger bony fishes, mainly croakers, may
have been taken off the Salaverry rock mass which afforded access to
deeper water.

Marine mammals, specifically the sea lion {Otaria byronia), contrib-
uted substantially to the meat protein consumed at Alto Salaverry.
While the proportion of the animals in the diet (about 7%, Tables 1
and 3) falls far short of that of both shellfish and fish, it is sufficient to
indicate that these animals were regularly killed and not fortuitous
finds. The sea lions probably frequented the Salaverry rock mass
where they could be taken by small hunting parties. In addition to sea
lion remains, a few whale bones, probably from animals stranded or
washed ashore, were discovered at the site. The only other mammal
remains were near-whole skeletons of a single viscacha {Lagidium per-
uanum) and a rat.

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VOL. 48

Plant Remains

Food plant evidence from Alto Salaverry reveals that, despite the
site’s location and emphasis on marine resources, cultigens formed a
substantial portion of the total diet. Since many species are consis-
tently underrepresented archaeologically, plants may have been even
more important than the Alto Salaverry data suggest.

The plant cultigens identified at Alto Salaverry, while variable in
abundance, represent a surprising range of species from a Cotton Pre-
ceramic context. Cotton {Gossypium barbadense), gourd {Lagenaria
siceraria), and squash {Curcurbita species), far surpass other species
in abundance (Tables 2 and 4). Other cultivated species include pepper
{Capsicum species), lima bean {Phaseolus lunatus), possible common
bean {Phaseolus vulgaris), pacae {Inga feuillei), lucuma {Lucuma
obovata), guava {Psidium guajava), avocado {Perse a americana), and
the plum-like cansaboca {Bunchosia armeniaca) also called ciruela
delfraile (Towle, 1961:60-61). Cotton, gourd, squash, beans, and pep-
per were probably actually cultivated, whereas the fruits were from
tended plants. All the species were likely brought from the valley bot-
tom where floodplain cultivation was possible and where moisture was
available for wild or tended plants.

In addition to the cultivated species, several wild plants were iden-
tified from Alto Salaverry. A few wild legumes were present (Sagas-
tegui, 1973, personal communication). Tillandsia species, a grass-like
plant with burrs {Cenchrus echinatus), and algarroba {Prosophis chi-
lensis) were moderate in frequency, whereas cane {Gynerium sagit-
tatum) was rare. Because Tillandsia species is an epiphytic plant, de-
pendent only on surface moisture, it was available locally and probably
used as fuel. Algarroba also grows well in marginal areas, which re-
ceive infrequent water, whereas cane would have been available nearer
the Moche River.

Two aspects of the Alto Salaverry plant cultigen inventory are es-
pecially significant. First, the predominance of the industrial plants,
cotton and gourd, relative to other cultigens, is suggestive of a pre-
vailing coastal preceramic attitude toward plant cultivation (S. Pozor-
ski, 1976). To people without pottery and with a marine subsistence
focus, gourd containers and floats and cotton net and cord would have
been extremely important-more important than plant food since an-
imal protein was so readily available. As a result, people initially
viewed plant cultivation essentially as a means for obtaining necessary
raw materials. Such a bias in favor of industrial plant cultivation could
easily have resulted in the neglect and limited production of most food
species.

However, the Alto Salaverry plant inventory is also noteworthy be-
cause of the variety of species present. The amount of care and plan-

1979

POZORSKI AND POZORSKI — AlTO SaLAVERRY

369

ning necessary for such a range of species varies greatly. At the time
when Alto Salaverry was occupied, the agriculturalists were dealing
successfully with: 1) plants such as gourd, squash, and beans which
mature and die each year; 2) perennials such as cotton and pepper
which mature rapidly and continue to produce; and finally 3) perennials
such as lucuma and avocado which mature very slowly, but eventually
produce annually.

How can the presence and apparent cultivation of so many useful
species be reconciled with the continued emphasis on industrial plants
at the expense of these food species? The areal limits of floodplain
land suitable for cultivation may ultimately have been the key variable.
With usable land in short supply and the resulting limited production
concentrating on essential industrial plants, food plant production
would have been secondary. The Alto Salaverry plant inventory sup-
ports this interpretation.

Alto Salaverry in Perspective

Alto Salaverry is a small, though carefully studied, example among
many early sites known on the north Peruvian coast. These sites share
numerous characteristics, especially with respect to subsistence sys-
tems and architectural patterns. The specific data available for Alto
Salaverry add to the collective information on early sites and make
possible further speculation concerning internal features of the site.

Subsistence

Within the north coast region, only three coastal Cotton Preceramic
sites have been described in any detail — Huaca Prieta, Huaca Negra,
and Las Haldas. Huaca Prieta lies at the mouth of the Chicama Valley
near an irregularity in the shoreline where the beach is slightly pro-
tected, Huaca Negra is located close to the modern fishing village of
Guanape near the north margin of the Viru Valley, and Las Haldas is
situated south of the mouth of the Casma Valley near a bay with both
sandy and rocky beach areas.

Huaca Prieta was excavated by Bird in 1946 as part of the Viru
Valley Project because the site was better preserved than early sites
in Viru. Within the refuse. Bird noted considerable numbers of fire-
cracked rocks, and a large amount of plant and animal remains was
recovered during screening of excavated midden. Most of the animal
protein came from fish, shellfish, and birds, while sea lion and porpoise
bone was present, but rare. Bird mentions starfish, sea urchins, crab,
clams, and large purple mussels as part of the shellfish inventory.
Excavations also yielded a nearly complete haul seine with gourd
floats, a net sinker, and a single shell fishhook fragment (Bird,
1948^:302, 19481?: 22-25). Cultivated plant species recovered from Hu-

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Annals of Carnegie Museum

voE. 48

aca Prieta include cotton, gourd, squash, pepper, Canavalia beans,
and lima beans. Tended tree fruits are lucuma, cansaboca (called
ciruela del fraile by Bird, but identified as Bunchosia armeniaca) and
guava (called guayaba by Bird). The roots of wild achira {Canna
eduiis), junco , and cattail were also apparently eaten (Bird, 1948^:302,
1948/?:24; Pickersgill, 1969:55-56; Sauer, 1964; Stephens, 1975; Towle,
1961; Whitaker and Bird, 1949). Although none of the plant species
were quantified, the frequency of cotton twined textiles and gourd
containers and floats confirms the importance of at least these two
cultigens.

At Huaca Negra in Viru, excavated by Strong and Evans in 1946,
few vegetal remains survived due to moist local conditions. Plant
species present include cotton, gourd, and cane^-all identified from
artifacts. Considerable shell and bone were collected from test pits;
and Strong and Evans (1952:17-23) mention clam, snail, large mussel
shells, bird bones, the sea urchin-like pieure, and fish bones which
were mostly from sharks or rays. Subsistence related artifacts include
net fragments, very few shell fishhooks, a grinding stone, and fire-
cracked rocks.

At Las Haldas the preceramic portion is overlain by a much larger
early ceramic component, making assessment difficult. Engel (1970:32)
recorded sea lion bones and legumes, notably lima beans, in the refuse.
Fung (1969:60-62) describes the midden in greater detail. The matrix
is ashy, with many fire-cracked rocks, few marine mollusks and almost
no land snails. The marine species identified include clams {Mesodes-
ma donacium and Eurhomalea rufa), mussels {Aulocomya ater and
Brachidontes purpuratus), gastropods {Teguia atra, Concholepas con-
cholepas, and Oliva species), limpets {Fissurella maxima, F. crassa,
and others). The presence of mammal, bird, and small-fish bones is
also documented. Gourd, cotton, and Tillandsia are the only plant
species recorded by Fung. One possible grinding stone is also de-
scribed.

The lack of quantitative subsistence information from other prece-
ramic sites makes direct comparisons with Alto Salaverry difficult,
although the data available suggest a pattern of similar marine-oriented
procurement systems. All the sites studied in detail, especially Alto
Salaverry and Las Haldas, as well as most other preceramic coastal
settlements, are located well away from river areas where floodplain
cultivation is feasible. This emphasis on marine resources at the ex-
pense of agriculture is further reinforced by both the predominance of
industrial plants useful for marine exploitation and the faunal inventory
at each site. Common technological aspects such as the use of haul
seines for near shore fishing and heated rocks for cooking point to
standard exploitation and processing procedures. However, on a very

1979

POZORSKI AND POZORSKI — AlTO SaLAVERRY

371

local level, differences in the species inventories for each site reveal

a degree of specialized adaptation to specific coastal situations.

Architecture

Preceramic domestic architecture is little known archaeologically.

Recently there has been considerable interest in early nondomestic
architecture, resulting in considerably more data about this aspect of

early settlements. However, most of this information comes from sur-
vey rather than excavation.

Preceramic domestic architecture was encountered in the excava-
tions by Bird at Huaca Prieta and Strong and Evans at Huaca Negra.
Huaca Prieta is a single oval midden 125 by 150 m by about 12 m high.
Within the ashy refuse matrix. Bird encountered both retaining walls
and a series of subterranean structures built of rounded beach cobbles
(Bird, 1948/7:22-23). At the site of Huaca Negra, a low circular mound
about 300 m in diameter. Strong and Evans (1952) report semisubter-
ranean structures with plastered clay walls within the refuse matrix.
Based on these descriptions, the domestic architecture of Alto Sala-
verry would appear to be consistent with the nearby known examples.

The semidomestic structures at Alto Salaverry are probably most
comparable to the low mounds described for several preceramic sites
south of the Moche Valley. Almost adjacent to the Pan American
highway in the vicinity of Culebras is a mound which may also be
preceramic. It is slightly set apart from the main body of refuse. Eur-
ther south, the site of Aspero near Supe contains a series of six large
mounds in association with areas of very deep refuse and considerable
domestic architecture. Excavations in several of the mounds revealed
architectural combinations of walls and platforms which had been re-
peatedly reconstructed (Robert Eeldman, personal communication;
Moseley, 1975). Piedra Parada, also in Supe, has but a single mound
(Robert Eeldman, personal communication; Moseley, 1975). Bandur-
ria, south of the Huaura Valley, consists of several artificial mounds
associated with deep refuse some distance away. Rio Seco, just north
of the Chancay Valley, is characterized by six such mounds closely
associated with refuse (Wendt, 1964). Excavations into the Rio Seco
mounds also revealed buried architecture — walls, filled platforms, and
often associated refuse. The mounds described for other preceramic
coastal sites and the complex structures above ground at Alto Salav-
erry are similar only in a very general way. Excavation data available
from Aspero and Rio Seco reveal that such large artificial mounds
represent much greater labor investments as well as technologically
more sophisticated construction techniques than were documented for
Alto Salaverry. Nevertheless, the public nature of Structures D and
E at Alto Salaverry would indicate that their function within the site

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may be analogous to that of early nondomestic and monumental struc- |
tures elsewhere. '

Architectural forms similar to the nondomestic or ceremonial cir- I
cular Structure F discovered at Alto Salaverry are known from only
two other preceramic sites, although the form becomes widespread on |
the coast in early ceramic times. Several examples are visible in the
air photographs published by Kosok (1965:194, 223, 225), but the first
study of this architectural form was published by Williams (1972). Both j
preceramic examples lie south of the Moche Valley. The Universidad '
Catolica survey of the Chao Valley recently discovered a preceramic
site called Salinas near the salt works south of Chao which has two
circular sunken structures. Considerably further south, near the mouth ‘
of the Supe Valley, the site of Piedra Parada contains a single circular
structure (Robert Feldman, personal communication). [

Excavations at Salinas revealed architectural details of the two cir- |
cular structures which are similar to features of the Alto Salaverry
example (Alva, 1978). Though larger and more elaborate with respect 1;
to the stone construction, the Salinas circular structures are quite sim- :
ilar in plan, with each having two opposing step-down entrances and ^
one having a pit in the center of the circular floor. However, at both
Salinas and Piedra Parada, the sunken circular structures are associ- i
ated with a mound facing toward the circle, a feature which sets the |!
Alto Salaverry circle apart from the known examples.

Circular sunken courts are common in early ceramic times, and all
known examples are also associated with mounds. These include a site ||
near Tanguche in the Santa Valley (Kosok, 1965:194); Taukachi, Kon- I
kan, Pallka, and Sechin Alto in the Casma Valley; Las Haldas south
of the Casma Valley, Bermejo north of the Patavilca Valley (Silva, ||
1975, 1978); numerous sites in the Patavilca-Fortaleza-Supe complex
such as Chupa Cigarro Grande (Kosok, 1965:223, 225; Robert Feld-
man, personal communication); and Garagay as far south as the Rimac
Valley (Ravines, 1975:197; Ravines and Isbell, 1975). Another example
was discovered at the sierra site of Chavin de Huantar (Lumbreras, li
1977). ‘

Summary and Conclusions 1

Alto Salaverry can be evaluated in terms of its place in north coast
prehistory on the basis of both data available from excavations within
the site and similarities shared with other sites along the coast. First, '
examining the site in detail makes one aware of both a complex sub- [
sistence system and an elementary ranked social structure. The plant |
and animal remains collected at Alto Salaverry document a mainly
marine subsistence orientation in conjunction with the knowledge and ,
use of a great variety of plant species. On the basis of such data the i

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373

people of Alto Salaverry are seen as agriculturally fully adapted to
make use of most cultivated and tended species, but still circumscribed
by the spatial limits of areas suitable for floodplain cultivation. Con-
sequently, the marine subsistence focus correlated with the site loca-
tion and the restricted land area available are seen as the key variables
in determining that production of industrial crops is high at the expense
of most food species.

Also within the site of Alto Salaverry, certain characteristics provide
information about the local social structure. On the basis of size, con-
struction techniques, general configuration and association with ref-
use, the types of architecture present at the site are ranked on a con-
tinuum from purely domestic through semidomestic to clearly
nondomestic. Although there is evidence for sequential construction
within the domestic and semidomestic sectors, neither the midden con-
tents nor associated artifacts indicate great time depth within the site.
The two structures designated as semidomestic architecture provide
the main evidence of elementary social ranking within the site. In the
case of both Structures D and E, the association of refuse with portions
of the structure confirm a domestic aspect of the function of each.
Additionally, most of each complex is comprised of rooms and plat-
forms without refuse which architecturally emphasize exterior finish-
ing, thereby suggesting a second public, or nondomestic function. In
contrast, the domestic structures are small, simple, and closely asso-
ciated with the main body of midden at the site. Such differences
between the complexes and the domestic structures suggest status
inequalities between the inhabitants of each architectural type. The
burial of an older adult incorporated in semidomestic architecture and
a child in the main refuse within the site whereas other adults are
placed in a nearby cemetery may also be correlated with differential
individual status. Certainly all these differences are slight and probably
reflect no more than a simple chiefdom level of development.

When Alto Salaverry is compared to other early coastal sites, two
aspects seem especially significant. First, with respect to both general
subsistence activities and domestic architectural forms, Alto Salaverry
conforms well to data available for the north coast both north and
south of the site. However, the presence of semidomestic architecture
and especially the circular structure makes Alto Salaverry distinct
from sites in the immediate vicinity and suggests ties with settlement
types only known further south. On the basis of this, Alto Salaverry
is seen as generally more similar to preceramic sites further south
which are characterized by nondomestic mounds and circular struc-
tures, and inconsistencies are viewed as a result of the peripheral lo-
cation of the site. This is not a reflection of far-reaching suprasocietal
linkages as early as preceramic times, but more likely the result of a

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Annals of Carnegie Museum

VOL. 48

gradual diffusion of a social system or set of beliefs manifest architec-
turally as a recognizable pattern.

Acknowledgments

I

The site of Alto Salaverry was discovered, surveyed, and recorded in 1970 by Michael j
E. Moseley as part of the Chan Chan-Moche Valley Project settlement pattern survey, j'
The associated cemetery, discovered somewhat later, was surveyed and recorded in i
1973 by Eric E. Deeds. Excavations at the site were conducted in May 1974 under the |
supervision of Thomas G. Pozorski and Shelia G. Pozorski and with the permission of i
the Instituto Nacional de Cultura granted to the Chan Chan-Moche Valley Project. |
Drafting was done by Eduardo Martell. Funding was provided by the National Science
Foundation, the National Geographic Society, and the Institute of Latin American Stud-
ies of the University of Texas. The authors gratefully acknowledge the contributions of |
all these individuals and institutions.

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i!

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cultura, Actas y Trabajos del III Congreso Peruano, 1:275-276. I

Bird, J. B. 194&g America's oldest farmers. Nat. Hist., 57:296-303, 334-335. j

. 1948^. Preceramic cultures in Chicama and Viru, in A reappraisal of Peruvian ji

archaeology. Mem., Soc. Amer. Arch., 4:21-28. |i

Collier, D. 1962. Archaeological investigations in the Casma Valley, Peru. 34th In- ::

ternat. Congress Americanists, 1:411-417. [

Craig, A. K., and N. P. Psuty. 1971. Paleoecology of shell mounds at Otuma, Peru. !|
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Donnan, C. B., AND C. J. Mackey. 1978. Ancient burial patterns of the Moche Valley,
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Engel, F. 1957^1. Early sites on the Peruvian coast. Southwestern J. Anthro., 13:
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. 19576. Sites et etablissments sans ceramique de la cote peruvienne. Journal de i

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. 1970. Las lomas de Iguanil y el complejo de Haldas. Departamento de Publi- ji

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Feldman, R. A. 1977. Life in ancient Peru. Field Mus. Nat. Hist. Bull., 48(6): 12-17. i;
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Grieder, T. 1975. A dated sequence of building and pottery at Las Haldas. Nawpa !

Pacha, 13:99-112. i|

Keen, A. M. 1971. Sea shells of tropical West America. Stanford Univ. Press, Stan- i
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Kosok, P. 1965. Life, land and water in ancient Peru. Long Island Univ. Press, New 1;

York, 264 pp. |

Lanning, E. P. 1967. Peru before the Incas. Prentice-Hall, New Jersey, 216 pp. jl
Lumbreras, L. G. 1977. Excavaciones en el templo antiguo de Chavi'n (sector R); I
informe de la sexta campaha. Nawpa Pacha, 15:1-38. j

Matsuzawa, T. 1978. The Formative site of Las Haldas, Peru: architecture, chro- ||
nology, and economy. Amer. Antiquity, 43:652-673. ji

Moseley, M. E. 1975. The maritime foundations of Andean civilization. Cummings !

Publ. Co., Menlo Park, California, 131 pp. I

Moseley, M. E., and L. K. Barrett. 1969. Change in Preceramic twined textiles |
from the central Peruvian coast. Amer. Antiquity, 34:162-165. ]

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Moseley, M. E., and G. R. Willey. 1973. Aspero, Peru: a reexamination of the site

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Olsson, a. a. 1961. Mollusks of the tropical Eastern Pacific: Panamic-Pacific Pele-
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PiCKERSGiLL, B. 1969. The archaeological record of chili peppers {Capsicum) and the
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PozoRSKi, S. G. 1976. Prehistoric subsistence patterns and site economics in the Moche
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Ravines, R. 1975. Garagay: un viejo templo en los Andes. Revista del Instituto Na-
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Richards, H. G., and W. Broecker, 1963. Emerged Holocene South American
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Richardson, J. B., III. 1974. Holocene beach ridges between the Chira River and
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Sagastegui A., A. 1973. Manual de las malezas de la costa norperuana. Universidad
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Sauer, J. 1964. Revision of Canavalia. Brittonia, 16:106-181.

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Actas y Trabajos del III Congreso Peruano, 1:310-325.

Stephens, S. G. 1975. A reexamination of the cotton remains from Huaca Prieta, north
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ANNALS

of CARNEGIE MUSEUM

CARNEGIE MUSEUM OF NATURAL HISTORY

4400 FORBES AVENUE « PITTSBURGH, PENNSYLVANIA 15213
I VOLUME 48 30 NOVEMBER 1979 ARTICLE 20

PALEONTOLOGY AND GEOLOGY OF THE BADWATER
CREEK AREA, CENTRAL WYOMING. PART 18.
REVISION OF LATE EOCENE HYOPSODUS

Leonard Krishtalka

Assistant Curator, Section of Vertebrate Fossils

Abstract

I

F

Review of late Eocene Hyopsodus material, including a larger sample from Badwater,
indicates that Hyopsodus fastigatus Russell and Wickenden, 1933, is conspecific with
Hyopsodus uintensis Osborn, 1902. Material from Montana previously referred to H.
fastigatus represents a new species of Hyopsodus .

Introduction

In the last major review of late Eocene Hyopsodus, Gazin (1968)
recognized two species—//, uintensis Osborn, 1902, and H . fastigatus
Russell and Wickenden, 1933— from sparse remains in Wyoming,
Utah, Montana and Saskatchewan. Additional collecting from the se-
ries of Uintan Badwater localities has since enhanced this record.
Analysis of these and other remains warrants revision of the system-
atics of late Eocene Hyopsodus . Also, the absence of a record of
Hyopsodus from the Duchesnean Badwater locality 2(1 — one of the
best sampled of the Badwater sites— has significant paleoecological
implications.

Osborn (1902) described H. uintensis from the holotype, a partial
maxilla (AMNH 2079), and two fragmentary lower jaws (AMNH 2078,
2078a), all from Uinta C deposits, Utah. Gazin (1956, 1968) referred
to this species material from the Badwater localities 5, 5A, 5 Front, 5
Back, 6, and 7, and six unnumbered specimens from the Uinta Basin
in the CMNH, USNM, YPM, and MCZ collections. Of the latter, only

Submitted for publication 20 April 1979.

377

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Annals of Carnegie Museum

VOL. 48

the CM material is definitely known to have been recovered from the
Uinta B White River Pocket locality. The provenance of the unnum-
bered Hyopsodus remains in the MCZ and USNM is probably the
same, but the YPM material may be from either Uinta B or Uinta C
sediments. Material identified as Hyopsodus cf. uintensis includes an !
isolated molar from the Beaver Divide Conglomerate (Gazin, 1956;
Van Houten, 1964; Emry, 1975) and beautifully preserved dentitions
from the Tepee Trail Formation, East Fork Basin, Wyoming (Mc-
Kenna, 1972).

Russell and Wickenden (1933) identified a larger species, H. fasti-
gatus, from four isolated lower molars from the Swift Current Creek
beds, Saskatchewan. Later, Russell (1965) added four more isolated
molars from the same deposits to the hypodigm and assigned three
teeth to Epihippus? sp. Gazin (1968) referred to H. fastigatus more
complete material from the Shoddy Springs locality. Climbing Arrow
Formation, Montana, in the CMNH collections.

Analysis of this material, remains of Bridgerian Hyopsodus , as well
as new collections from the Badwater Uintan deposits, implies that (1)
in Russell’s study (1965), one of the teeth identified as H . fastigatus
does not belong to Hyopsodus , and two of the three alleged Epihippusl
sp. teeth are upper molars of Hyopsodus’, (2) the holotype and referred
material of H . fastigatus from the Swift Current Creek beds are con-
specific with H. uintensis’, (3) the referred material of H. fastigatus
from Montana represents a new species; and (4) collections of Hyop-
sodus from the Uinta B White River Pocket locality belong to H.
paulus, hitherto known only from Bridgerian horizons.

Abbreviations used are as follows: AMNH, American Museum of Natural History;
CMNH(CM), Carnegie Museum of Natural History; NMC, National Museum of Can-
ada; ROM, Royal Ontario Museum; MCZ, Museum of Comparative Zoology, Harvard
University; USNM, Smithsonian Institution; YPM, Yale Peabody Museum, Yale Uni-
versity; L, length; W, width; N, number; SD, standard deviation; CV, coefficient of
variation. All measurements in text and tables are in millimeters.

Systematics

Hyopsodus Leidy, 1870
Hyopsodus uintensis Osborn, 1902
(Figs. 1™5, Tables 1, 2)

Hyopsodus fastigatus Russell and Wickenden, 1933.

Hyopsodus , cf. uintensis Gazin, 1956.

Hyopsodus sp. Van Houten, 1964.

Hyopsodus fastigatus Russell, 1965 (in part).

Epihippusl sp. Russell, 1965 (in part).

Hyopsodus uintensis Gazin, 1968 (in part).

Hyopsodus cf. H. uintensis Emry, 1975.

Holotype. — ^AMNH 2079, partial right maxilla with P^-M^, Uinta
Formation (Uinta C), Utah.

1979

Krishtalka — Badwater Hyopsodus

379

Fig. 1. — Hyopsodus uintensis, type, AMNH 2079 (part), RPM^ part of

Referred specimens. AMNH 2078; P4-M,, AMNH 2078a, CM 14576; M,_.>,
CM 14418, 16860; CM 16054; P4, CM 14520, 28864, 29210; M, , CM 15250, 15256,
15252, 14574, 14521, i5251, 15265; M.,, CM 25325, 16816, 14523, 15253, 15246, 16857,
15262, 15264, 15263, ROM 1682, 1683, NMC 8654, 8655; M3, CM 15247; p■'-^ CM
28865, 28889,; M^-^, CM 14419, 21985; CM 18251; P, CM 29209; dP, CM 14524,
15254, 28843; M‘, CM 14458, 14519, 29026, 14752, 15248, 15249, 15255, 15257, 15260,
15261, 16858, 25323, 28840; USNM 21089, ROM 1681, 1686, 1687; M\ CM 14514, 14515,
14517, 14518, 14575, 15556, 16859, 25324, 28838, 28839, USNM 23743, 181389; M^,
CM 14753, 15259, 16856, 19739, 25328, 28841, 28842, 28888, 29027.

Localities. — Badwater localities 5, 5A, 5 Front, 5 Back, 6, 7, Hendry Ranch deposits.

380

Annals of Carnegie Museum

VOL. 48

Fig. l.—Hyopsodus uintensis, type, AMNH 2079 (part),

Wyoming; Beaver Divide Conglomerate Member, Wiggins Formation, Wyoming; Swift
Current Creek beds, Saskatchewan; Uinta Formation, Utah.

Known distribution.— AJintdin of Wyoming, Utah, and Saskatche-
wan.

Emended diagnosis. — Differs from all previously described species
of Hyopsodus as follows: more molariform, with wider trigonid

Krishtalka — Badwater Hyopsodus

Figs. 3-4.~Hyopsodus uintensis. 3) CM 28889, 4) CM 14418, LM,_2.

382

Annals of Carnegie Museum

VOL. 48

Fig, 5. — Hyopsodus uintensis, AMNH 2078, LP;j-M,.

and talonid; on trigonid oriented more obliquely, entoconid

greatly enlarged, hypoconulid reduced, valley between these two
cusps obliterated; with broader postcingulum lingually, stronger
preprotocrista; paraconule on P; with hypocone almost as large
as protocone; deep lingual valley completely separating two cusps and
extending labially to metaconule; with wider hypoconal shelf;
marked lophodonty on the molars. Significantly smaller than H. sho~
iemi, new species (described below), with triangular shorter tri-
gonid on P3_4, narrower talonid on P4 and more transverse M^““.

Remarks .—H . uintensis is derived with respect to all previously
described species of Hyopsodus in the degree of molarization of the
premolars, the degree of lophodonty on the molars, the deep valley
separating the large hypocone and protocone on M^~“, and the en-
largement of the entoconid on

1979

Krishtalka— -Badwater Hyopsodus

383

Table Dimensions of upper teeth o/ Hyopsodus uintensis /rom Wyoming, Utah and
Saskatchewan, and H, sholemi new species, from Montana.

Teeth

statistics

Hyopsodus uintensis

Hyopsodus sholemi

L

w

L

w

F Range

3.0-3 J

5.1

3.4

5.6

Mean

3.066

5.1

—

Number

3

2

1

1

Range

4.0-4.6

4.9-5. 7

5.0-5. 1

6.4-6.5

Mean

4.368

5.363

5.066

6.466

Number

19

19

3

3

SD

0.1916

0.2564

CV

4.386

4.781

—

—

Range

4. 1-4.9

5. 7-6.6

5.0-5. 1

7. 1-7.5

Mean

4.538

6.213

5.066

7.333

Number

16

16

3^

3

SD

0.2629

0.3052

___

CV

5.793

4.912

—

—

Range

3. 5-3. 8

4.5-5.5

3. 6-4.4

6.0-6.8

Mean

3.656

5.191

4.025

6.325

Number

9

11

4

4

SD

0.1236

0.3207

CV

3.380

6.170

West (1979) has convincingly shown that Bridgeriae material of
Hyopsodus from the Green River Basin represents only three species,
one of which is H. paulus. In these and known Wasatchian species of
Hyopsodus, molar lophodonty is absent or weak, the hypocone on
is small, the protocoee^hypocoee valley is shallow so that a strong
labial connection between the two cusps is maintained, and the ento-
conid on Mj„3 is not enlarged.

Accordingly, CM 17150, the (previously unnumbered) partial maxilla
from the Uinta B White River Pocket locality that Gazin (1968) iden-
tified as H. uintensis is referred to H, paulus, along with CM 14924,
19648, 19947, and 19948, additional material recovered from the same
locality.

Similarly, the unnumbered specimens in the MCZ and YPM from
the Uinta Basin that Gazin (1968) assigned to H, uintensis are iden-
tified here as H. paulus. There is no record of Hyopsodus material
from the Uinta Basin in the USNM collections.

USNM 181398 from Beaver Divide is, as Gazin (1956) and Emry
(1975) tentatively noted, an M^ of H. uintensis.

Russell’s (1965; Russell and Wickeedee, 1933) recognition of H.
fastigatus from isolated lower molars stemmed partly from his stated

384

Annals of Carnegie Museum

VOL. 48

Table 2, — Dimensions of lower teeth o/ Hyopsodus uintensis from Wyoming,
Saskatchewan, and H. sholemi new species, from Montana.

Utah and

Teeth

statistics

Hyopsudus uintensis

Hyopsodus sholemi

L

w

L

w

P4 Range

3. 3-3. 6

2. 5-3. 2

4.2

3.3

Mean

3.5

2.817

— .

—

Number

5

6

1

1

SD

0.1124

0.2316

_

cv

3.497

8.221

—

—

Ml Range

4. 2-4. 8

3.0-3. 8

— .

3.8

Mean

4.5

3.318

—

—

Number

12

11

_

1

SD

0.1906

0.2135

_

__

CV

4.235

6.434

—

—

M2 Range

4.9-5.4

3. 2-3.9

5.5

4.3

Mean

5.04

3.633

—

— .

Number

15

15

1

1

SD

0.1505

0.2058

—

—

CV

2.987

5.666

—

—

M3 Range

5. 4-5. 6

3. 3-3. 5

—

—

Mean

5.5

3.4

—

Number

2

2

—

Holotype (CM 18851).

absence of a record of the lower dentition of H. ulntensis. Osborn
(1902) had, however, referred two lower jaws to H. uintensis along
with the type partial maxilla. Although Russell (1965) correctly noted
that a newly recovered of H . fastigatus (ROM 1681) was indistim
guishable from of H. uintensis, he and Gazin (1968) maintained H.
fastigatus for the Swift Current Creek material, citing the larger size
of the isolated lower molars in comparison with those of H. uintensis
from Badwater. Analysis of the greater sample of H. uintensis from
Badwater and remeasurement of the Swift Current Creek material in-
dicates that the type and referred specimens of H. fastigatus (except
ROM 1684) from Saskatchewan do not differ significantly in size or
crown morphology from H. uintensis. Accordingly, H. fastigatus is
synonymized here with H. uintensis. Additionally, two (ROM 1686,
1687) of the three upper molars that Russell (1965) identified as Epi~
hippus? sp. represent H . uintensis. ROM 1684 is not a molar of Hyop-
sodus.

The material from the Shoddy Springs locality that Gazin (1968)
assigned to H. fastigatus is, as described below, distinct from H.
uintensis, and referred to a new species. The Hyopsodus remains from

1979

Krishtalka — Badwater Hyopsodus

385

386

Annals of Carnegie Museum

VOL. 48

the East Fork Basin (McKenna, 1972) are under study elsewhere and
not considered here.

Hyopsodus sholemi, new species
(Figs. 6=8; Tables 1, 2)

Hyopsodus fastigatus Gazin, 1968 (in part).

Holotype. — CM 18851, partial left dentary with P:rM2, from Shoddy
Springs locality. Climbing Arrow Formation, Montana.

1979

Krishtalka — Badwater Hyopsodus

387

Fig. ^.—Hyopsodus sholemi, USNM 23744 (part); a) b)

388

Annals of Carnegie Museum

VOL. 48 *

Referred specimens. — Mj or Mg, CM 21257; P^-M^, USNM 23744; M^“^, CM 18849,
18850; Mf CM 21258.

Locality. — Shoddy Springs, Climbing Arrow Formation, Montana (coordinates on file
in the Section of Vertebrate Fossils, CMNH).

Known distribution. — Duchesnean of Montana. |

Diagnosis .--LdiVgQsi known species of Hyopsodus; most closely re- j
sembles H. uintensis in morphology of differs from H. uin-

tensis as follows: Pg more elongate, with stronger paracristed and def- j
inite metaconid; P4 with broader talonid, especially at the lingual part j
of the base of the metaconid; P^ quadrate (rather than triangular), more
molariform, with expanded, squared-off, posterointernal corner of the
crown; more nearly square, with larger L/W ratio. i'

Etymology . — Named for Mr. Sholem Krishtalka.

Remarks. — -Like H. uintensis, the molars of H. sholemi differ from |i
all other species of Hyopsodus in exhibiting marked lophodonty, '
oblique skewing of the trigonid, an enlarged entoconid, and a deep 1
valley that completely separates the protocone and large hypocone. •
Compared to H. uintensis, Pg^-Mg^ of H. sholemi are significantly |
larger, P^ is squared-off lingually by an expansion of the posterointer- i:
nal corner of the crown, and Pg_4 are more molariform. |

H. sholemi is known only from Shoddy Springs, Montana, from ■
material previously (Gazin, 1968) identified as H. fastigatus. The ho- :
lotype and remaining specimens of H. fastigatus from the Swift Cur- [
rent Creek beds, Saskatchewan, were referred above to H. uintensis. |

ij

Conclusions i'

The Badwater Uintan localities preserve remains of a single species j:
of Hyopsodus , H. uintensis, the type of which is from Uinta C de- '
posits, Utah. Material from the Uinta B White River Pocket locality p
previously described as H. uintensis shares the diagnostic features of
and is referred to H. paulus, a species formerly known only from i
Bridgerian sediments. H. fastigatus, described from Saskatchewan li
and Montana, is not valid. The Swift Current Creek material, including ■
the holotype, belongs to H. uintensis. The remains from the Climbing
Arrow Formation compose the hypodigm of a new species, H. sho- !|
lemi. [

H. uintensis and H. sholemi are derived compared to all other |
known species of Hyopsodus— are more molariform, and the |
molars extremely lophodont. Additional specializations are a large hy-
pocone and deep hypocone-protocone valley on M^-^, and an oblique- [
ly oriented trigonid on M^.g. H. sholemi is significantly larger than H.
uintensis, with a more molariform P^ and P3_4. |

H. uintensis is currently known from late Uintan localities in Wy- ji

1979

KrISHTALKA — BaDWATER //F0«'0/)(75

389

oming, Utah, and Saskatchewan. H. sholemi, from Montana, may be
Duchesnean. Other elements of the Shoddy Springs fauna imply an
age comparable to that of the Duchesnean Badwater locality 20 (Krish-
talka and Black, 1975) which has been dated at 41 my. The absence
of Hyopsodus from locality 20 has paleoecological implications that
will be discussed elsewhere, in a final review of the Badwater fauna.

Acknowledgments

Drs. Mary R. Dawson and Robert M. West provided helpful discussions and reviewed
the manuscript. Dr. West also greatly facilitated comparisons between Bridgerian and
Uintan material of Hyopsodus . Dr. Malcolm C. McKenna kindly made available spec-
imens of Hyopsodus in the American Museum of Natural History, including undescribed
material from the Tepee Trail Formation. This study was supported by NSF grants GB-
1266, GB-4089, GB-7081, GB-30840X, and DEB-76-18760.

Literature Cited

Emry, R. J. 1975. Revised Tertiary stratigraphy and paleontology of the western Bea-
ver Divide, Fremont County, Wyoming. Smithsonian Contrib. Paleobiol., 25:1-20.
Gazin, C. L. 1956. The geology and vertebrate paleontology of Upper Eocene strata
in the northeastern part of the Wind River Basin, Wyoming. Part 2. The mammalian
fauna of the Badwater area. Smithsonian Misc. Coll., 131:1-35.

— — 1968. A study of the Eocene condyiarthran mammal Hyopsodus . Smithsonian
Misc. Coll., 153:1-90.

Krishtalka, L., and C. C. Black. 1975. Paleontology and geology of the Badwater
Creek area, central Wyoming. Part 12. Description and review of late Eocene Mul-
tituberculata from Wyoming and Montana. Ann. Carnegie Mus., 45:287-297.
McKenna, M. C. 1972. Vertebrate paleontology of the Togwogtee Pass area, north-
western Wyoming. Pp. 80-101, in Guidebook — Field Conference on Tertiary Bio-
stratigraphy of Southern and Western Wyoming (R. M. West, ed.).

Osborn, H. F. 1902. American Eocene primates, and supposed rodent family Mixo-
dectidae. Bull. Amer. Mus. Nat. Hist,, 16:169-214.

Russell, L. S. 1965. Tertiary mammals of Saskatchewan, Part 1: The Eocene fauna.

Royal Ontario Mus., Life Sci. Contrib., 67:1-33.

Russell, L. S., and R. T. D. Wickenden. 1933. An Upper Eocene vertebrate fauna
from Saskatchewan. Trans. Royal Soc. Canada, ser. 3, sect. 4, 27:53-65.

Van Houten, F. B. 1964. Tertiary geology of the Beaver Rim area, Fremont and
Natrona Counties, Wyoming. U.S. G. S. Bull., 1164:1-99.

West, R. M. 1979. Paleontology and geology of the Bridger Formation, southern Green
River Basin, southwestern Wyoming. Part 4. Notes on Hyopsodus . Contrib. Biol.
Geol., Milwaukee Publ. Mus., in press.

Back issues of many Annals of Carnegie Museum articles are
available, and a few early complete volumes and parts are listed
at half price. Orders and inquiries should be addressed to:
Publications Secretary, Carnegie Museum, 4400 Forbes Avenue,
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ISSN 0097-4463

ANNALS

of CARNEGIE MUSEUM

I CARNEGIE MUSEUM O'F NATURAL HISTORY

,| 4400 FORBES AVENUE • PITTSBURGH, PENNSYLVANIA 15213

VOLUME 48 30 NOVEMBER 1979 ARTICLE 21

PALEONTOLOGY AND GEOLOGY OF THE BADWATER
CREEK AREA, CENTRAL WYOMING.

PART 19. PERISSODACTYLA

I Craig C. Black

1 Research Associate, Section of Vertebrate Fossils

i

■j Abstract

Eight species of peiissodactyis, representing six families, are found in the Badwater
faunas. All of these except a titanothere are absent from the locality 20 assemblage,
suggesting both temporal and ecological differences between that and the other Badwater
assemblages.

Introduction

Certain late Eocene perissodactyls have been reviewed by Radinsky
(1963, 1964, 1967). Even so, material in the Badwater collection adds

I significantly to our knowledge of the late Eocene helaietid Dilophodon,
and raises questions about our understanding of late Eocene titaeotU
eres and horses. Thorough review of both these groups is now badly-
needed.

The species of perissodactyls and their distribution in the Badwater
localities are given in Table L I believe it is significant that no peris-
sodactyl other than a titanothere is found at locality 20. The faueule
I from locality 20 has been considered on other evidence (Black, 1978:

231) to be younger than that from the other Badwater localities but
j slightly older than the LaPoint and Pearson Ranch faunas. The absence
of Dilophodon and Epitriplopus at locality 20 may reflect the younger
age of that assemblage, while the absence of Epihippus, Colodon, and
Amynodon is due either to depositioeal or to paleoecological factors.

Submitted for publication 20 April 1979.

391

392

Annals of Carnegie Museum

VOL. 48

Table 1. — Perissodactyls in the Badwater Fauna.

Locality

Taxa

6

5, 5A, 6, 7, Wood

Dry Creek

Family Equidae

Epihippus gracilis

Epihippus parvus

3, 20

Family Brontotheriidae

? Telmatherium cf. 7. cult ride ns

Dry Creek

Family Eomoropidae

Grangerial anarsius

5, 5A, 6, 7

5, 5A, 6, 7, Rodent

Family Helaletidae

Dilophodon leotanus

Colodon woodi

5, 6

Family Hyracondontidae

Epitriplopus uintensis

3, 5, 6

Family Amynodontidae

Amynodon sp.

The questions of faunal correlation and paleoecology will be dealt with
in detail in the final paper of the Badwater series.

All measurements given are in millimeters. Abbreviations used are as follows: CM,
Carnegie Museum of Natural History; KUMNH, Museum of Natural History, Univer-
sity of Kansas; PU, Princeton University Museum; USNM, Smithsonian Institution; L,
length; W, width; N, number; SD, standard deviation; CV, coefficient of variation.

Acknowledgments

Drs. Mary Dawson and R. M. West provided helpful discussion and reviewed the
manuscript.

This study was supported by NSF grants GB-1266, GB-4089, GB-7081, GB-30840X,
and DEB-76- 18760.

Systematic Review

Order Perissodactyla
Family Equidae Gray, 1821
G^rm^ Epihippus Marsh, 1878
Epihippus gracilis (Marsh), 1871

Referred specimens. — -USNM 21092, RP^-P; 21094, lower molar; CM 16861, isolated

upper molar; 14423, 14426 isolated lower cheek teeth.

Locality. — All CM specimens from locality 6 (Black and Dawson, 1966).

Discussion. — Epihippus gracilis is a rare form at Badwater. Only
five specimens are known, all but one of which are isolated teeth.
However, the larger size of these few specimens clearly sets them off
from specimens of Epihippus parvus, a species which is more com-

1979

Black-— Badwater Perissodactyls

393

moely found at Badwater. Measurements for dentition of E. gracilis
(length followed by width) are as follows: 16861 LM, 9.5, 1L8; 14423
RP4, 7J, 6J; 14426 RM, 9.2, 6.5.

Epihippus parvus Granger, 1908

Referred specimens .—USNM 21091, RP^-F; 21093, isolated lower molariform tooth;
CM 14424, 14427^14429, 14459--14461, 14464^14466, 14526-= 14529, 14531-14533, 14581,
14582, 14842, 15267-15275, 16069, 16808, 16815, 16862-16864, 18189, 18190, 18259,
18260, 19740, 19741, 19753, 21992, 21993, 28863, 28881, 28882, 29016-29018, all isolated
molariform teeth.

Localities.— 5, 5A, 6, 7, Wood, Dry Creek.

Discussion.— Thh smaller species of Epihippus is much more com-
mon. at Badwater than, is E. gracilis and it is found at most localities.
As all the material is isolated teeth little can be said except that the
Badwater specimens of E. parvus compare favorably in size and mor-
pliology with specimens of this species from the Uinta C Mytoe Mem-
ber of the Uinta Formation in northeastern Utah.

Family Brontotheriidae Marsh, 1873
Genus Teimatherium Marsh, 1872
? Teimatherium cf. T, cultridems
(Figs. 1-2)

Referred specimens.— CM 21211, partial left maodible with Ps-Mg; 25381, femur, tibia,
proximal end of scapula, partial innominates.

Locality.— 2Q.

Discussion.— In the absence of diagnostic skull material it is difficult
to arrive at a reliable generic determination of this form. On the basis
of size and overall dental morphology the jaw is quite similar to the
type of Teimatherium cultridens (PU 10027), a partial skull and man-
dible with complete dentition from Heiir}'’s Fork Divide, Bridger Ba-
sie, Wyoming (Osborn, 1929:168). The Badwater mandible is smaller
than that of species of Teimatherium known from the late Eocene and
it is also smaller than any of the other late Eocene titanotheres of the
genera Diplacodon, Eotitanotherium, DoUchorhinus, Metarhinus,
Sthenodectes, and Manteoceras. The dentition is bueodoet with the
crests rounded, not elevated and sharp as in the known late Eocene
species.

Wood et al. (1936) cited the presence of Teimatherium cf. cultridens
at Badwater locality 3 but this reference was made on the basis of half
a lower molar (Gazin, 1956). Until ail late Eocene titanotheres are
reviewed, a review which is badly needed, assignment of this Badwater
specimen must remain questionable.

On the basis of its dental morphology this specimen appears to rep-
resent persistence into the late Eocene of a more generalized, middle
Eocene species. Measurements are given in Table 2.

394

Annals of Carnegie Museum

VOL. 48

1979

Black— Bad WATER Pewssodactyls

395

Table 2.~~Measurements o/ ?Teimatherium c/. T. cultridens (CM 2/2// LP;i-M_^).

Teeth

L

w

P3

20.3

!2.2

P4

22.5

14.5

M,

28.0

20.0

M2

38.0

25.5

M3

50.7

25.0

Family Eomoropidae Matthew, 1929
Genus Gramgeria Zdaesky, 1930
Grangeria? anarsius (Gazin, 1956)

Eomoropus anarsius Gazie, 1956:12.

Locality Dry Creek locality, SE >4, S.9, T.39N., R.92W., Fremont County, Wyo-
mi eg.

Discussion,—No additional material of this chalicothere has been
found since Gazin's original description of the only known specimen,
USNM 21097, a partial skull with left C\ P^, and and a left
mandible with Ps-Mg. Radinsky (1964:16) transferred the species from
Eomoropus to Grangeria.

Grangerial anarsius is not yet known to occur in the main suite of
Badwater localities along Bad water Creek in Natrona County. The
single known specimen comes from the Dry Fork locality several miles
to the west in Fremont County.

Family Helaletidae Osborn, 1892
Ganm Dilophodom Scott, 1883
Diiophodmt ieotanus (Peterson), 1931
(Figs. 3-^5)

Heteraletes ieotanus Peterson, 1931:68.

Referred specimens.— USNM 21098 both maxillae with 20207 right mandible

with P2"M3; about a dozen isolated cheek teeth in the USNM collections; KUMNH
17188, 17207^17209, isolated molars; CM 28837, partial RP^M^ 14462, 18241, F; 14425,
15998, Ff 14580, 19757, P4 14578, 16893, F; 14475, partial RM"-M4 16060,

14583, 14837, 18239, 19756, 25329, Mg 14577, M^-, 14535, 15277, 15588, 18240, 21995,
M®; 23652, left mandible with dP2”dP4”Mi-M2 , right mandible with dP3”dP4”Mi”M2; 19772,
LP3^P4; 14457, 14591, P3; 28892, P4; 14454, 14478, 14579, Mi; 18186, M3.

Localities.- — -5, 5 A, 6 and 7.

Fig. l.—7Teimatherium cf. T. cultridens, CM 21211, lateral view left mandible, xVi.
Fig. 2.~7Telmatherium cf. T. cultridens, CM 21211, occlusal view LP3-M3, xVi.

396

Annals of Carnegie Museum

VOL. 48 I

Table 'i. —Measure merits of teeth o/ Dilophodon leotanus.

Catalog nos.

Teeth

L

w

CM 14457

P.3

5.3

4.7

CM 14591

Pa

5.2

4.6

CM 19772

Pa

6.1

4.4

CM 19772

P4

6.5

5.1

CM 28892

P4

6.0

5.5

CM 23652

M,

8.2

6.0

CM 23652

M,

8.1

6.0

CM 14579

M,

8.3

5.9

CM 14454

M,

8.5

6.4

CM 14478

M,

8.5

6.3

CM 23652

M,

9.2

6.5

CM 23652

M,

9.1

6.4

CM 18186

M3

10.1

6.8

CM 14462

pi

4.6

4.6

CM 18241

pi

4.5

4.0

CM 14425

p2

5.2

6.1

CM 15998

p2

5.2

6.2

CM 19757

pa

5.7

8.6

CM 14580

P

5.9

8.9

CM 14578

P

6.3

8.7

CM 16893

P

6.1

9.0

CM 16060

P

5.9

8.8

CM 25329

Ml

7.5

8.5

CM 18239

Ml

7.6

9.5

CM 19756

Ml

6.6

8.2

CM 14582

Ml

7.7

8.8

CM 16060

Ml

7.1

8.8

CM 14577

M2

8.3

9.2

CM 16060

M2

8.2

9.5

CM 18240

M3

9.0

10.5

CM 15588

M3

8.8

10.3

CM 21995

M3

9.5

10.3

CM 16060

M3

8.6

10.1

Discussion. — Radinsky (1963:53-56) has provided a thorough de-
scription of the Dilophodon dentition and has compared the dental
morphology of D. leotanus with that of D. minusculus. He character-
ized D. leotanus as being slightly smaller than D. minusculus with a
small hypocone on and with shortened lower premolar trigonids.

Fig. 3. — Dilophodon leotanus, CM 16060, LP-M^, xl.

Figs. 4-5. — Dilophodon leotanus, CM 23652, occlusal and medial view, left mandible, |
dP3=M,,xl. i'

1979

Black—Badwater Perissodactyls

397

398

Annals of Carnegie Museum

VOL. 48

1979

Black — Badwater Perissodactyls

399

The larger sample of D. leotanus from Badwater confirms that P3-
P4 are shorter than P;rP4 of D. minus cuius and that they have antero-
posteriorly compressed trigonids. There is no hypocone on P^. One
shows a short metaloph (CM 19757), whereas another (CM 14580) has
no metaloph at all. The three Ps in the sample show the same metaloph
variation from no metaloph (CM 16060) to a complete metaloph (CM
16893). A small hypocone is discernable on one P (CM 14578) but no
hypocone is present on the other Ps.

The upper and lower molars are closer in size to those of D. min-
usculus than are the premolars (Table 3). The differences in the an-
terior dentition and in the symphysis would seem to warrant specific
separation of D. minusculus and D. leotanus as suggested by Gazin
(1956) and Radinsky (1963).

Genus Colodon Marsh, 1890
Colodon woodi (Gazin), 1956

Desmatotherium woodi Gazin, 1956:17.

Holotype. —VSNM 20200, RP^-M^.

Referred USNM 20201, 20202, RF-F, CM 28873, RF;

14534, partial RMC 14463, partial LM^; 15279, LM^; 15593, RM^; 15594, RMC 15595,
LM3; 21994, LMj.

Localities.-— 5, 5A, 6, 7, and Rodent.

Discussion.— Tht additional Carnegie Museum specimens add little
to what was already known of this species (Radinsky, 1963:62) except
to support that Colodon woodi (Table 4) is smaller than Colodon kayi.

The single lower molar is relatively narrower than similar teeth of
Colodon kayi. The paralophid is weak and there is no metalophid.

Family Hyracodontidae Cope, 1879

Genus Epitriplopus Wood, 1927
Epitriplopus uintemis (Peterson, 1919)

(Figs. 6-7)

Referred specimens. —CM 14430, right lower molar; 14530, left lower molar; 16052,

left M^; 17501, left lower molar; USNM, 21099, left lower molar.

Localities. — CM 14530, locality 5, all other Carnegie specimens from locality 6.

Discussion.— GwLm (1956:23) recorded the presence of a hyraco-
dontid at Badwater on the basis of one complete and two fragmentary
lower molars. He assigned the material to Epitriplopus stating that he
could not be certain whether the form represented Epitriplopus, Tri-

plopus, or Prothyracodon.

Figs. 6-7. — Epitriplopus uintensis, (6) CM 14430, RM, x2. (7) CM 16052, LM^, x2.

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Annals of Carnegie Museum

VOL. 48

Table 4. — Measurements of teeth of Colodon woodi.

Catalog nos.

Teeth

L

w

CM 28873

RP"

8.7

11.4

CM 15594

RM*

11.1

14.5

CM 15279

LM2

12.4

14.4

CM 15593

RM2

12.9

15.8

CM 15595

LM"

14.3

16.2

CM 21994

LM,

11.4

8.5

Prothyracodon is a synonym of Triplopus while Peterson’s Proth-
yracodon uintensis (Peterson, 1919) was made the type of Epitriplopus
by Wood (1927). Among other diagnostic characters the cheek teeth
of Epitriplopus are higher crowned than those of Triplopus and there
is no trace of a metacone on in Epitriplopus, whereas the metacone
is distinct in Triplopus (Radinsky, 1967:27).

The Badwater material includes an with no metacone and the
posterior extension of the ectoloph extremely reduced and displaced
far lingually. In this character the tooth is intermediate between that
of the most reduced of Triplopus, seen in T. rhinocerinus (Radin-
sky, 1967: Plate 1) and the of Epitroplopus uintensis where no pos-
terior ectoloph extension is seen. This reduction together with the
height of crown of the worn lower molar (CM 14430) and of the
lead me to assign the Badwater hyracodont to Epitriplopus uintensis.
Measurements are given in Table 5.

Family Amynodontidae Scott and Osborn, 1883
Gtmi^ Amynodon Marsh, 1877
Amynodon sp.

Material. — CM 15447, fragment of left maxilla with partial M^ and base of M^.

Locality. — 3.

Discussion.— Wood, Seton, and Hares (1936) recorded the presence
of Amynodon advenus from Badwater, Wyoming, in a series of in-
terbedded tuffaceous silts and pond limestones. This was later record-
ed as locality 3 by Tourtelot (1957). The original Badwater collection
made by Wood, Seton, and Hares has not been seen by us and our
material is too fragmentary for specific allocation.

Table 5. — Measurements of teeth o/ Epitriplopus uintensis.

Catalog nos.

Teeth

L

w

CM 16052

M=^

16.5

18.9

CM 14430

Mj or 2

18.0

11.1

CM 17501

Ml or 2

16.0

11.1

1979

Black — Badwater Perissodactyls

401

In addition to the fragmentary maxilla, there are in the Carnegie
Museum collections fragments of molar enamel from localities 5 and
6 which probably are those of an amynodont.

Literature Cited

Black, C. C. 1978. Paleontology and geology of the Badwater Creek area, central
Wyoming. Part 14. Artiodactyls. Ann. Carnegie Mus., 47:223-259.

Black, C. C., and M. Dawson. 1966. Paleontology and geology of the Badwater
Creek area, central Wyoming. Part 1. History of field work and geological setting.
Ann. Carnegie Mus., 38:297-307.

Gazin, C. L. 1956. The geology and vertebrate paleontology of upper Eocene strata
in the northeastern part of the Wind River Basin, Wyoming. Part 2. The mammalian
fauna of the Badwater area. Smithsonian Misc. Coll., 131:1-35.

Osborn, H. F. 1929. The titanotheres of ancient Wyoming, Dakota and Nebraska.
U.S. Geol. Surv. Monogr. 55, vol. 1:1-698.

Radinsky, L. 1963. Origin and early evolution of North American Tapiroidea. Bull.
Peabody Mus. Nat. Hist., Yale Univ., 17:1-106.

. 1964. Paleomoropus, a new early Eocene chalicothere (Mammalia, Perisso-

dactyla), and a revision of Eocene chalicotheres. Amer. Mus. Novit. 2179:1-28.

. 1967. A review of the rhinoceratoid family Hyracondontidae (Perissodactyla).

Bull. Amer. Mus. Nat. Hist., 136:1-46.

Tourtelot, H. a. 1957. The geology and vertebrate paleontology of upper Eocene
strata in the northeastern part of the Wind River Basin, Wyoming. Part I: Geology.
Smithsonian Misc. Coll., 134:1-27.

Wood, H. E., H. Seton, and C. J. Hares. 1936. New data on the Eocene of the
Wind River Basin, Wyoming. Proc. Geol, Soc. Amer., 1935:394-395 (abstract).

Back issues of many Annals of Carnegie Museum articles are
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ANNALS

0/ CARNEGIE MUSEUM

CARNEGIE MUSEUM OF NATURAL HISTORY

4400 FORBES AVENUE « PITTSBURGH, PENNSYLVANIA 15213

VOLUME 48 30 NOVEMBER 1979 ARTICLE 22

THE RHYACOPHILA OF PENNSYLVANIA, WITH LARVAL
DESCRIPTIONS OF R. BANKSI AND R. CARPENTERI
(TRICHOPTERA: RHYACOPHILIDAE)

John S. Weaver IIP

Jan L. Sykora^

Research Associate, Section of Insects and Spiders

Abstract

Over 3,000 immature and adult specimens of the genus Rhyacophiia were collected
in Pennsylvania. This collection consists of 17 species most of which are new state
records. The larvae of R. banksi Ross and R. carpenteri Milne were associated by the
metamorphotype method. The larva of banksi is indistinguishable from known larvae
of the other species of the invaria subgroup (Schmid, 1970). The larva of carpenteri is
identical to R. sp. 3 {banksi ?) of Flint (1962). A revised key to the 23 known larvae of
eastern Nearctic Rhyacophiia is given.

Introduction

There are 34 eastern Nearctic species of Rhyacophiia, of which four
were described from female types. Since Flint’s (1962) key to the lar-
vae of eastern Nearctic Rhyacophiia, additional larval associations of
this group have been given by Smith (1968) for angelita, Sherberger
and Wallace (1971) for vuphipes, and Neves (1977) for acutiloba and
Carolina. The larvae of banksi and carpenteri are described herein,
and were associated by the metamorphotype method of Milne (1938).

^ Department of Entomology and Economic Zoology, Clemson University, Clemson,
South Carolina 29631.

^ Graduate School of Public Health, University of Pittsburgh, Pittsburgh, Pennsylvania

15261.

Submitted for publication 15 March 1979.

403

404

Annals of Carnegie Museum

VOL. 48

Therefore, the larvae of 23 of the 30 species recognized by male gen-
italia are presently known, leaving accola, appalachia, montana, myc~
ta, otica, parantra, and teddyi unassociated.

In the past, little work has been done documenting the occurrence
of Rhyacophila in Pennsylvania. Hyland (1948), in a list of 29 species
of Trichoptera from the state, recorded only one Rhyacophila— nigri-
ta. Ross (1941) mentioned three, fuscula, invaria, and nigrita, and
Roback (1975) listed immatures of fuscula, and two other tentatively
identified taxa.

For this study the adults were collected along streams and springs
using light traps and sweeping nets. The larvae and pupae were hand-
picked from stones and debris submerged in water or collected by a
Hess type quantitative sampler.

The majority of specimens examined in this investigation were col-
lected from Pennsylvania. However, a few records from adjacent areas
of New Jersey are also included as they provide insight into the dis-
tribution of Rhyacophila in Pennsylvania. Over 3,000 immature and
adult specimens, consisting of 17 species of Rhyacophila, were pro-
cured, most of which are new state records. The accumulation of these
data from 23 counties has contributed to the knowledge of the local
distribution, seasonal occurrence, and natural history of several
species.

The bibliography of each species listed is not complete, and includes
only descriptive references of the larvae and adult males. A list of the
Rhyacophila (Schmid, 1970) occurring in eastern North America is
given, because it includes species which probably later will be record-
ed from Pennsylvania.

List of Eastern Nearctic Rhyacophila
fuscula Group

*t fuscula (Walker 1852) Neuronia
t vuphipes Milne (1936)

montana Group
montana Carpenter (1933)

glaberrima Group
t glaberrima Ulmer (1907)

Carolina Group

*t Carolina Banks (1911)
fenestra Ross (1938fl)
kiamichi Ross (1944)
t ledra Ross (1939)

1979

Weaver and Sykora — Pennsylvanian Rhyacophila

405

t otica Etnier and Way (1973)
teddyi Ross (1939)

invaria Group
invaria complex

t banksi Ross (1944)
t invaria (Walker 1852) Polycentropus
parantra Ross (1948)
shenandoahensis Flint (1958)

*t vibox Milne (1936)

nigrita complex
accola Flint (1972)
acutiloba Morse and Ross (1971)
appalachia Morse and Ross (1971)
t carpenteri Milne (1936)
mycta Ross (1941)

*t nigrita Banks (1907)

lobifera Group
t lobifera Betten (1934)

angelita Group
angelita Banks (1911)

siberica Group

amicis Ross (1956)
t atrata Banks (1911)
t manistee Ross (1938i3)

* melita Ross (1938«)
t minor Banks (1924)

torva Group
t torva Hagen (1861)

acropedes Group

acropedes Banks (1914)
ignorata Schmid (1974) 9 holotype

ungrouped species
for mo s a Banks (1911) $ holotype
mainensis Banks (1911) 9 holotype
soror Provancher (1878) 9 holotype

t Species recorded from Pennsylvania.
* Species recorded from New Jersey.

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Annals of Carnegie Museum

VOL. 48

Key to Eastern Ne arctic Rhyacophila Larvae
(modified from Flint, 1962)

(Larvae of accola, appalachia, montana, mycta, otica, parantra, and teddyi

unknown)

i. Anal proleg with an apicolateral spur (Flint, 1962: Fig. lb) . 2

Anal proleg without such a spur (Flint, 1962: Fig. 2b) ........... 7

2(1). Apicolateral spur of proleg short and sharply decurved (Flint, 1962:

Fig. 6b) 3

Spur long and evenly curved (Flint, 1962: Fig. lb) 5

3(2). Body segments with alternating light and dark bands ....... melita

Body segments not contrastingly banded 4

4(3). Frontoclypeus with dark “H” pattern (Sherberger and Wallace, 1971:

Fig. 1) vuphipes

Frontoclypeus not patterned as above amicis

5(2). Frontoclypeus with dark transverse bands forming a “W" pattern

(Flint, 1962: Fig. Ic) fuscula

Frontoclypeus without such a pattern 6

6(5). Head uniformly colored (Flint, 1962: Fig. 4c) ............... atrata

Head with a dark transverse band posteriorly (Flint, 1962: Fig.
6c) ................................................... angeiita

7(1). Abdomen bearing many branched gills .... acropedes

Abdomen without gills 8

8(7). Anal proleg with a free basoventral hook (Flint, 1962: Fig. 2b) ... 9
Anal proleg without a free basoventral hook (Flint, 1962: Fig. 8b, d)

12

9(8). Anal claw without ventral teeth (Flint, 1962: Fig. 5d) 10

Anal claw with ventral teeth (Flint, 1962: Fig. 2b) 11

10(9). Head equal in length and width, with lateral margins obliquely angled

(Flint, 1962: Fig. 5a) minor

Head slightly longer than wide, with lateral margins curved (Flint,
1962: Fig. 5c) ........................................ manistee

11(9). Frontoclypeus with a dark ‘‘V” pattern (Flint, 1962: Fig. 2c); mandi-
bles edentate (Flint, 1962: Fig. 2a) torva

Frontoclypeus not patterned as above; mandibles toothed apically

(Flint, 1962: Fig. lib) lobifera

12(8). Second segment of maxillary palps subequal to first (Ross, 1944: Fig.

114) glaberrima

Second segment of maxillary palps longer than first (Fig. 4d here-
in) 13

13(12). Anal claw without ventral teeth (Flint, 1962: Fig. lid), Carolina

Group 14

Anal claw with ventral teeth (Flint, 1962: Fig. 8b), invaria Group ... 16
14(13). Head with distinct pattern of infuscations and muscle scars (Flint,
1962: Fig. 10b) .............................. fenestra and ledra

Head nearly concolorous 15

1979

Weaver and Sykora — Pennsylvanian Rhyacophila

407

15(14). Head slightly darkened posteriorly (Flint, 1962: Fig. 11a), (Arkansas

and Oklahoma) kiamichi

Head uniformly golden brown (Flint, 1962: Fig. 11c; Fig. 4a herein),

(Appalachian and Piedmont regions) Carolina

16(13). Head and pronotum nearly black with a conspicuous pattern of pale

muscle scars and spots (Flint, 1962: Fig. 9b) acntiloba

Head not marked as above . 17

17(16). Head narrowing on posterior half (Flint, 1962: Fig. 10a; Fig. 8a

herein) carpenteri

Head either parallel sided or broadening posteriorly 18

18(17). Head nearly parallel sided (Flint, 1962; Fig. 9a) nigrita

Head distinctly broadening posteriorly (Flint, 1962: Fig. 8a, c; Fig. 6a
herein) — invaria subgroup: hanksi, invaria, shenandoahensis and vi-
box

Tm Rhyacophila of Pennsylvania
fuscula Group

Rhyacophila fuscula (Walker)

Neuroma fuscula Walker 1852.

Rhyacophila fuscula (Walker) Ross, 1944 (male and female genitalia, larva). Flint, 1962
(larva). Schmid, 1970 (male and female genitalia). Wiggins, 1977 (larva).

There is no other species of Rhyacophila from North America as
frequently documented in the literature as fuscula. It is known from
nearly every state and province in the Appalachian Mountains and the
surrounding Great Lakes regions. Being a montane species, it is very
widespread in the Pennsylvania highlands (Fig. 1). Both Flint (1962)
and Roback (1975) discovered it to be the most common species of
Rhyacophila in their collections of immatures from the East. How-
ever, from the material examined it appears to be no more widespread
than Carolina in the Alleghenies of Pennsylvania and much less com-
mon at elevations below 460 m.

The larvae were often collected from swiftly flowing riffles of pristine
mountain streams, ranging from approximately 1.5 to 12 m in width.
In the brook (3 to 6 m wide) issuing from Maul Spring, a constant
temperature, well-oxygenated spring (9°C ± LC), larvae were never
taken near the source, but only at distant reaches where water tem-
perature regimes fluctuated diurnally. These observations suggest that
the larvae of fuscula prefer a eury thermic environment. Seasonally,
larvae were c ollected in the spring and adults throughout the summer.

Material examined. — Pennsylvania. Elk County: Ridgway, South Branch Island
Run, 30 May 1977, 1 larva. Fayette County: Seven Springs, Back Run, 2 June 1977, 1
<3. Franklin County: Turnpike between Blue and Kittatinny Mountains, 12 April 1977,

1 larva; 18 May 1977, 3 larvae. Forest County: Kellettville, Ross Run, 5 June 1976, 5
larvae; 27-28 August 1976, 3 9 9; Marienville Sportsmens Club, Spring Creek, 3-4 July

408

Annals of Carnegie Museum

VOL. 48

1976, 1 larva, 4 6 S, 1 9; 28-29 August 1976, 1 9; Tionesta Station, Hunters Creek, 5 !j
June 1976, 3 larvae, 1 (5 , 1 9 ; 25 July 1976, 1 9 ; 28 August 1976, 4 66, 1 9. Fulton j

County: McConnellsburg, Aughwick Creek, 30 June 1977, 2 larvae, 3 66. Lancaster |

County: New Holland Watershed Area, 4 June 1977, 2 larvae. Pike County: Milford, ;
Sawkill Creek, 1 May 1977, 1 larva. Potter County: Harmontown, south branch of |

Genesee River, near Route 449, 2 July 1978, 3 larvae, 1 6. Somerset County: Kooser j

State Park, 7-8 June 1976, 4 6 6: 22-23 June 1976, \ 6, I 9: 20-21 August 1976, 1
6: 25-26 September 1976, 1 6: Laurel Hill State Park, Jones Mill Run, 7-8 June 1976, j
1 6: 22-23 June 1976, 10 (3(3, 4 9 9. Venango County: Utica, French Creek, 30 June j
1976, 1 9. Westmoreland County: Linn Run State Park, 20-21 August 1976, 13 c3c3, 5
9 9: Laughintown, Furnace Run, 15 May 1976, 2 larvae; 7-8 June 1976, 4 c3(3, 2 9 9; jj
14-15 July 1976, 13 (3(3, 3 9 9; 20-21 August 1976, 18 c3(3, 10 9 9; 25-26 September ;i
1976, 2 6 6,2 9 9; Rector, Powdermill Nature Reserve, Maul Spring, 31 May 1976, 1 i'

(3 ; 19-20 June 1975, 1 (3,3 9 9; 16-17 July, 1975, 3 (3 (3 ; 13-14 August 1975, 10 (3 (3 , jl

23 9 9 ; 10-11 September 1975, 1 (3,4 9 9 ; 27 February 1976, 3 larvae. New Jersey. ||

Sussex County: North of Highpoint State Park near Route 23, Clove Run, 16 April 1977, 'i

9 larvae: 19 May 1977, 2 larvae, 1 prepupa. ;

Rhyacophila vuphipes Milne ji

Rhyacophila vuphipes Milne, 1936 (male genitalia). Ross, 1956 (male genitalia). Schmid, i
1970 (male genitalia). Roback, 1975 (larva). j

Rhyacophila vuphiphes Sherberger and Wallace, 1971 (lapsus), (larva). ii

This species is known from the eastern slopes of the Appalachian |

Mountains of the middle and southern Atlantic coastal states. It is |

reported here for the first time from Pennsylvania, while other records j

exist from Georgia, New York, North Carolina, South Carolina, and

1979

Weaver and Sykora — Pennsylvanian Rhyacophua

409

Fig. 2,— Distribution of the species of Rhyacophila of the lobifera, glaberrima, and
torva groups.

Tennessee . Its most northern occurrence is in Quebec (Roy and Harp-
er, 1975). However, it is more frequently collected in the southern
Appalachian Piedmont region.

In Pennsylvania only one larva was collected from the riffle of a
large meandering stream approximately 15 m in width (Fig. 1). Im-
matures may be collected in the early summer, and adults are on the
wing from August to September.

Material Pennsylvania. Cumberland County: Boiling Springs, Yellow

Breeches Creek, 12 June 1976, 1 larva; 17-18 September 1976, 2 6 6.

glaberrima Group
Rhyacophila glaberrima Ulmer

Rhyacophila glaberrima Ulmer, 1907 (male genitalia). Ross, 1944 (male and female
genitalia, larva). Ross, 1956 (male genitalia). Flint, 1962 (larva). Schmid, 1970 (male
genitalia). Roback, 1975 (larva).

This species, the only member of the glaberrima group, is widely
distributed throughout eastern and central North America. It is re-
corded for the first time from Pennsylvania, where it was collected
commonly from the central and western highlands (Fig. 2). Other state
and provincial records include Georgia, Illinois, Massachusetts, New
Hampshire, New York, North Carolina, Nova Scotia, Quebec, Ten-
nessee, and Virginia. In Pennsylvania larvae were found in small

410

Annals of Carnegie Museum

VOL. 48

Fig. 3. — Distribution of the species of Rhyacophila of the Carolina group in Pennsyl- |;
vania. [i

streams and springs. Immatures may be collected in the spring and I
summer, and adults in the late summer and early autumn.

Material examined. — Pennsylvania. Clearfield County; S. B. Elliott State Park, |
Lick Run, 30 May 1977, 7 larvae, 1 prepupa; 1 August 1977, 3 S pupae; 1 June 1978, j

1 larva. Fulton County; Emmaville, Chestnut Bridge Hollow, 5 dri. 5 9 9 ; 29 May !

1977, 1 larva; 1 July 1977, 1 larva, 1 prepupa. Washington County: Amity, small spring
near Ten Mile Creek, 15 August 1976, 1 S . Westmoreland County: Laurel Hill Summit, i

Spruce Run, 21-22 August 1976, 2 Linn Run State Park, 20-21 August 1976, 3
(3(3, 1 9; Laughintown, Furnace Run, 14-15 July 1976, 1 3; 20-21 August 1976, 14 j-

3 3,8 9 9; Rector, Powdermill Nature Reserve, Maul Spring. 16-17 July 1975, 2 3 3; i

13-14 August 1975, 14 3 3, 6 9 9; 10-1 1 September 1975, 3 33,5 9 9:6 June 1977, ji

2 larvae; 20 July 1977, 1 prepupa. jl

Carolina Group S

Rhyacophila Carolina Banks

Rhyacophila Carolina Banks, 191 1 (male genitalia). Ross, 1938/? (male genitalia), Schmid, |
1970 (male genitalia). Neves, 1977 {R. sp. 5 Flint, 1962, larva is Carolina). j|

Rhyacophila sp. 5 {Carolina ?), Flint, 1962 (larva). |

Rhyacophila Carolina was the most commonly collected species in [
this study, occurring throughout the western slopes of the Ohio River |i
Valley, the central ridge and valley system of the Alleghenies, and the I
rolling foothills of the southeastern Piedmont (Fig. 3). It is also known '|
from Kentucky, Maine, Massachusetts, New Hampshire, New York, i
North Carolina, Quebec, South Carolina, and Tennessee.

1979

Weaver and Sykora — Pennsylvanian Rhyacophila

41 1

Fig. 4. — Rhyacophiia Carolina, larva: A = head aod pronotum, dorsal view; B = anal
proleg, lateral view; C = mandibles, dorsal view; D = maxillary palpus (MP).

Description of larva (Fig. 4). — Overall length of mature larva 16 mm. Head: Dis-
tinctly narrowing anteriorly; coloration variable; in some specimens golden brown, im-
maculate; in others, slightly darker with faint muscle scars; right mandible with three
apical teeth, middle tooth slightly longer than dorsal tooth and much longer than ventral
tooth. Left mandible with two apical teeth subequal in length. Abdomen: Anal proleg
with very short basoventral and apicolateral processes, the latter barely noticeable; anal
claw with no ventral teeth. Neves (1977) has associated this species with the larva! form,
R. sp. 5 of Flint (1962). It is the authors’ opinion that the larval forms of R. sp. 6 and
R. sp. 7 of Roback (1975) are both Carolina.

Habitat .—Mdiny larvae vyere collected from small pristine streams
0.6 to 4.8 mm in width. Occasionally, immatures were also found in
springs. Mature larvae were collected from February to August, pre-
pupae from February to May, pupae from May to August, and adults
from April to October. These observations make it difficult to deter-
mine either if most mature larvae discovered in the late summer pupate
and emerge in the autumn, or if some may overwinter as pupae. These
overwintering pupae may be part of a second generation that emerge
in the spring, or there may be two cohorts involved.

412

Annals of Carnegie Museum

VOL. 48

Material “-Pennsylvania. Beaver County: Raccoon Creek State Park,

stream at the marina, 18-19 July 1976, 1 (5,1 $ . Chester County: Atgien, Glen Run, 21
March 1977, 4 larvae, 4 prepupae; 11 April 1977, 6 larvae, 6 prepupae; 24 ApriJ 1977, t

1 larva, 2 prepupae; 7 May 1977, 1 larva, 7 prepupae, 2 S pupae, 6 <^(5, 1 9; 17 May

1977, 10 <7 (7 , 4 9 9 ; 6-7 June 1977, 18 d d , 20 9 9 ; 18-19 June 1977, 41 c7 <7 , 9 9 9 ; 23- ?

26 June 1977, 1 larva, I (7 pupa, 25 c7(7, 15 9 9; 12-14 July 1977, 5 c7(7; 25 October !;

1977, 1 (7, 1 9; Atgien, Spring Grove Dairy, 6 March 1977, 2 larvae; 21 March 1977, 9
prepupae; 8 May 1977, 7 <7 pupae, 7 9 pupae; 12 July 1977, 8 6 pupae, 8 9 pupae, 1

9; 27 February 1978, 3 larvae, 1 prepupa. Clearfield County: S. B. Elliott State Park, |

Lick Run, 30 May 1977, 7 larvae, 1 prepupa; 1 August 1977, 3 6 pupae. Cumberland i'

County: Mount Holly Springs, Whisky Spring, ! July 1977, 2 <7 <7 . Elk County: Ridgway, 1

spring near Route 949, 23 June 1977, 1 larva. Fayette County: Seven Springs, Neals I

Run Spring, 2 June 1977, 1 S . Franklin County: Turnpike between Blue and Kittatinny |j

Mountains, Trout Run, 12 June 1976, 1 (7, 1 9. Forest County: Kellettville, Ross Run,

18 June 1976, 1 larva; 23-28 August 1976, 3 c7 c7 , 5 9 9 ; 8-9 October 1976, 2 c7(7; I

Marienville Sportsmens Club, Spring Creek, 28-29 August 1976, 9 <7 (7 , 6 9 9; Tionesta j!

Station, 25 July 1976, 1 9. Fulton County, Emmaville, Chestnut Bridge Hollow, 23-24
July 1976, 5 (7(7, 2 9 9; 31 July 1976, 16 c7c7, 3 9 9; 15-16 September 1976, 1 (7, 1
9; 1 July 1977, 1 6 pupa, 1 (7; 29 July 1977, 1 <7 pupa; McConnelisburgh, Aughwick
Creek, 30 June 1977, 22 S6, 1 9. Lancaster County: Christiana, Pine Creek, 18-19
September 1976, 5 (7 (7 , 4 9 9; New Holland Watershed Area, 4-5 April 1977, 2 larvae,

2 (7(7; 26-27 June 1977, 1 (7; 16-17 July 1977, 17 66, 1 9; 5 October 1977, I (7; 23
March 1978, 4 prepupae. Pike County: Metamoras, Bushkill Creek, 12 June 1977, 1
6 . Somerset County: Kooser State Park, 22-23 June 1976, 1 <7; Laurel Hill State Park,
Jones Mill Run, 7-8 June 1976, 2 6 6,1 9 ; 22-23 June 1976, 2 <7 <7 , I 9 . Warren County:
Cobham, Rams Horn Spring, 24-25 July 1976, 1 6i 18 April 1977, 1 prepupa. Washington
County: Amity, small spring near Ten Mile Creek, 10 June 1976, 4 (7 (7 , 3 9 9; 16 June

1976, 5 6 6,3 9 9; 15 August 1976, 1 9. Westmoreland County: Laurel Hill Summit,
Spruce Run, 12 June 1976, 1 larva; 14-15 July 1976, 1 larva, 35 <7 c7 , 8 9 9; 21-22 August

1977, 1 larva, 18 <7(7; Linn Run State Park, 30 July 1976, I <7; 20-21 August 1976, 5
(7(7, 4 9 9; Laughintown, Furnace Run, 7-8 June 1976, 3 (7 c7 ; 14-15 July 1976, 14
(7 (7, 5 9 9; 20-21 August 1976, 16 66, 5 9 9; Rector, Powdermill Nature Reserve,
Maul Spring, 19-20 June 1975, 1 c7; 16-17 July 1975,3 <7(7,3 9 9; 13-14 August 1975,

13 6 6 ,35 9 9; 10-1 1 September 1975, 1 <7 . New Jersey. Sussex County: Stokes State
Forest, stream near Sawmill Road, 30 April 1977, 1 prepupa; 11 June 1977, I larva.

RhyacopMla ledra Ross

Rhyacophila ledra Ross, 1939 (male genitalia). Ross, 1944 (male genitalia, larva). Ross,

1956 (male genitalia). Flint, 1962 (larva). Schmid, 1970 (male genitalia).

This species has been recorded previously from Georgia, Illinois,
North Carolina, Ohio, and Tennessee. It is not common in Peensyb
vania, from which it is recorded for the first time (Fig. 3). Its overall
distribution apparently entails the Ohio River Valley and the southern
Appalachian Mountains. There are insufficient data available to con-
clude v/hat is the definitive aquatic habitat of this species. Ross (1944)
collected a few mature pupae in May from a small temporary stream.
Adults have been collected on the wing in June.

Material examined.— Fennsylvania. Armstrong County: Vandergrift, Carnahan
Run, 1 June 1977, 1 <7. Westmoreland County: Rector, Powdermill Nature Reserve,
Powdermill Run, 19-20 June 1975, 1 <7; Maul Spring, 23-24 June 1977, 1 <7, 1 9.

1979

Weaver and Sykora— Pennsylvanian Rhyacophila

413

Fig. 5. — Distribution of the species of Rhyacophila of the invaria subgroup in Penn-
sylvania.

Rhyacophila otica Etnier and Way

Rhyacophila otica Etnier and Way, 1973 (male genitalia).

Rhyacophila pennsylvanica Sykora and Weaver, 1976, N . syn.

This species previously recorded from Tennessee by Etnier and Way
(1973) is relatively uncommon in Pennsylvania where only a few adult
specimens have been collected at an ultraviolet light trap (Fig. 3). The
immature stages of this species remain unknown.

Material examined. — Pennsylvania. Fulton County: Emmaville, Chestnut Bridge
Hollow, 23-24 July 1976, 1 cJ, 1 $. Westmoreland County: Rector, Powdermill Nature
Reserve, Maul Spring, 14-15 July 1975, 1 d; Powdermill Run, 14-15 July 1975, 2 66.

invaria Group

This group is composed of two subgroups, the invaria subgroup and
the nigrita subgroup (Schmid, 1970) both of which are restricted to
eastern Nearctica.

invaria Subgroup
Rhyacophila banksi Ross

Rhyacophila banksi Ross, 1944 (male and female genitalia).

Rhyacophila sp. 3 [carpenteri ?), Flint, 1962.

This interesting species was found to be rather common in the Ap-
palachians of western Pennsylvania (Fig. 5). Other state records in-
clude New Hampshire, Tennessee, and Vermont.

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VOL. 48

Fig. 6. — Rhyacophila hanksi, larva: A = head and pronotum, dorsal view; B = anal
proleg, lateral view; C = mandibles, dorsal view; D = maxillary palpus (MP).

Description of larva (Fig. 6). — Overall length 16 mm. Head: Coloration variable; in
some, golden brown, in others dark brown with an irregular row of pale muscle scars
extended on each side of the front and the coronal suture, muscle scars faintly distin-
guishable on the frontocephalis, lateral and posterior portions lighter; right mandible
with three apical teeth, mesal tooth longest; left mandible with two apical teeth, ventral
tooth longer; maxillary palpi with second segment twice as long as the first. Abdomen:
Proleg with basoventral and apicolateral processes, neither free of membrane; claw with
one large ventral tooth and sometimes a smaller tooth.

Flint (Roback, 1975) was of the opinion that "it is impossible to separate the larvae
of R. invaria, R. shenandoahensis, and R. vihox." Likewise, we are unable to distin-
guish the larvae of R. hanksi from these three species.

Discussion. — These species, besides sharing a hypothetical common
ancestor and many synapomorphic characters, also appear to inhabit
similar aquatic environments, specifically small streams and springs.
Furthermore, it is uncommon for more than one species of the invaria
subgroup to occur at the same locality, therefore the species involved
appear to be allopatric.

1979

Weaver and Sykora — Pennsylvanian Rhyacophila

415

Habitat. — Many larvae of hanksi were collected from small streams
and springs (0.3 to 1.2 m wide). However, a few were collected near
the confluence of the hatchery spring and Spring Creek at Marienville.
Immatures were also taken from Spruce Run, a naturally acidic stream
(pH = 3.5 in July). Seasonally, immatures have been collected in the
spring and early summer, and adults are on the wing from June to
August.

Material examined. — Pennsylvania. Elk County: Ridgway, small spring near Route
949, 3-4 July 1976, 5 dd, 2 9 $; 16 April 1977. 9 larvae; 30 May 1977, 2 larvae, 2
prepupae; 23 June 1977, 1 prepupa, 4 6 pupae; 1 July 1977, 4 <? pupae, 1 $ pupa; 12
July 1977, 3 S pupae, 3 9 pupae, 1 9. Fayette County: Seven Springs, Neals Run
Spring, 15 April 1977, 4 larvae, 2 prepupae; 2 June 1977, 3 6 pupae, 1 6 . Forest County;
Marienville Sportsmens Club, hatchery spring, 19 July 1976, 1 larva; 28-29 August 1976,
1 c3; 18 April 1977, 3 larvae, 2 prepupae; Spring Creek, 18 April 1977, 1 larva, 8 pre-
pupae. Warren County: Cherry Grove, Sandstone Spring Picnic Grounds, 6 June 1976,
1 larva, 1 6 pupa; 18 April 1977, 2 larvae. Washington County: Amity, spring near Ten
Mile Creek, 10 June 1976, 6 d d ; 16 July 1976, 1 S. Westmoreland County; Laurel Hill
Summit, Spruce Run, 7 June 1973, 3 6 pupae, 2 9 pupae; 16 May 1976, 5 larvae, 5
prepupae; 12-13 June 1976, 3 6 pupae, 2 9 pupae, 18 dd, 4 9 9; 14-15 July 1976, 41
dd, 1 9; 20 March 1977, 3 larvae.

Rhyacophila invaria (Walker)

Poiycentropus invarius Walker, 1852.

Rhyacophila invaria (Walker) Ross, 1938/? (male genitalia). Flint. 1962 (larva). Schmid,
1970 (male genitalia).

In Pennsylvania this species was collected only in the eastern re-
gions of the Tuscorora Mountains and the Piedmont (Fig. 5). It is
apparently separated in distribution from hanksi which is common in
the western mountains of the Appalachians. It is also known from
Maine, Massachusetts, New York, Quebec, and Nova Scotia. Several
larvae were collected in springbrooks from February to July, mature
pupae in June, and adults were on the wing in June and July. The
larvae are very similar to those of other species of the invaria subgroup
(see hanksi).

Material examined. — Pennsylvania. Chester County: Atglen, small spring near
Glen Run, 11 April 1977, 1 larva; 24 April 1977, 21 larvae, 6 prepupae; 7 May 1977, 5
prepupae, 1 6 pupa; 6-7 June 1977, 5 larvae, 1 prepupa, 1 S pupa, 1 9 pupa; 19 June
1977, 11 dd, 5 9 9; 23 June 1977, 2 9 9; 25-26 June 1977, 23 dd, 1 9; 12 July 1977,
1 larva; 27 February 1978, 2 larvae. Cumberland County: Mount Holly Springs, Whisky
Spring, 1 July 1977, 2 S S . Fulton County: McConnellsburgh, Aughwick Creek, 30 June
1977, 1 larva, 3 prepupae, 1 S pupa, 2 9 pupae, 2 dd, 1 9; 2 July 1977, 1 9 pupa, 1
d; 17 July 1977, 3 d pupae, 2 9 pupae. Lancaster County: New Holland Watershed
Area, 4-5 April 1977, 2 larvae, 2 9 pupae; 26-27 June 1977, 7 prepupae, 1 d pupa, 3
9 pupae, 1 9; 16-17 July 1977, 8 dd; 23 March 1978, 1 larva.

Rhyacophila vibox Milne

Rhyacophila vibox Milne, 1936 (male and female genitalia). Ross, 1944 (male and female
genitalia, larva). Flint, 1962 (larva). Schmid, 1970 (male genitalia).

416

Annals of Carnegie Museum

VOL. 48

Fig. 7. — Distribution of the species of Rhyacophila of the nigrita subgroup in Pennsyl-
vania.

This is the most widely dispersed species of the invaria subgroup, I
being known from Illinois, Maine, Massachusetts, Michigan, New
Hampshire, New York, North Carolina, Ontario, Quebec, Tennessee,
Vermont, Wisconsin, and recorded herein for the first time from Penn- i

sylvania and New Jersey (Fig. 5). Its distribution in Pennsylvania ap- '

parently is not restricted to a specific geographic region of the state as !

observed for two other species of the invaria subgroup, banksi and j

invaria. The larvae of vibox are very difficult to distinguish from the |

other known larvae of this subgroup (see banksi). The immatures of '

vibox have been collected mostly from small eury thermic streams, 0.6 |

to 1.5 m wide from April to June. Mature pupae and adults occur in
May and June. 1

Material examined. — Pennsylvania. Beaver County: Raccoon Creek State Park, I
Mineral Run, 30 May 1976, 2 SS \ stream near marina, April 1975, 2 larvae; 28 May
1976, 5 6 pupae, 1 d; 19 May 1977, 1 larva, 5 6 pupae. Clearfield County: S. B. Elliott
State Park, Lick Run, 18 April 1977, 9 larvae, 2 prepupae; 30 May 1977, 2 larvae, 6 j

prepupae, 14 6 pupae, 1 6, 1 9 ; 23 June 1977, 12 6 pupae, 8 9 pupae; 1 May 1978, 'i

3 larvae, 2 prepupae; 1 June 1978, 2 larvae, 1 prepupa, 3 S pupae, 4 9 pupae, 6 S . i|

Forest County: Kellettville, small spring near Ross Run, 7 June 1978, 6 6 pupae, 7 9 ij

pupae. Potter County: Harmontown, small spring near Route 449, 2 July 1978, 4 larvae,

5 prepupae, 1 S pupa. Westmoreland County: Rector, Powdermill Nature Reserve, l

White Oak Run, 31 May 1975, 1 6.2 9 9. New Jersey. Sussex County: Stokes State i

Forest, small stream near Sawmill Road, 4 April 1977, 3 larvae; 11 April 1977, 1 c?, 2
9 9 ; 30 June 1977, 1 larva, 1 prepupa.

1979

Weaver and Sykora — Pennsylvanian Rhyacophila

All

Fig. 8. — Rhyacophila carpenteri, larva: A = head and pronotum, dorsal vievi/; B = anal
proleg, lateral view; C = mandibles, dorsal view; D = maxillary palpus (MP).

nigrita Subgroup
Rhyacophila carpenteri Milne

Rhyacophila carpenteri Milne, 1936 (male and female genitalia). Ross, 1956 (male gen-
italia). Schmid, 1970 (male genitalia).

Rhyacophila sp. 3 (banksil), Flint, 1962.

This interesting species is distributed throughout the Appalachian
Mountains, being documented from Kentucky, Massachusetts, New
Hampshire, North Carolina, and Quebec. It is a new state record for
Pennsylvania, where three localities were discovered to be frequented
by adults, and subsequent samplings procured several larvae and meta-
morphotypes (Fig. 7). It is identical to i?. sp. 3 larval form of Flint

418

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VOL. 48

(1962), who remarks, “This species is easily recognized by the shape
of the head, for no other species yet found in the East has a head
capsule that gradually narrows toward the rear.”

Description of larva (Fig. 8). — -Overall length, !4 mm. Head: Light chestnut color,
nearly parallel sided, but distinctly narrowing posteriorly; right mandible with three
apical teeth, middle tooth longest; left mandible with two apical teeth, ventral tooth
longer; maxillary palpi with second segment 2.5 times as long as the first. Thorax:
Pronotum with a row of setae on the anterior margin, shorter mesad and gradually
lengthening anterolaterally. Abdomen: Anal proleg with basoventral and apicolateral
processes, neither free of the membrane; anal claw with one small ventral tooth.

Immatures were collected from a small spriegbrook, 0.9 |

to 1 .5 m wide, approximately 60 m from the source. Larvae were taken |
in April and May; prepupae, May and June; pupae, June and July; i

adults, June, July, and August. |

1

Materia! examined. — Pennsylvania. Elk County: Ridgway, spring near Route 949,

3-4 July 1976, 12 dd; 16 April 1977, 1 larva;' 30 May 1977, 2 larvae, 2 prepupae; 23
June 1977, 1 prepupa, 2 S pupae; 1 July 1977, 4 S pupae, I 9 pupa; 12 July 1977, 3
S pupae, 3 9 pupae, I 9 ; Ridgway, South Branch Island Run, 3 July 1976, 2 S 6 . Tioga
County: Ansonia, Colton Point, Right Branch Run, 1-2 July 1978, 5 6 6,2 9 9. West-
moreland County: Rector, Powdermill Nature Reserve, Maul Spring, 19-20 June 1975,

1 d; 16-17 July 1975, 19 riri, 3 9 9; 13-14 August 1975, A 66, 1 9; 23-24 June 1977,

5 dri.

Rhyacophila nigrita Banks

Rhyacophila nigrita Banks, 1907. Betten, 1934 (male genitalia). Flint, 1962 (larva).

Schmid, 1970 (male genitalia).

This species is very common throughout the Appalachians, being
previously recorded from Kentucky, Massachusetts, New Hampshire,
North Carolina, Pennsylvania, Quebec, and Tennessee; specimens re-
ported herein are a new state record from New Jersey, In Pennsylvania
it was collected from many different localities (Fig. 7), its larvae typ-
ically inhabiting clear mountain rivulets concurrently with fuscula and
Carolina. Many immatures were collected at Maul Spring— first in-
stars, from June to August; fifth (last) instars, from November to June;
adults, from May to September.

Material examined. — Pennsylvania. Bedford County: Schellsburg, spring near |
Route 30, 1 August 1976, 2 larvae. Elk County: Ridgway, South Branch Island Run, 29 j
May 1977, 1 6 pupa. Fayette County: Seven Springs, Neals Run Spring, 2 June 1977, j
A 6 6:1 June 1977, 2 6 pupae, 2 9 pupae. Franklin County: Turnpike between Blue j
and Kittatinny Mountains, Trout Run, 12 June 1976, 1 S'; 18 May 1977, 2 larvae, 1
prepupa. Forest County: Kellettville, Ross Run, 5-6 June 1976, 1 larva, 1 6: Tionesta,
small stream at Camp Tionesta, 6 June 1976, 1 6: Truemans Corner, Minister Creek,

5 June 1976, 1 prepupa. Fulton County: Emmaville, Chestnut Bridge Hollow, 28-29
May 1977, 1 larva, 1 ri , 1 9 : 1 July 1977, 2 6 pupae, 2 6 6: McConnellsburgh, Aughwick |
Creek, 30 June 1977, 2 6 6. Lancaster County: New Holland Watershed Area, 4-5 June i
1977, 5 larvae, 3 pupae, 2 S J ; 26-27 June 1977, 1 9 pupa, 3 ri ri , 9 9 9 ; 16-17 July I
1977, 3 (3 (3 , 4 9 9 . Pike County: Milford, Sawkill Creek, 1 1 June 1977, ! prepupa. Tioga j

1979

Weaver and Sykora — Pennsylvanian Rhyacophila

419

County; Ansonia, Colton Point, Right Branch Run, 1 July 1978, 8 11 99. Warren

County; Cobham, Rams Horn Spring, 24-25 July 1976, 2 6 6 2 9 9; 18 April 1977, 2
larvae. Westmoreland County; Laughintown, Furnace Run, 14-15 July 1976, 1 c? ; Rec-
tor, Powdermill Nature Reserve, Maul Spring, 28 January 1975, 51 larvae; 6 February
1975, 2 larvae; 28 February 1975, 20 larvae; 27 March 1975, 37 larvae; 24 April 1975, 34
larvae; 6 May 1975, 3 larvae, 3 prepupae, 3 pupae; 22 May 1975, 44 larvae, 8 prepupae,
26 pupae; 3 1 May 1975, 3 d c? , 3 9 9 ; 19-20 June 1975, 46 larvae, 8 prepupae, 18 pupae,
102 c5c5, 66 9 9; 16-17 July 1975, 12 larvae, 2 pupae, 23 d c3 , 50 9 9 ; 13-14 August
1975, 21 larvae, 1 prepupa, 18 dd, 21 9 9; 10-11 September 1975, 39 larvae, 1 d, 4
9 9; 11 October 1975, 54 larvae; 5 November 1975, 52 larvae; 3 December 1975, 93
larvae; 3 January 1976, 38 larvae; 6 May 1977, 3 larvae; 22 January 1977, 4 6 pupae,
6 9 pupae, 16 66, 14 9 9; 1 July 1977, 2 prepupae, 4 6 pupae, 2 9 pupae. New
Jersey. Sussex County; north of High Point State Park near Route 23, Clove Run, 19
May 1977, 1 prepupa. Stokes State Forest, small stream near Sawmill Road, 4 April
1977, 1 larva; 30 April 1977, 15 larvae, 7 prepupae; 1 1 June 1977, 2 larvae, 3 prepupae,
3 6 pupae, 6 9 pupae, 1 9 .

lobifera Group
Rhyacophiia lobifera Betten

Rhyacophila lobifera Betten, 1936 (male and female genitalia, larva). Ross, 1944 (male
and female genitalia, larva). Flint, 1962 (larva). Ross, 1956 (male genitalia). Schmid,
1970 (male genitalia).

This lone member of the lobifera group occurs throughout the Ohio
River Valley and the central United States. It is recorded for the first
time from Pennsylvania (Fig. 2) while other state records include Illi-
nois, Kentucky, Ohio, Oklahoma, and Tennessee. In Pennsylvania,
only one specimen was collected from the riffle area of a stream ap-
proximately 6 m wide. However, Ross (1944) mentioned that they were
common in small springs and temporary streams. It is probable that
this is their definitive habitat as Stern and Stern (1969) found many
larvae amid the moss and algae in a springbrook. In Kentucky, Resh
et al. (1975) found its flight period to be limited to two weeks in late
April and early May.

Material examined. — -Pennsylvania. Armstrong County: Vandergrift, Carnahan
Run, 16 March 1975, 1 larva.

sibirica Group
Rhyacophila atrata Banks

Rhyacophila atrata Banks, 1911. Ross, 1956 (male genitalia). Flint, 1962 (larva).

This montane species is indigenous to the Appalachian Mountains
of Massachusetts, North Carolina, New Hampshire, and New York.
It is also a new state record for Pennsylvania where adults have been
collected from three counties in May and June (Fig. 9). Flint (1962)
discovered that the larvae live in riffles of streams 1.8 to 4.8 m wide.

Material examined. — Pennsylvania. Forest County: Kellettville, Ross Run, 5-6
June 1976, 1 <5 , 1 9 . Franklin County: Turnpike between Blue and Kittatinny Moun-

420

Annals of Carnegie Museum

VOL. 48

Fig. 9. — Distribution of the species of Rhyacophila of the sibirica group.

tains. Trout Run, 12 June 1976, 1 c?, 1 9. Westmoreland County: Rector, Powdermill
Nature Reserve, White Oak Run, 31 May 1975, 1 S .

Rhyacophila manistee Ross

Rhyacophila manistee Ross, 1938fl (male genitalia). Flint, 1962 (larva).

This species has been recorded previously from Michigan, New
Hampshire, and New York. This caddisfly is recorded here for the
first time from Pennsylvania where we believe it to be relatively rare
(Fig. 9). The larvae live in meandering trout streams approximately

4.8 to 5.5 m wide (Flint, 1962). Adults have been collected in late May.

Material examined. — Pennsylvania. Westmoreland County: Rector, Powdermill
Nature Reserve, White Oak Run, 31 May 1975, 2 SS , 1 9.

Rhyacophila melita Ross

Rhyacophila melita Ross, 1938(r/ (male genitalia). Ross, 1956 (male genitalia). Flint, 1962
(larva). Roback, 1975 (larva).

This species has been recorded from Michigan, New Hampshire,
New York, and Vermont. R. melita probably occurs in Pennsylvania,
because it has been collected in New Jersey (new state record) only
a few miles from the Pennsylvania state line (Fig. 9). These immatures
were discovered in a large, clear mountain stream approximately 9 m
wide, only a few km east of the Appalachian Trail.

1979

Weaver and Sykora — Pennsylvanian Rhyacophila

421

Material examined. — New Jersey. Sussex County: north of High Point State Park,
Clove Run, 16 April 1977, 3 larvae.

Rhyacophila minor Banks

Rhyacophila minora Banks, 1924 (lapsus). Ross, 1956 (male genitalia). Flint, 1962 (lar-
va). Roback, 1975 (larva).

This is the most familiar species of the siberica Group in eastern
North America, It has been recorded previously from Maine, Massa-
chusetts, North Carolina, New Hampshire, Nova Scotia, New York,
South Carolina, West Virginia, and Tennessee. It is herein recorded
for the first time from Pennsylvania where it is relatively common (Fig.
9). Many immature s were collected from Maul Spring, but other rec-
ords indicate that it inhabits larger streams as well.

Material examined.— VEnnsyicvAniA. Beaver County: Raccoon Creek State Park, Min-
eral Run, 30 May 1976, 2 6 S , 1 9. Fayette County: Seven Springs, Back Run, 2 June
1977, 2 6 6,3 9 9. Franklin County: Turnpike between Blue and Kittatinny Mountains,
Trout Run, 12 June 1976, 1 6. Forest County: Kellettviile, Ross Run, 5-6 June 1976,
1 6,2 9 9; Tionesta Station, Hunters Creek, 5 June 1976, 4 66, 9 9 9; Truemans
Corner, Minister Creek, 5 June 1976, 1 larva, 1 9 . Fulton County: Emmaville, Chestnut
Bridge Hollow, I July 1977, 2 66, 1 9; McConneilsburg, Aughwick Creek, 30 June
1977, 25 6 6,7 9 9. Pike County: Sawkill Creek, 1 May 1977, 5 prepupae, 2 pupae.
Somerset County: Kooser State Park, 7-8 June 1976, 1 6. Westmoreland County:
Laughintown, Furnace Ron, 7-8 June 1976, 1 ri, 1 9 ; Laurel Hill Summit, Spruce Run,
18 July 1976, 1 ri; Rector, Powdermill Nature Reserve, Maul Spring, 28 January 1975,
21 larvae; 6 February 1975, 1 larva; 28 February 1975, 16 larvae; 27 March 1975, 18
larvae; 24 April 1975, 14 larvae; 6 May 1975, 9 larvae, 1 prepupa; 19-20 June 1975, 2
larvae, 6 pupae, 4 rid; 16-17 July 1975, 3 riri, 3 9 9; 13-14 August 1975, 3 larvae, 3
6 6,3 9 9; 10 September 1975, 75 larvae; 11 October 1975, 93 larvae; 5 November
1975, 112 larvae; 3 December 1975, 69 larvae; 3 January 1976, 9 larvae; 23 June 1977,
10 ri ri , 8 9 9 ; White Oak Run, 15-16 May 1977, 1 ri , 2 9 9 ; 23-24 June 1977, 8 ri ri .

torva Group

Rhyacophila torva Hagen

Rhyacophila torva Hagen, 1861. Ross, 1956 (male genitalia). Flint, 1962 (larva). Schmid,
1970 (male genitalia).

This is the only species of the torva group (Schmid, 1970) and is
restricted to eastern Nearctica, where its occurrence is common
throughout the Appalachian Mountains. It is known from Maine, Mas-
sachusetts, North Carolina, New Hampshire, New Jersey, New York,
Quebec, Tennessee, Virginia, Vermont, Washington, D.C., and West
Virginia. This is the first report of this species from Pennsylvania,
where its larvae were found under large stones in a fast-flowing, limpid
stream approximately 6 m wide. Flint (1962) stated that they appear
to dwell in streams about 1.8 to 2.7 m in width. Mature larvae may be
collected in the spring and adults throughout the summer (Fig. 9).

422

Annals of Carnegie Museum

VOL. 48

Material examined. — Pennsylvania. Franklin County: Turnpike between Blue and
Kittatinny Mountains, Trout Run, 12 June 1976, 1 6, 1 9; 18 May 1977, 2 larvae, 1
prepupa. Forest County: Kellettville, Ross Run, 18-19 June 1976, 3 cJ . Fulton County:
McConneilsburg, Aughwick Creek, 30 June 1977, 2 c3 , 3 9 9. Somerset County: Laurel
Hill State Park, 23 June 1976, I 9. Tioga County: Ansonia, Colton Point, Right Branch
Run, 1 July 1978, 1 (5 , 2 9 9 . Westmoreland County: Laurel Hill Summit, Spruce Run,
22 August 1976, 1 larva; Laughintown, Furnace Run, 7-8 June 1976, 2 SS , 1 9; 14-15
July 1976, 8 (3(3, 10 9 9; 20-21 August 1976, 4 c3c3, 4 9 9.

Acknowledgments

We wish to express our sincere gratitude to Dr. Craig C. Black, the Director of the
Carnegie Museum of Natural History, who has encouraged this research and provided
facilities at the Powdermill Nature Reserve of the Carnegie Museum.

Appreciation is also expressed to Dr. Bradford Owen who provided light trap material
from Washington County, Pennsylvania.

Literature Cited

Banks, N. 1907. Descriptions of new Trichoptera. Proc. Ent. Soc. Washington, 8:117-
133.

. 1911. Descriptions of new species of North American neuropteroid insects.

Trans. American Ent. Soc., 37:335-360.

. 1914. American Trichoptera — notes and distiibutions. Canadian Ent., 46:149-

156, 201-205, 252-258, 261-268.

. 1924. Descriptions of new neuropteroid insects. Bull. Mus. Comp. Zook,

45:421-455.

Be li en, C. 1934. The caddisflies or Trichoptera of New York State. New York State
Mus. Bull., 292:1-576.

Carpenter, F. M. 1933. Trichoptera from the mountains of North Carolina and Ten-
nessee. Psyche, 40:32-47.

Etnier, D. a., and J. D. Way. 1973. New southeastern Trichoptera. J. Kansas Ent.
Soc., 46:422-430.

Flint, O. S., Jr. 1958. Descriptions of several species of Trichoptera. Bull. Brooklyn
Ent. Soc., 53:21-24.

. 1962. Larvae of the caddis fly genus Rhyacophila in eastern North America

(Trichoptera: Rhyacophilidae). Proc. U.S. Nat. Mus., 113:465-493.

. 1972. Three new caddisflies from, the southeastern United States. J. Georgia

Ent. Soc., 7:79-82.

Hagen, H. 1861. Synopsis of the Neuroptera of North America, with a list of the
South American species. Smithsonian Misc. Coll., 4:1-347.

Hyeand, K. 1948. New records of Pennsylvania caddis flies (Trichoptera). Ent. News,
59:38-40.

Miene, L. j. 1936. Studies of the North American Trichoptera. Cambridge, Massa-
chusetts, 3:56-128.

Milne, M. J. 1938. The ‘‘Metamorphotype Method” in Trichoptera. J. Nev/ York Ent.
Soc., 46:435-437.

Morse, J. C., and H. H. Ross. 1971. Two new species of Rhyacophila from eastern
North America. J. Kansas Ent. Soc., 44:403-405.

Neves, R. 1977. The larval identity of Rhyacophila acutiloha and R. Carolina (Tri-
choptera, Rhyacophilidae). J. Kansas Ent. Soc., 50:148.

Provancher, M. a. 1878. Additions et corrections aux Neuropteres de la Province de
Quebec. Le Nat. Canadien, 10:124-147, 367-369.

1979

Weaver and Sykora — Pennsylvanian Rhyacophila

423

Resh, V. H., K. H. Haag, and S. E. Neff. 1975. Community structure and diversity
of caddisfly adults from the Salt River, Kentucky. Env. Ent., 4:241-=253.

Roback, S. S. 1975. New Rhyacophilidae records with some water quality data. Proc.
Acad. Nat. Sci. Philadelphia, 127:45-50'.

Ross, H. H. 1938fl. Descriptions of Nearctic caddis flies (Trichoptera) with special
reference to the Illinois species. Illinois Nat. Hist. Surv. Bull., 21:101-183.

. 1938/1. Lectotypes of North American caddis flies in the Museum of Compar-
ative Zoology. Psyche, 45:1-61.

— . 1939. New species of Trichoptera from the Appalachian region. Proc. Ent.

Soc. Washington, 41:65-72.

— . 1941. Descriptions and records of North American Trichoptera. Trans. Amer-

ican Ent. Soc., 67:35-126.

— . 1944. The caddis flies, or Trichoptera, of Illinois. Illinois Nat. Hist. Surv. Bull.,

23:1-326.

— — . 1948. New nearctic Rhyacophilidae and Philopotamidae (Trichoptera). Ann.
Ent. Soc. America, 41:17-26.

— — — . 1956. Evolution and classification of the mountain caddisflies. Urbana, Illinois,
1-213.

Roy, D., and P. P. Harper. 1975. Nouvelles mentions de trichopteres du Quebec et
description de Limnephilus nimmoi sp. oov. (Limnephilidae). Canadian J. ZooL,
53:1080-1088.

Schmid, F. 1970. Le genre Rhyacophila et la famille des Rhyacophilidae (Trichoptera).
Mem. Soc. Ent. Canada, 66:1-230.

1974. Un Rhyacophila nearctique meconnu (Trichoptera, Rhyacophilidae). Le

Nat. Caeadien, 101:933-934.

Sherberger, F. F., and J. B. Wallace. 1971. Description of the larval stage of
Rhyacophila vuphiphes Milne (Trichoptera: Rhyacophilidae). New York Ent. Soc.,
79:43-44.

Smith, S. D. 19'68. The Rhyacophila of the Salmon River drainage of Idaho with
special reference to larvae. Ann. Ent. Soc. America, 61:655-674.

Stern, M. S., and D. H. Stern. 1969. A limnological study of a Tennessee cold
spriegbrook. Amer. Midland Nat., 82:62-82.

Sykora, J. L., and J. S. Weaver III. 1976. A new species of Rhyacophila (Trichop-
tera, Rhyacophilidae) from western Pennsylvania. Ann. Carnegie Mus., 46:29-32.
Ulmer, G. 1907. Trichopteren. In, Selys-Longchamps, Collections zoologiques du
Baron Edm. de Selys-Longchamps catalogue systematique et descriptif, Fasc VI (Pt.
1):1-102.

Walker, F. 1852. Catalogue of the specimens of neuropterous insects in the collection
of the British Museum. London, 1:1-192.

Wiggins, G. B. 1977. Larvae of the North American caddisfly genera (Trichoptera).
Toronto Univ., Toronto Press, 401 pp.

Back issues of many Annals of Carnegie Museum articles are
available, and a few early complete volumes and parts are listed
at half price. Orders and inquiries should be addressed to:
Publications Secretary, Carnegie Museum, 4400 Forbes Avenue,
Pittsburgh, Pa. 15213.

S

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ISSN 0097-4463

<07, 73

ANNALS

o/ CARNEGIE MUSEUM

CARNEGIE MUSEUM OF NATURAL HISTORY

4400 FORBES AVENUE ® PITTSBURGH, PENNSYLVANIA 15213
VOLUME 48 30 NOVEMBER 1979 ARTICLE 23

CHECKLIST OF CALIFORNIA MAMMALS
Daniel F. Williams^

Research Associate, Section of Mammals

Abstract

A checklist, consisting of 216 species of mammals found within California, is pre-
sented. Changes in nomenclature subsequent to the list of Jones et al. (1975, Revised
checklist of North American mammals north of Mexico) are documented.

Introduction

This checklist is intended to provide an updated reference to the
mammals of California. Since the publication of Ingles’ (1965) Mam-
mals of the Pacific States,” several systematic studies have resulted in
changes in the names of species of mammals found in the wild in
California. Additionally, five exotic species may have become estab-
lished, and the wolf is now considered to be extinct within the State
(McCullough, 1967). This brings the number of species to become ex-
tinct within California in the last century to three.

The vernacular names used here are primarily based upon Jones et
al. (1975). Although a single vernacular name should apply to all pop-
ulations of a species, there is more than one name in common use
for some of the mammals found within California. Cetaceans are most
notable in this respect. For a few species, I have adopted vernacular
names different from those of Jones et al. (1975). In most of these
cases, the names selected are widely known and generally accepted

^ Department of Biological Sciences, California State College Stanislaus, Turlock, Cal-
ifornia 95380.

Submitted for publication 5 June 1979.

425

I

426 Annals of Carnegie Museum vol. 48 j

among mammalogists. In a few others, such as the hares of the genus
Lepus, and Dipodomys nitratoides, more appropriate or less confusing
names are used in preference to those in common use. [

The scientific names in the list generally follow those of Jones et al. I
(1975). The New World rats and mice are regarded as a subfamily of ||
the family Muridae, two orders of cetaceans are recognized, and the |i
pinnipeds are treated as a suborder of the order Carnivora. Systematic ;
studies that have resulted in changes in taxonomy subsequent to the *
list of Jones et al. (1975), and other departures from their nomenclature
are documented below. i

Hennings and Hoffmann (1977) showed that the dusky shrew {Sorex ;
obscurus) is specifically distinct from Sorex vagrans, and pointed out S
that its proper name is S. monticolus. They also implied that 5. sin- ji
uosus is a subspecies of S. ornatus. I concur with their view on this i
latter point, primarily because of the evidence given by Rudd (1955). il
I have retained S. willetti as a species since its status has not yet been ;
investigated. I recognize Myotis occultus as a species distinct from M. i
lucifugus despite the works of Findley and Jones (1967) and Barbour |i
and Davis (1970). This decision is based primarily upon the evidence II
given by Findley (1972). Hibbard (1963) presented compelling evidence |
that the pygmy rabbit, Brachylagus idahoensis, is not congeneric with '
the cottontails {Sylvilagus). Nadler et al. (1977) regarded Eutamias as
a subgenus of Tamias. Callahan (1977) demonstrated that Tamias
{Eutamias) merriami consisted of two species, T. merriami and T.
obscurus. Patton et al. (1976) showed that Dipodomys californicus is j
specifically distinct from D. heermanni. I agree with Johnson’s (1973) i'
recognition of Arborimus as a genus distinct from Phenacomys, and
despite the lack of clarity in his discussion concerning the classification I
of the white-footed vole, I have included albipes in the genus Arbor-
imus. I followed Orr (1972) in recognizing Globicephala scammonii.
Van Gelder (1977, 1978) presented formidable arguments for the inclu- ;
sion of the foxes in the genus Canis. '

Methods and Acknowledgments

In the following list, species not native to California are indicated by an asterisk I
(*) following their scientific name. Some of the introduced species may not have estab-
lished viable populations in the wild; those with uncertain status are indicated by a
question mark (?) following their vernacular name.

Dr. Aryan I. Roest supplied information on the status of several introduced species. |
Clare Brooks aided me in preparing drafts of this paper.

Checklist

Order MARSUPiALiA—marsupials

Family Didelphidae— -New World opossums
Didelphis virginiana*

Virginia opossum

1979

Williams—California Mammals

427

Order Insectivora— insectivores
Family Soricidae— shrews

Sorex lyelli ............................... Mt. Lyell shrew

Sorex vagrans .............................. vagrant shrew

Sorex monticolus ............................ dusky shrew

Sorex pacificus .............................. Pacific shrew

Sorex ornatus ............................... ornate shrew

Sorex willetti ......................... Santa Catalina shrew

Sorex tenellus ................................ Inyo shrew

Sorex palustris ................................ water shrew

Sorex bendirii ............................... marsh shrew

Sorex trowbridgii ...................... Trowbridge’s shrew

Sorex merriami .......................... Merriam’s shrew

. Notiosorex crawfordi ......................... desert shrew

Family Talpidae— moles

Neurotrichus gibbsii ........................... shrew-mole

Scapanus townsendii ..................... Townsend’s mole

Scapanus orarius .............................. coast mole

Scapanus latimanus ..................... broadTooted mole

Order Chiroptera— bats
Family Phyllostomatidae— leaf-nosed bats
Macrotus californicus .............. California leaf-nosed bat

Choeronycteris mexicana .................. long-tongued bat

Family Vespertilionidae— mouse-eared bats
Myotis lucifugus ........................ little brown myotis

Myotis occultus ............................ Arizona myotis

Myotis yumanensis ........................... Yuma myotis

Myotis velifer ................................. cave myotis

Myotis evotis ............................ long-eared myotis

Myotis thysanodes .......................... fringed myotis

Myotis volans .......................... long-legged myotis

Myotis californicus ....................... California myotis

Myotis leibii ........................... small-footed myotis

Lasionycteris noctivagans .................. silver-haired bat

Pipistrellus hesperus .................... western pipistrelle

Eptesicus fuscus ............................ big brown bat

Lasiurus borealis ................................. red bat

Lasiurus cinereus ............................... hoary bat

Lasiurus ega .......................... southern yellow bat

Euderma maculatum ........................... spotted bat

Plecotus townsendii .............. Townsend’s big-eared bat

Antrozous pallidus .............................. pallid bat

Family Molossidae— free-tailed bats

Tadarida brasiliensis ............... Brazilian free-tailed bat

428

Annals of Carnegie Museum

VOL. 48

Tadarida femorosacca .............. pocketed free-tailed bat

Tadarida macrotis ....................... big free-tailed bat

Eumops perotis ........................ western mastiff bat

Order Lagomorpha— lagomorphs
Family Ochotonidae— pikas

Ochotona princeps ................................... pika

Family Leporidae— hares and rabbits
Brachylagus idahoensis ....................... pygmy rabbit

Sylvilagiis bachmani .......................... brush rabbit

Sylvikigus nuttcdlii ...................... NuttalFs cottontail

SylvUagus audubonii ...................... desert cottontail

Oryctolagus cuniculus"^ .................. European rabbit ?

Lepus americaniis .......................... seowshoe hare

Lepus townsendii .......................... white-tailed hare

Lepus californicus ........................ black-tailed hare

Order Rodentia— rodents

Family Aplodontidae— mountain beaver
Aplodontia rufa ........................... mountain beaver

Family SciURiDAE—squirrels

Marmota flaviventris ................... yellow-bellied marmot

Spermophilus townsendii ........ Townsend’s ground squirrel

Spermophilus beldingi ............. Beldieg’s ground squirrel

Spermophilus mohavensis ........... Mohave ground squirrel

Spermophilus tereticaudus ...... round-tailed ground squirrel

Spermophilus variegatus ...................... rock squirrel

Spermophilus beecheyi ........... California ground squirrel

Spermophilus lateralis ........ goldee-mantied ground squirrel

Ammospermophilus leucurus . . . white-tailed antelope squirrel
Ammospermophilus nelsoni .... San Joaquin antelope squirrel

Tcunias alpinus ........................... alpine chipmunk

Tamias minimus ........................... least chipmunk

Tamias amoenus ..................... yellow-pine chipmunk

Tamias panamJntinus , ................... Panamint chipmunk

Tamias umhrinus ......................... Uinta chipmunk

Tamias speciosus ...................... lodgepole chipmunk

Tamias siskiyou ........................ Siskiyou chipmunk

Tamias senex ............................ Allen’s chipmunk

Tamias ochrogenys ............... yellow-cheeked chipmunk

Tamias quadrimaculatus ............... long-eared chipmunk

Tamias sonomae ........................ Sonoma chipmunk

Tamias merriami ...................... Merriam’s chipmunk

Tamias ohscurus ....................... chaparral chipmunk

Sciurus griseus ....................... western gray squirrel

Sciurus carolinensis'^ ......................... gray squirrel

1979

Williams — California Mammals

429

Sciurus niger* fox squirrel

Tamiasciurus douglasii ................... Douglas’ squirrel

Glaucomys sabrinus ................. northern flying squirrel

Family Geomyidae— pocket gophers
Thomomys bottae .............. southwestern pocket gopher

Thomomys talpoides ............... northern pocket gopher

Thomomys townsendii ........... Townsend’s pocket gopher

Thomomys montlcola ............... mountain pocket gopher

Thomomys mazama ................. western pocket gopher

Family Heteromyidae— heteromyid rodents
Perognathus longimembris .............. little pocket mouse

Perognathus inornatus ............ San Joaquin pocket mouse

Perognathus parvus .............. Great Basie pocket mouse

Perognathus xanthonotus ........ yellow-eared pocket mouse

Perognathus aiticola ............. white-eared pocket mouse

Perognathus formosus ............. long-tailed pocket mouse

Perognathus baileyi ................. Bailey’s pocket mouse

Perognathus penicillatus ............... desert pocket mouse

Perognathus fallax ................ San Diego pocket mouse

Perognathus californicus ............ California pocket mouse

Perognathus spinatus spiny pocket mouse

Microdipodops megacephalus ......... dark kangaroo mouse

Microdipodops palUdus ................ pale kangaroo mouse

Dipodomys heermanni ............. Heermaen’s kangaroo rat

Dipodomys panamintinus ............ Paoamint kangaroo rat

Dipodomys stephensi ................ Stephen’s kangaroo rat

Dipodomys ingens ...................... giant kangaroo rat

Dipodomys merriami ............... Merriam’s kangaroo rat

Dipodomys nitratoides ............. San Joaquin kangaroo rat

Dipodomys ordii ........................ Ord’s kangaroo rat

Dipodomys agilis ...................... Pacific kangaroo rat

Dipodomys venustus ............. narrow-faced kangaroo rat

Dipodomys elephantinus ............. big-eared kangaroo rat

Dipodomys microps ............. chisel-toothed kangaroo rat

Dipodomys deserti ..................... desert kangaroo rai

Dipodomys californicus .............. California kangaroo rat

Family CASTORiDAE~=~beaver

Castor canadensis ................................. beaver

Family Muridae— mice and rats
Reithrodontomys megalotis .......... western harvest mouse

Reithrodontomys raviventris ........ salt marsh harvest mouse

Peromyscus californicus .................. California mouse

Peromyscus crinitus ......................... canyon mouse

Peromyscus eremicus ........................ cactus mouse

430

Annals of Carnegie Museum

VOL. 48

Peromyscus maniculatus deer mouse

Peromyscus boylii brush mouse

Peromyscus truei pinyon mouse

Onychomys leucogaster ......... northern grasshopper mouse

Onychomys torridus ............ southern grasshopper mouse

Sigmodon hispidus hispid cotton rat

Sigmodon arizonae Arizona cotton rat

Neotoma albigula white-throated wood rat

Neotoma lepida ........................... desert wood rat

Neotoma fuscipes dusky-footed wood rat

Neotoma cinerea bushy-tailed wood rat

Clethrionomys occidentalis western red-backed vole

Phenacomys intermedius heather vole

Arborimus albipes ........................ white-footed vole

Arborimus longicaudus red tree vole

Microtus oregoni creeping vole

Microtus montanus montane vole

Microtus californicus California vole

Microtus townsendii Townsend’s vole

Microtus longicaudus long-tailed vole

Lagurus curtatus sagebrush vole

Ondatra zibethicus muskrat

Rattus norvegicus"^' Norway rat

Rattus rattus'^' black rat

Mus musculus* house mouse

Family Zapodidae— jumping mice

Zapus princeps western jumping mouse

Zapus trinotatus Pacific jumping mouse

Family Erethizontidae — New World porcupines

Erethizon dorsatum porcupine

Family Capromyidae — capromyid rodents
Myocastor coy pus"'" nutria ?

Order Odontoceti— toothed whales
Family Ziphiidae— beaked whales
Berardius bairdii ..................... Baird’s beaked whale

Mesoplodon stejnegeri North Pacific beaked whale

Mesoplodon carlhubbsi Moore’s beaked whale

Ziphius cavirostris goose-beaked whale

Family Physeteridae — sperm whales

Physeter catodon sperm whale

Kogia breviceps pygmy sperm whale

Family Delphinidae — porpoises and dolphins

Stenella graffmani Eastern Pacific spotted dolphin

Stenella caeruleoalba blue and white dolpin

1979

Williams— California Mammals

431

Delphinus delphis ...............

Tursiops gilli .....................

Lagenorhynchus obliquidens . . . . .
Lissodelphis borealis ...........

Orcinus orca ...................

Grampus griseus ................

Pseudorca crassidens ............

Globkephala scammonii .........

Steno bredanensis ...............

Phocoena phocoena .............

Phocoenoides dalli ..............

Order Mysticeti— baleen whales
Family Eschrichtidae— gray whale
Eschrichtius robustus ............

Family Balaenopteridae— rorquals
Balaenoptera physalus ...........

Balaenoptera borealis ...........

Balaenoptera acutorostrata ......

Balaenoptera musculus ..........

Megaptera novaeangUae .........

Family Balaenidae— right whales
Balaena glacialis ................

Order Carnivora— carnivores
Family Canidae— canids
Canis iatrans ...................

Cams lupus ....................

Canis vulpes ....................

Cams macrotis .................

Canis cinereoargenteus ..........

Canis littoralis ..................

Family Ursidae— bears

Ursus americanus ...............

IJrsus arctos ...................

Family Procyonidae— procyoeids
Bassariscus astutus .............

Procyon lotor ...................

Family Mustelidae— mustelids
Martes americana ...............

Martes pennanti ................

Mustela erminea ................

Mustela frenata .................

Mustela vison ..................

Gtt/o gw/o ......................

Taxidea taxus ..................

.......... common dolphin

. Pacific bottle-nosed dolphin
Pacific white-sided dolphin
northern right-whale dolphin
.............. killer whale

........... Risso’s dolphin

......... false killer whale

. . North Pacific pilot whale
. , . . . rough-toothed dolphin
.......... harbor porpoise

........... Dali’s porpoise

........ gray whale

......... fin whale

......... Sei whale

...... mieke whale

........ blue whale

hump-backed whale

northern right whale

............ coyote

. gray wolf (extinct)
........... red fox

............ kit fox

.......... gray fox

. . . insular gray fox

........ black bear

grizzly bear (extinct)

........... ringtail

.......... raccoon

........... marten

............. fisher

........... ermine

. . long-tailed weasel
............. mink

......... wolverine

............ badger

432 Annals of Carnegie Museum vol. 48

Spilogale gracilis ................... western spotted skunk

Mephitis mephitis ........................... striped skunk

Lutra canadensis .............................. river otter

Enhydra lutris ................................... sea otter

Family Felidae— felids

Felis concolor .............................. mountain lion

Feiis rufus ........................................ bobcat

Family Otari idae— eared seals
Callorhinus ursinus ........................ northern fur seal

Arctocephalus townsendi ................ Guadalupe fur seal

Eumetopias jubata ....................... northern sea lion

Zalophus californianus .................. California sea lion

Family Phocidae— hair seals

Phoca vituUna ................................. harbor seal

Phoca fasciata ................................. ribbon seal

Mirounga angustirostris .............. northern elephant seal

Order Perissodactyla— odd-toed ungulates
Family Equidae— horses and asses
Equus asiniis"^' ................................. wild burro

Eqiius caballus'"''' ............................... wild horse

Order Artiodactyla— even-toed ungulates
Family Suidae— pigs

Siis scrofa"^ ...................................... wild pig

Family Cervidae— deer

Cervus elaphus .................................... wapiti

Cervus nnicolor* ................................ sambar ?

Cervus dama * ............................... fallow deer ?

Cervus axis* ................................... axis deer ?

Odocoileiis hemionus ............................ mule deer

Odocoileiis virginianus ................... white-tailed deer

Family Antilocapridae- — pronghorns
Antilocapra americana ......................... pronghorn

Family Bovidae— bovids

Ovis canadensis .......................... mountain sheep

Capra hircus* ..................................... goat ?

Ammotragus lervia* ......................... Barbary sheep

Hemitragus jemkihicus* .................... Himalayan tahr

Bison bison ................................. bison (extinct)

Literature Cited

Barbour, R. W., and W. H. Davis. 1970. The status of Myotis occulms. J. Mamm.,
51:150-151.

Callahan, J. R. 1977. Diagnosis of Eutamias obscurus (Rodentia: Sciuridae). J.
Mamm., 58: 1 88-201.

1979

Williams— -California Mammals

433

Findley, J. S. 1972. Phenetic relationships among bats of the genus Myotis. Syst.
Zool., 21:31-52.

Findley, J. S., and C. J. Jones. 1967. Taxonomic relationships of bats of the species
Myotis fortidens, M. lucifugus, and M. occultus. J. Mamm., 48:429-444.

Hennings, D., and R. S. Hoffmann. 1977. A review of the taxonomy of the Sorex
Vagrans species complex from western North America. Occas. Papers Mus. Nat.
Hist., Univ. Kansas, 68:1-35.

Hibbard, C. W. 1963. The origin of the P;5 pattern of Sylvilagus, Caprolagns, Oryc-
tolagns and Lepus. J. Mamm., 44:1-15.

Ingles, L. G. 1965. Mammals of the Pacific States. Stanford Univ. Press, Stanford,
California, 506 pp.

Johnson, M. L. 1973. Characters of the heather vole, Phenacomys, and the red tree
vole, Arborimiis. J. Mamm., 54:239-244.

Jones, J. K., Jr., D. C. Carter, and H. H. Genoways. 1975. Revised checklist of
North American mammals north of Mexico. Occas. Papers Mus., Texas Tech
Univ., 28: 1-14.

McCullough, D. R. 1967. The probable affinities of a wolf captured near Woodlake,
California. California Fish and Game, 53:146-153.

Nadler, C. F., R. S. Hoffmann, J. H. Honacki, and D. Pozini. 1977. Chromosomal
evolution in chipmunks, with special emphasis on A and B karyotypes of the sub-
genus Neotamias. Amer. Midland Nat., 98:343-353.

Orr, R. T. 1972. Marine mammals of California. Univ. California Press, Berkeley, 64
PP-

Patton, J. L., H. MacArthur, and S. Y. Yang. 1976. Systematic relationships of
the four-toed populations of Dipodomys heermanni. J. Mamm., 57:159-163.

Rudd, R. L. 1955. Population variation and hybridization in some California shrews.
Syst. Zool., 4:21-34.

Van Gelder, R. G. 1977. Mammalian hybrids and generic limits. Amer. Mus. Novi-
tates, 2635: 1-25.

— . 1978. A review of canid classification. Amer. Mus. Novitates, 2646:1-10.

Back issues of many Annals of Carnegie Museum articles are
available, and a few early complete volumes and parts are listed
at half price. Orders and inquiries should be addressed to:
Publications Secretary, Carnegie Museum, 4400 Forbes Avenue,
Pittsburgh, Pa. 15213.

r<^7 73

ISSN 0097-4463

ANNALS

0)' C A i<N E C I E K-l H S E LI iV1

CARNEGIE MUSEUM OF NATURAL HISTORY
4400 FORBES AVENUE ^ PITTSBURGH, PENNSYLVANIA 15213
VOLUME 48 ' ■ 30 NOVEMBER 1979 ARTICLE 24

EASTERN NORTH AMERICAN PLEISTOCENE
OCHOTONA (LAGOMORPHA: MAMMALIA)

John E. Guilday

Associate Curator, Section of Vertebrate Fossils

Abstract

Fossil remains of Ochotona sp. from four Appalachian cave deposits in Maryland,
West Virginia, and Virginia are compared with Recent North American O. princeps and
O. coUaris. They appear to be slightly smaller but the fossil material is insufficient for
subgeneric allocation. O. spangki from the late Pliocene of Oregon is discussed and its
type description emended.

Introduction

The Holarctic genus Ochotona comprises a closely related group of
perhaps 12 Palaearctic (Ellerman and Morrison-Scott, 1951) and two
(Hall and Kelson, 1959) Nearctic species. They are morphologically
conservative, differing among themselves in details of proportion, size,
and pelage. The dental pattern varies little throughout the known Plio-
cene to Recent History of the genus. Despite their basic anatomical
similarities some species of Ochotona, depending upon the degree of
local geographic isolation, break up into races described primarily on
the basis of pelage coloration. Ochotona princeps from the central
Rocky Mountain Cordillera has 36 currently recognized subspecies
(Hall and Kelson, 1959) reflecting the strong isolating role of post-
Pleistocene, low-latitude montane refugia. The northern North Amer-
ican O. coUaris or O. p. coUaris (see Yourigman, 1975) reported from
Alaska, Yukon, Northwest Territories, and northwestern British Co-
lumbia, in a boreal region where posUPleistocene limiting factors are

Submitted for publication 22 May 1979.

435

436

Annals of Carnegie Museum

voE. 48

less intense than in lower latitudes, does not exhibit this polymorphic

pattern.

All North American and most Palaearctic species are largely con-
fined to rocky skree, usually at higher altitudes, a circumstance re-
flecting perhaps more upon the distribution of skree than upon the
physiological limits of pikas. In Alaska, O. collaris occurs in dry alpine
tundra, or dry microhabitats within high boggy tundra meadows above
2,000 ft (Guthrie, 1973). Ochotona pusilla, occurring on the Russian
Steppes from approximately latitude 45°N to 55°N, often is found
. .in dense thickets, wheatfields, wild cherry brush, among weeds,
along fences in the village” (Ognev, 1940:88).

The Ochotonidae, once the most varied and successful lagomorphs,
first appeared in the fossil record in the Oligocene of Asia. They sub-
sequently spread to Europe, Africa, and North America. The genus
Ochotona, possibly of Asian origin, appeared in the Pliocene of Eu-
rope, Asia, and North America. Ochotona is the only pika known
from mid-latitude North America from late Pliocene (Hemphillian)
times to the present (Dawson, 1967). There is a gap in the known
North American Ochotona record of perhaps three million years, from |
O. spanglei from the Hemphillian McKay Reservoir local fauna of i

Oregon, to Ochotana sp. from the mid-Pleistocene (Irvingtonian) of |

the mid- Appalachians of eastern North America. All other North |
American records south of the sub-Arctic are from Rancholabrean to
Recent times in western North America.

A fossil ”ochotonid” was described by E. D. Cope (1871:93-94, fig.

20) from Port Kennedy Cave, Montgomery County, Pennsylvania, un- i

der the name Praotherinm palatinnm, later Lagomys palatinus (Cope, >

1899:209-210). The interpretation of this specimen, AMNH 8574, a |

partial palate with four molariform teeth on each side, has always been |

controversial. Cope himself was uneasy about its taxonomic affini- ;

ties. Dice (1923) examined the specimen and stated that it was not ,

Ochotona. It is definitely a young leporid (M. Dawson, personal com-
munication). Cope states that Lepus sylvatlcus Bach. {^Sylvilagus
sp.) was common in the Port Kennedy deposit. '

In order to assess the taxonomic affinities of the Appalachian ma-
terial, the holotype and only known specimen of O. spanglei was ex-
amined, a partial left jaw with P.rM^ (M^ mentioned in the diagnosis
has apparently been lost). The original diagnosis reads, “A pike [sicj
abou^the^ize of the living North American species. The character of
the P4-M3 is similar, but there is a rounded external lobe on P3. Pos- :

terior and anterior faces of this tooth are straight.” (Shotwell, |

1956:726-727). The diagnosis does not adequately describe the char- |

acteristics of the species and is here emended. The specimen is not |

near modern North American Ochotona in size, but is smaller than O. |

Table I. — Dental measurements, in mm, various Ochotona, Recent and fossil.

1979

Guilday — Pleistocene Ochotona

437

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438

Annals of Carnegie Museum

VOL. 48

Fig. 1. — X-rays, lingual aspect, anterior to left. Line indicates posterior limits of lower
incisor. Top: Ochotona spanglei. University of Oregon F-4083. Late Pliocene (Hemp-
hillian), P;5-M2 and root of incisor. Bottom: Ochotona princeps. Recent, CM 20198, full
dentition.

princeps in all dimensions (see Table 1). The details of the occlusal
pattern of P3 , as stated by Shotwell, do not serve to distinguish it from
other species of Ochotona. Ochotona spanglei does differ markedly
from all modern Ochotona in the position of the posterior end of the
lower incisor. The incisor extends back in the jaw to a position under
the mid“point of Mj. This can be seen in the X-ray (Fig. 1) and is
apparent externally by a swelling on the lingual aspect of the jaw. In
modern Ochotona the origin of the incisor has migrated forward to a
position under the P4/M1 junction. This forward migration is a contin-
uation of a trend noted in older, more primitive ochotonids where the
incisor is even more posterior than in O. spanglei and is apparently
a device to accommodate increasing hypsodonty of the cheek teeth
(M. Dawson, personal communication). Therefore, although the orig-

1979

Guilday — Pleistocene Ochotona

439

Fig. 2. — Ochotona sp. Cumberland Cave, Maryland. Anterior to right. Top: right maxilla
with USNM 12379; premaxillary fragment with partial right incisor, USNM un-

catalogued, ventral views. Center: partial right mandible, USNM 12380, labial view.
Bottom: right lower dentition with Pa-M.^, USNM 12380, occlusal view (from Dawson,
1967:301, Figs. 4-5).

440

Annals of Carnegie Museum

VOL. 48

inal diagnosis was inadequate, O. spanglei, by virtue of its small size
and incisor placement, is a well marked and primitive species of Och-
otona, with no obvious relationship to any particular modern species,
but nothing to disbar it from a direct ancestral position in the genus.

Eastern North America

Ochotona sp. is presently known from four mid-Appalachian sites—
Cumberland Cave, Maryland (lat. 39°41'N, long. 78 47'W), where it
was common, represented by complete upper and lower dentitions
(Fig. 2) and isolated teeth of at least 13 individuals; Trout Cave, West
Virginia (lat. 38°36'N, long. 79°22'W), jaw fragments and isolated
teeth; Rapp's Cave, West Virginia (lat. 37°58'N, long. 80°23'W), one
complete immature mandible; Jasper Saltpeter Cave, Virginia (lat.
36°46'N, long. 82°48'W), one upper molar and incisor fragment (see
list of specimens examined).

All known eastern fossil specimens were examined and compared
directly with 56 crania and lower jaws of O. princeps from Utah,
Colorado, Montana, and Alberta, and four Ochotona collaris from
Alaska. Only dental and mandibular comparisons were made due to
the fragmentary nature of the fossil material. The upper and lower
teeth were indistinguishable from those of modern O. princeps, which
does not necessarily imply conspecificity because of the lack of intra-
generic variation. The position of the lower incisor is as in O. princeps,
differing from the more primitive condition seen in O. spanglei.

The Cumberland Cave fossil material averages significantly smaller
in alveolar length of lower tooth row than Recent O. princeps (Stu-
dent’s t = 2.59, P = <.01— .02) but the observed range falls well within
that of modern adult O. princeps, so that the slightly smaller size of
the Appalachian specimens cannot be appreciated except in series
(Table 1). Given their geological age, their (now) isolated eastern dis-
tribution, and slight size difference, better fossil material may warrant
taxonomic recognition, but pending future discoveries the mid-Appa-
lachian Pleistocene Ochotona cannot be identified beyond the generic
level.

Discussion

Did the present North American species of Ochotona evolve in situ
from an O. spanglei stock, or represent a post-Pliocene invasion, or
invasions, of more progressive species of Ochotona from the Palaearc-
tic? The present distributional hiatus and morphological (but not
karyotypic, Youngman, 1975) distinction between O. collaris to the
north and O. princeps to the south in the Rocky Mountain Cordillera
suggests that more than one stock is represented, that the southern O.
princeps population antedates at least the Wisconsinan glaciation, and

1979

Guilday — Pleistocene Ochotona

441

that several glacial refugia may have been involved from which post-
glacial range adjustments took place, Beringian and western perigla-
ciaL

There is no pre-Wiscoesiean Pleistocene record of Ochotona, south
of Alaska, from the American West or Midwest. In the western mon-
tane areas this may be due to a lack of suitable preservation sites, but
in the numerous Great Plains Pleistocene faunas Ochotona is conspic-
uously absent (Hibbard et aL, 1965). The Great Plains served as an
ecological barrier to Ochotona throughout the Pleistocene, as it does
today.

The most plausible immigration route into the Appalachians for Och-
otona would have been from a northwesterly direction, circumventing
the Great Plains. To make this traverse the animal would have had to
spread across areas free of rocky talus, implying greater ecological
freedom than it now has in North America, perhaps as a consequence
of less competition from other similar sized herbivores such as the
many terrestrial sciurids persent at lower latitudes. Dawson (personal
communication) points out that the decline of the varied Neogene och-
otonids in post-Miocene times coincides with the proliferation of ar-
vicolids, and suggests that the voles may have been an important lim-
iting factor in ochotonid fortunes. The recent Ontario find of Ochotona
(Churcher and Dods, 1979) from an undated, possibly pre-Wisconsinan
cave breccia, may lie along this route. Surprisingly, however, the On-
tario specimen, a partial femur from Kelso Cave, in the Niagara es-
carpment, 45°30'N, 79°55'W, is from a large animal about the size of
the extinct O. whartoni Guthrie and Matthews, 1971, from the early
Pleistocene Cape Deceit local fauna, Alaska, and the Old Crow Basin
of the Yukon (Harington, 1978), and, on the basis of size alone, is
unrelated to the smaller Appalachian Ochotona sp. of Maryland, West
Virginia, and Virginia.

Ecology of Eastern Ochotona

Two of the four known mid-Appalachian sites containing Ochotona
have accompanying species that make some ecological inferences pos-
sible. The Rapp’s Cave specimen is a chance find of unknown pro-
venience by a private collector. The Jasper Saltpeter Cave specimens
were in sediments disturbed by Civil War mining activities and the
meager accoro.paeyieg small mammal fauna is both chronologically
suspect and noncommittal. However, Cumberland Cave, Maryland
(Gidley and Gazin, 1938; Guilday and Handley, 1967; Holman, 1977;
Van der Meulen, 1978; Zakrzewski, 1975), and Trout Cave, West Vir-
ginia (Guilday, 1971; Zakrzewski, 1975), have extensive accompanying
faunas. On the evidence primarily of arvicoiid dental morphology, the
Cumberland Cave local fauna has been allocated to the Irvingtonian,

442

Annals of Carnegie Museum

VOL. 48

ca. 700,000 years BP, slightly older than the late Kansan Cudahy fauna
(Van der Meulen, 1978). Ochotona is not present in any of the nu-
merous Wisconsinan to early Holocene cave faunas from the Appa-
lachians (Guilday et al., 1977, 1978) and was apparently gone from
eastern North America by Rancholabrean times.

Present with Ochotona at Cumberland and Trout caves were sev-
eral vertebrates of western affinities; a thomomine pocket gopher Tho-
momys potomacensis and an ictidomyine ground squirrel Spermophi-
lus cf. tridecemlineatus; and at Cumberland Cave a coyote Canis
priscolatrans and a badger Taxidea marylandica. An extensive am-
phibian and reptile fauna of at least 30 species including one of mid-
western affinities, the fox snake, Elaphe vulpina, has been reported
from Cumberland Cave (Holman, 1977). Cumberland Cave also pro-
duced a variety of large browsers and grazers, a horse Equus, a tapir
Tapirus, two peccaries Platygonus and Mylohyns, an ovibovid Eu~
ceratherium, two cervids Odocoileus , Cervus, a ground sloth Mega-
lonyx, and the mastodon Mammut. Two arboreal sciurids, Tamiasciu-
rus and Glaucomys, and a woodland zapodid, Napaeozapus, were
present. Voles of boreal affinities were present at both Trout and Cum-
berland csiYes—Phenacomys, Clethrionomys , Synaptomys {Micto-
mys). However, these boreal rodents were a rare element in the nu-
merically large vole sample from each site, which consisted primarily
of Microtus, Pedomys and Pity mys. A wolverine Gulo was present at
Cumberland Cave.

This partial listing of species from these two eastern -bear-

ing sites, Cumberland Cave and Trout Cave, suggests a more varied
fauna than the closed, largely oak-dominated forests of the central
Appalachians were capable of supporting in Holocene times, indicating
a greater variety in the ecological landscape. Such faunas could best
have been accommodated by a cool-temperate environment, relatively
drier than it is today. The presence of so many extinct species makes
it impossible to suggest a modern analog, but the broad analogy of a
varied grassland/deciduous-coniferous forest situation superimposed
upon the regional Ridge and Valley topography (Hunt, 1974) seems
most appropriate.

This environmental interpretation is not consistent with the usual
concept of what Ochotona 'dikes” based upon modern habitats of
North American pikas, implying either that the Eastern North Amer-
ican Irvingtonian form was indeed a separate species with its own set
of environmental requirements, and/or that Ochotona today is closely
confined to its rocky niche by interspecific pressures, and not by evo-
lutionary specialization-— that it lives where it does because of com-
petitive factors rather than physiological or anatomical strictures.

1979

Guilday — Pleistocene Ochotona

443

Acknowledgments

I thank Harold W. Hamilton and Allen D. McCrady, whose field efforts resulted in
the recovery of most of the Appalachian specimens here discussed; Hugh H. Genoways
and Susan McLaren, Section of Mammals, Carnegie Museum of Natural History, for
the loan of Recent O. princcps specimens, Don E. Wilson, National Museum of Natural
History, for specimens of O. collaiis, and Eric Gustafson, University of Oregon, for
the loan of the holotype specimen of O. spanglei. X-ray by Allen D. McCrady. I also
thank Alice M. Guilday and Elizabeth Hill for secretarial assistance, Mary R. Dawson
for sharing her ochotonid expertise, and C. S. Churcher, Royal Ontario Museum, for
a preprint of his paper on the Niagara Escarpment ochotonid.

Specimens Examined

Recent. — Ochotona collads, Alaska: USNM 148590, 157225, 157227, 512900. O.prin-
ceps, Utah: CM Recent Mammal no. 9469-9471, 9473, 9475, 9477-9484, 12021-12024,
14719-14720, 14726-14733, 16020, 16022-16024, 16026-16028, 16030, 16032, 20200-
20201, 20203-20204, 20607-20608; Colorado: 20193-20198; Montana: 22143-22144,
22146, 22374; Alberta: 22906-22908, 22912.

Pleistocene. — O. pusilla, Great Doward Cave, England, CM 12666. Ochotona sp.,
Jasper Saltpeter Cave, Virginia, CM 30264; Rapp's Cave, West Virginia, CM 24290;
Trout Cave, West Virginia, CM 12722, 12793, 12818, 12837, 12864, 12879; Cumberland
Cave, Maryland, USNM 7689, 7768-7770, 12378-12379, 12381, CM 20242-20243, 20247,
20249-20250, 20477, 20479-20480.

Pliocene. — O. spanglei, McKay Reservoir, Oregon, UOMNH F-4083.

Literature Cited

Churcher, C. S., and R. R. Dods. 1979. Ochotona and other vertebrates of possible
Illinoian age from Kelso Cave, Halton County, Ontario. Canadian J. Earth Sci., in
press.

Cope, E. D. 1871. Preliminary report on the Vertebrata discovered in the Port Kennedy
bone cave. Proc. Amer. Phil. Soc., 12:73-102.

. 1899. Vertebrate remains from Port Kennedy bone deposit. J. Acad. Nat. Sci.

Philadelphia, ser. 2, 11:193-267.

Dawson, M, R. 1967. Lagomorph history and the stratigraphic record. Pp. 287-316,
in Essays in paleontology & stratigraphy, Raymond C. Moore Commemorative
Volume (C. Teichert and E. L. Yochelson, eds.), Univ. Kansas, Dept. Geol. Spec.
Publ., 2:1-626.

Dice, L. R. 1923. Notes on Praotherium palatinum Cope. J. Mamm., 4:260.
Ellerman, j. R., and T. C. S. Morrison-Scott. 1951. Checklist of Palaearctic and
Indian mammals. British Mus. (Nat. Hist.), London, 810 pp.

Gidley, j. W., and C. L. Gazin. 1938. The Pleistocene vertebrate fauna from Cum-
berland Cave, Maryland. Bull. U.S. Nat. Mus., 171:1-99.

Guilday, J. E. 1971. The Pleistocene history of the Appalachian mammal fauna. Pp.
233-262, in The distributional history of the biota of the southern Appalachians,
Part III: Vertebrates (P. C. Hold, ed.), Virginia Poly. Inst, and State Univ., Re-
search Div. Mono., 4:ix + 1-306.

Guilday, J. E., H. W. Hamilton, E. Anderson, and P. W. Parmalee. 1978. The
Baker Bluff Cave deposit, Tennessee, and the late Pleistocene faunal gradient. Bull.
Carnegie Mus. Nat. Hist., 11:1-67.

Guilday, J. E., and C. O. Handley, Jr. 1967. A new Peromyscus (Rodentia: Cri-
cetidae) from the Pleistocene of Maryland. Ann. Carnegie Mus., 39:91-103.

444

Annals of Carnegie Museum

VOL. 48

Guilday, J. E., P. W. Parmalee, and H. W. Hamilton. 1977. The Clark's Cave
bone deposit and the late Pleistocene paleoecology of the central Appalachian
Mountains of Virginia. Bull. Carnegie Mus. Nat. Hist., 2:1-87.

Guthrie, R. D. 1973. Mummified pika {Ochotona) carcass and dung pellets from Pleis-
tocene deposits in interior Alaska. J. Mamm., 54:970-971.

Guthrie, R. D., and J. V. Matthews, Jr. 1971. The Cape Deceit fauna — early
Pleistocene mammalian assemblage from the Alaskan Arctic. Quaternary Res.,
1:474-510.

Hall, E. R., and K. R. Kelson. 1959. The mammals of North America. The Ronald
Press Co., New York, l:xxx -f- 1-546 -h 79; 2:viii -h 547-1083 + 79.

Harington, ( . R. 1978. Quaternary vertebrate faunas of Canada and Alaska and their
suggested chronological sequence. Syllogeus No. 15, The National Museum of Can-
ada, Ottawa, 105 pp.

Hibbard, C. W., C. E. Ray, D. E. Savage, D. W. Taylor, and J. E. Guilday.
1965. Quaternary mammals of North America. Pp. 509-525, in The Quaternary of
the United States (H. E. Wright, Jr. and D. D. Erey, eds.), Princeton Univ. Press,
922 pp.

Holman, J. A. 1977. The Pleistocene (Kansan) herpetofauna of Cumberland Cave,
Maryland. Ann. Carnegie Mus., 46:157-172.

Hunt, C. B. 1974. Natural regions of the United States and Canada. W. H. Freeman
& Co., San Francisco, xii + 725 pp.

Ognev, S. I. 1940. Mammals of the U.S.S.R. and adjacent countries. Vol. 4. Israel
Program for Scientific Translations, Jerusalem (1966).

Shotwell, j. a. 1956. Hemphillian mammalian assemblage from northeastern Oregon.
Bull. Geol. Soc. Amer., 67:717-738.

Van der Meulen, A. J. 1978. Microtiis and Pityniys (Arvicolidae) from Cumberland
Cave, Maryland, with a comparison of some New and Old World species. Ann.
Carnegie Mus., 47:101-145.

Youngman, P. M. 1975. Mammals of the Yukon Territory. Nat. Mus. Canada Publ.,
Zoo!., 10:1-192.

Zakrzewski, R. j. 1975. The late Pleistocene arvicoline rodent Atopomys. Ann. Car-
negie Mus., 45:255-261.

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