TEXAS TECH UNIVERSITY

Natural Science Research Laboratory

Occasional Papers

Museum of Texas Tech University

Number 328 2 October 2014

Sexual Dimorphism in the Ringtail (Bassariscus astutus) from Texas

Frederick B. Stangl, Jr., Sarah Henry-Langston, Nicholas Lamar, and Stephen Kasper

Abstract

Sexual dimorphism in the ringtail has been alluded to in the literature, but the phenomenon
has not been documented. We present a morphometric assessment of cranial and dental features
for 152 specimens of Bassariscus astutus flavus from Texas and contiguous regions of northern
Mexico. Sample composition relative to age and sex is biased heavily towards young adult
males, with juveniles and senile adults of both sexes poorly represented. Adult males averaged
larger than females, and were significantly so for 12 of 17 characters examined. No discernible
geographic variation was noted for the species in our study area.

Key words: Bassariscus astutus , cranial variation, ringtail, sexual dimorphism, Texas

Introduction

Cranial variation has been documented for some
species of the Procyonidae, such as the raccoon, Pro-
cyon lotor (Kennedy and Lindsay 1984; Ritke 1990;
Ritke and Kennedy 1993); mountain coatis, Nasuella
spp., (Helgen et al. 2009); and the kinkajou, Potosfla¬
vus (Kortlucke 1973). Yet, little comparable informa¬
tion is available for the ringtail ( Bassariscus astutus ),
a widespread and often common carnivore that ranges
across much of the southwestern United States and into
southern Mexico (Hall 1981; Baker 1999).

Males of the order Carnivora typically are larger
than their female conspecifics—a trend especially
pronounced in Mustelidae, Felidae, and Procyonidae
(Gittleman and Van Valkenburgh 1997). Nowak (1999)
stated that male procyonids generally are a fifth larger
and heavier than females, an observation supported by
each of the above cited studies except that of Helgen et
al. (2009) for a limited sample of Nasuella. Poglayen-

Neuwall and Toweill (1988) noted that the skulls of
male ringtails generally are larger than comparably
aged females, yet Baker (1999) remarked that little
significant sexual dimorphism exists for B. astutus.

Population studies of the ringtail are available for
Texas (Ackerson and Harveson 2006) and New Mexico
(Harrison 2012), and synopses of the natural history
of the species are provided by Poglayen-Neuwall and
Toweill (1988) and Gehrt (2003). However, the most
detailed morphometric assessment for B. astutus of
which we are aware is Hoffmeister’s (1986) presenta¬
tion of selected cranial measurements for 15 skulls
from Arizona, leading to his observation that males
averaged larger for all skull measurements except for
least interorbital breadth and Ml breadth. He suggested
that further studies likely would demonstrate that males
are significantly larger than females.

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Occasional Papers, Museum of Texas Tech University

This study presents the first comprehensive Texas, and to the extent that our sample size permits,
morphometric assessment of the ringtail. We attempt we address microgeographic variation within this north-
to detail variation by age and sex for the species in eastern most subspecies of the taxon, B. astutus flavus.

Materials and Methods

A total of 152 specimens (Appendix) of Bas-
sariscus astutus flavus from the collections of the
Department of Biology at Midwestern State University
(MWSU; n = 88) and the Museum of Texas Tech Uni¬
versity (TTU; n = 64) comprise the basis for this study.
A total of 17 cranial and mandibular measurements
(nine of which are illustrated in Fig. 1) were attempted
from each specimen: rostral breadth (across base of
upper canines), least interorbital breadth, postorbital
breadth, greatest zygomatic breadth, mastoid breadth,
length and width of auditory bullae, condylobasal
length, maxillary toothrow (C-M2) length, width and
height of foramen magnum, greatest breadth of Ml,
greatest breadth of ml, basal length and width of upper
canine, and basal length and width of lower canine. All
measurements were recorded with digital calipers to the
nearest 0.01 mm. Paired elements were taken from the
right side, unless missing or too damaged to assess.

Each specimen was classified by sex as recorded
on the collector’s tag. The few juveniles available,
judged on the basis of their diminutive and sometimes
fragmentary skulls and lack of complete adult dentition,
were not included in this study. All older specimens
were assigned to one of four relative age categories,
based on relative degrees of cranial ossification and
dental wear, as modified from Kortlucke’s (1973)
work on kinkajous, although our terminology varies
somewhat. For our purposes, juveniles lack their
complete adult dentition. Age class 1 (subadults) is
characterized by complete but unworn dentition and
basisphenoid unfused anteriorly and posteriorly; age
class 2 (young adults) have slightly worn dentition and
basisphenoid fused posteriorly; age class 3 (old adults)
possess moderately worn dentition and basisphenoid
fused anteriorly and posteriorly; and age class 4 (senile
adults) are marked by heavily worn dentition and basi¬
sphenoid sutures tightly fused to obliterated.

All computations were performed with NCSS,
Version 5.3 statistical package (Hintze 1990). Two¬

way analyses of variation (age, sex) were employed to
inspect for an interaction effect between age and sex,
and to assess secondary sexual dimorphism across age
classes. Age was found to be unimportant, permitting
the pooling of specimens regardless of age. However,
we judged the middle two age classes to be sufficiently
large (n = 80; 54 males, 26 females) to discard the
smaller samples comprised of the youngest (n = 9;
averaged smallest for most characters) and the oldest
age classes (n = 4; averaged largest for several char¬
acters). We then used ANOVAs to characterize sexual
dimorphism for each character of the pooled middle
two age classes.

One-way ANOVAs and Duncan’s multiple means
tests were applied to test each sex independently for
any microgeographic variation of individuals from
the middle two age classes within the study area. Our
sample originated entirely from within the geographic
range of B. astutus flavus (sensu Hall 1981), but en¬
compassed such a large and geographically variable
region that we sorted specimens by biotic province (as
figured by Blair 1950). These provinces are defined
by distinctive physiographic and climatic regimes, and
sufficient samples were available for four: Balconian
(broken limestone terrain of central Texas); Chihuahuan
(desert mountains and basins of mostly Trans-Pecos);
Texan (rugged Cross Timbers of north-central Texas);
and Kansan (mostly grassland and savanna, bisected
by Caprock Escarpment). The Tamaulipan sample
(subtropics of extreme southern Texas and contiguous
Tamaulipas, Mexico) was of insufficient size to include
in the geographic analysis. Because nearly a third of
our specimens lacked data regarding sex, we applied
discriminant function analysis to test the utility of sexu¬
ally dimorphic features in accurately predicting the sex
for those individuals not accompanied by such data.

Stangl et al.—Cranial Variation in Bassariscus astutus

3

Figure 1. Dorsal and ventral views of skull of an adult male specimen of Bassariscus astutus (MWSU 8802).
Representative measurements are: a) rostral breadth; b) interorbital breadth; c) postorbital breadth; d) zygomatic
breadth; e) maxillary toothrow; f) mastoid breadth; g) condylobasal length; and upper canine (h) and Ml (i), for which
dimensions were recorded. Other measurements not figured were dimensions of foramen magnum, auditory bullae,
lower canine, and ml. (Drawing by Nicholas Lamar.)

Results

Sexual dimorphism for Bassariscus astutus was
pronounced (Table 1). Only for postorbital breadth
did females average marginally larger than males,
albeit insignificantly so. For the remaining charac¬
ters, males averaged larger. These differences were
highly significant, with the exceptions of bullar width,
dimensions of foramen magnum, and breadth of Ml.
Discriminant function analysis of age classes 2 and 3
served to accurately predict the sex for 95% (74 of 78)
of the individuals. Misclassified specimens included
two animals listed as males (MWSU 7500, TTU 17409)
and two listed as females (TTU 1671, TTU 6663).

Males comprised the majority of known-sex
individuals (60 of 93), and outnumbered females for
each of the first three age categories. Age categories
and breakdown by sex were as follows: age class 1 (n =
13): 5 males, 4 females, 4 of unknown sex; age class 2

(n = 96): 42 males, 21 females, 33 of unknown sex; age
class 3 (n = 34): 12 males, 5 females, 17 of unknown
sex; and age class 4 (n = 9): 1 male, 3 females, 5 of
unknown sex. As noted above, there were no signifi¬
cant differences among each of the four age classes for
either sex, although subadults generally were smaller
than older aged specimens, and senile adults generally
larger than younger animals.

More than half of the specimens of B. astutus
initially analyzed in this study originated from the
Balconian (n = 88) Province. Remaining animals were
taken from the Chihuahuan (n = 40), Texan (n = 15),
Kansan (n = 7), and Tamaulipan (n = 1) provinces.
Samples were small for some provinces, but there was
no significant geographic variation noted of either sex
for any character.

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Occasional Papers, Museum of Texas Tech University

Table 1. Comparison of cranial and dental measurements (in mm) between sexes of adult (age classes 2, 3) ringtails
(Bassariscus astutus flavus) from Texas. Descriptive statistics are: sample size (n), mean, standard deviation (SD),
minimum and maximum measures, confidence intervals (C.L), and coefficient of variation (CV). Probability values
(P) derived from one-way ANOVAs; non-significant results indicated by n.s.

Sex (n)

Mean ± SD

Minimum-

maximum

95% C. I.

CV

P

Male (54)

14.9 ±0.6

Rostral breadth

13.6-16.3

14.7-15.1

4.2

Female (26)

13.6 ±0.8

12.6-15.5

13.2-13.9

5.8

<0.0001

Male (54)

16.6 ±0.8

Interorbital breadth

14.8-18.5

16.4-16.8

4.7

Female (26)

15.8 ±0.8

14.7-17.6

15.5-16.2

5.4

<0.0001

Male (54)

17.7 ±1.5

Postorbital breadth

13.5-22.0

17.3-18.1

8.3

Female (26)

18.0 ±1.7

14.3-21.5

17.3-18.7

9.6

n.s.

Male (53)

50.6 ±2.3

Zygomatic breadth

46.3-55.5

50.0-51.2

4.4

Female (26)

47.3 ±2.4

43.0-54.6

46.4—48.3

5.2

<0.0001

Male (53)

36.0 ± 1.0

Mastoid breadth

33.8-38.0

35.7-36.2

2.8

Female (26)

34.2 ± 1.0

32.3-36.7

33.8-34.6

2.9

<0.0001

Male (54)

12.6 ±0.5

Bullar length

11.6-13.7

12.4-12.7

3.8

Female (25)

12.2 ±0.5

11.4-12.9

12.0-12.4

3.8

<0.001

Male (54)

10.4 ±0.5

Bullar width

9.4-11.4

10.3-10.6

4.2

Female (26)

10.3 ±0.3

9.7-10.8

10.1-10.4

2.7

n.s.

Male (52)

79.4 ±2.2

Condylobasal length

74.1-83.4

78.8-80.0

2.7

Female (26)

76.2 ± 1.9

73.8-81.2

75.4-76.9

2.5

<0.0001

Male (50)

11.9 ± 0.6

Foramen magnum width

10.6-13.3

11.7-12.0

5.2

Female (26)

11.6 ± 0.5

10.7-12.5

11.4-11.8

4.3

n.s.

Stangl et al.—Cranial Variation in Bassariscus astutus

5

Table 1. (cont.)

Sex (n)

Mean ± SD

Minimum-max¬

imum

95% C. I.

CV

P

Male (50)

8.8 ±0.6

Foramen magnum height

7.7-10.4

8.6-8.9

6.7

Female (26)

8.8 ±0.6

7.7-10.6

8.6-9.0

6.5

n.s.

Male (47)

Maxillary toothrow length (C-M2)

31.7 ± 1.1 29.8-36.0 31.3-32.0

3.6

Female (25)

30.3 ±0.7

28.8-31.6

30.0-30.6

2.5

<0.0001

Male (50)

3.6 ±0.2

Upper canine basal length

3.1^1.3

3.6-3.7

6.1

Female (25)

3.2 ±0.2

2.9-3.7

3.2-3.3

6.1

<0.0001

Male (50)

2.7 ±0.1

Upper canine basal width

2.3-3.1

2.6-2.7

5.3

Female (25)

2.3 ±0.1

2.1-2.6

2.3-2.4

5.2

<0.0001

Male (52)

8.2 ±0.5

Ml breadth

12-9.2

8.0-8.3

5.8

Female (26)

8.0 ±0.4

7.1-8.7

7.9-8.2

5.1

n.s.

Male (51)

4.0 ±0.3

Lower canine basal length

3.4—4.8

3.9—4.1

7.0

Female (25)

3.5 ±0.2

3.0-3.9

3.4-3.6

5.9

<0.0001

Male (51)

2.9 ±0.3

Lower canine basal width

2.2-3.7

2.8-2.9

8.9

Female (25)

2.5 ±0.2

2.3-3.2

2.4-2.6

8.9

<0.0001

Male (53)

3.8 ±0.2

Breadth of ml

3.3^1.2

3.7-3.8

4.2

Female (25)

3.6 ± 0.1

3.4-3.9

3.6-3.7

3.8

<0.0001

6

Occasional Papers, Museum of Texas Tech University

Discussion

The documented sexual dimorphism for ring¬
tails is sufficiently pronounced to suggest that the
phenomenon is typical for the ringtail across its range.
Elucidation of other aspects of cranial variation must
await studies of larger sample sizes of greater spatial
and temporal representation. Attempts to fully char¬
acterize age variation are confounded by the dearth of
specimens representing the age extremes: juveniles
and senile adults. Few wild individual ringtails seem
to reach the 11-14 years-of-age reported for some cap¬
tives (Crandall 1974), as evidenced in the few senile
adults in our sample. Mortality rates for Bassariscus
astutus in nature are high, as evidenced by one Texas
study (Ackerson and Harveson 2006) where 6 of 17
tagged animals were lost to predation during the course
of their 15-month study. The great horned owl (Bubo
virginianus ) is thought to be a major predator (Poglay-
en-Neuwall and Toweill 1988), and larger carnivores
(e.g. coyote, Canis latrans ; bobcat, Lynx rufus ) are
common across the state. Juveniles are scarce in col¬
lections, as the young animals remain close to the den
and grow quickly—weaning occurs within 10 weeks of
birth, adult dentition acquired 10 weeks later, and adult
size is achieved by 30 weeks-of-age (Toweill and To¬
weill 1978; Poglayen-Neuwall and Toweill 1988). This
rather narrow window of availability to collectors also
coincides with late summer and early fall—a period in
the academic calendar of historical collecting inactivity,
as reflected in the paucity of such records for ringtails
of any age from the collections of Midwestern State
and Texas Tech universities (Stangl and Jones 1987).

Our own sample is comprised largely of speci¬
mens salvaged from fur trappers and as automobile
traffic fatalities. Population studies of B. astutus in
Texas (Ackerson and Harveson 2006) and contiguous
New Mexico (Harrison 2012) indicated occurrence
of the sexes in equal numbers, although our sample
was skewed heavily towards males. This increased
vulnerability likely emphasizes naive or inexperienced
yearlings and young adults of both sexes during disper¬
sal, but particularly males with their greater dispersal
distances, larger home ranges, and perhaps less cautious

behavior. Road-killed ringtails are not as frequently
encountered as one might expect for so common an
animal, but it is trapped easily and is often taken in traps
intended for more valuable furbearers (Schmidly 1984).

Hoffineister (1986) noted the need for a compre¬
hensive systematic review of B. astutus. Lack of any
discernible pattern of microgeographic variation from
within the geographic range of the nominal subspecies
B. a.flavus likely reflects continuous gene flow across
suitable habitat of a moderate-sized and vagile carni¬
vore. Adequate habitat exists from the Chihuahuan
Desert mountains and foothills of the Trans-Pecos to the
rocky terrain across central Texas and northward along
the rugged, wooded hills of the Cross Timbers. These
three areas represent our largest regional sample sizes,
and coincide with those parts of the state referenced by
Schmidly (2004) as supporting the largest populations.

Faunal treatments commonly offer a select series
of cranial measurements for included taxa, although
sample sizes generally are small, only a few measure¬
ments of unaged adults are usually provided, and se¬
lected measurements vary among authors. A literature
survey of representative skull measurements from
across the range of the species (Table 2) provides data
too limited for any taxonomic inferences, although they
are supportive of the suggestion that ringtails from our
study area are larger than animals from the Sonoran
Desert southwest, and smaller than those from southern
Mexico (Poglayen-Neuwall and Toweill 1988).

Finally, we noted during the course of this inves¬
tigation that there is some confusion in the literature
regarding the dental formula of B. astutus that could
adversely impact future studies. The P4 is a large tooth
that was interpreted incorrectly as an Ml by Elbroch
(2006) and perhaps others, perpetuating the original
error seemingly first applied by Poglayen-Neuwall
and Toweill (1988). Gehrt (2003) first reported this
discrepancy, but left the issue unresolved. As typical
for the family Procyonidae (Stains 1984), the correct
formula for the ringtail is: i 3/3, c 1/1, p 4/4, m 2/2 = 40.

Stangl et al.—Cranial Variation in Bassariscus astutus

7

Table 2. Selected available cranial measurements (means, to nearest 0.1 mm) of the ringtail ( Bassariscus astutus )
from the literature. Abbreviations are as follows: IOB, interorbital breadth; ZYG, zygomatic breadth; MAST, mastoid
breadth; CNB, condylobasal length; and MAXT, maxillary tooth row length.

Authority/subspecific taxon

n

IOB

ZYG

MAST

CNB

MAXT

Hoffmeister (1986) from Arizona

B. a. arizonensis

5 males

14.7

47.4

34.6

77.4

30.7

5 females

15.5

46.9

33.4

74.3

30.0

B. a. yumanensis

1 male

15.8

41.6

--

74.8

29.8

3 females

16.5

45.1

36.0*

73.9

30.2

Hall (1946) from Nevada

B. a. nevadensis

2 males

14.7

47.0

33.5

--

--

Verts and Carraway (1998) from Oregon

B. a. raptor 3 males

15.3

47.8

78.0

31.1

3 females

15.8

48.2

--

--

30.5

Durrant (1952) from Utah

B. a. nevadensis

1 male

13.9

46.7

32.4

74.3

--

1 female

14.4

44.4

32.9

75.6

--

Anderson (1972) from Chihuahua

B. a. consitus

1 unknown

14.8

47.6

--

73.9

--

Baker (1956) from Coahuila

B. a.flavus

3 males

16.5

44.2

-

-

31.0

Goodwin (1969) from Oaxaca

B. a. astutus

1 male

16.6

57.7

-

89.7

33.5

1 female

18.2

55.7

-

85.0

32.8

B. a. macdougalli

1 female

16.3

53.0

--

83.9

32.4

B. a. bolei

1 male

16.0

56.5

--

84.5

31.7

*n=l

8

Occasional Papers, Museum of Texas Tech University

Literature Cited

Ackerson, B. K., and L. A. Harveson. 2006. Characteris¬
tics of a ringtail (Bassariscus astutus) population
in Trans-Pecos Texas. Texas Journal of Science
58:169-184.

Anderson, S. 1972. Mammals of Chihuahua: Taxonomy and
distribution. Bulletin of the American Museum of
Natural History 148:151-410.

Baker, R. H. 1956. Mammals of Coahuila, Mexico. Uni¬
versity of Kansas, Museum of Natural History
9:125-335.

Baker, R. H. 1999. R i n gta i 1 1 Bassariscus astutus . Pp. 219—
221 in The Smithsonian book of North American
mammals (D. E. Wilson and S. Ruff, eds.). Smith¬
sonian Press, Washington, xxv + 750 pp.

Blair, W. F. 1950. The Biotic Provinces of Texas. Texas
Journal of Science 2:93-117.

Crandall, L. S. 1974. Management of wild mammals in
captivity. University of Chicago Press, xv + 769 pp.

Durrant, S. D. 1952. Mammals of Utah: Taxonomy and
distribution. University of Kansas, Museum of
Natural History 6:1-549.

Elbroch, M. 2006. Animal skulls: A guide to North Ameri¬
can species. Stackpole Books, Mechanicsburg,
Pennsylvania, xi + 727 pp.

Gehrt, S. D. 2003. Raccoon: Procyon lot or and allies. Pp.
611-634 in Wild mammals of North America:
Biology, management, and conservation (G. A.
Feldhamer, B. C. Thompson, and J. A. Chapman,
eds.). Johns Hopkins University Press, Baltimore,
xiii + 1216 pp.

Gittleman, J. L., and P. Van Valkenburgh. 1997. Sexual
dimorphism in the canines and skulls of carnivores:
Effects of size, phylogeny, and behavioural ecology.
Journal of Zoology, 242: 97-117.

Goodwin, G. G. 1969. Mammals from the state of Oaxaca,
Mexico, in the American Museum of Natural His¬
tory. Bulletin of American Museum of Natural
History 141:1-269.

Hall, E. R. 1981. The mammals of North America. John
Wiley & Sons, New York, 2:601-1181.

Hall, E. R. 1946. Mammals of Nevada. University of Cali¬
fornia Press, Berkeley, xi + 710 pp.

Harrison, R. L. 2012. Ringtail ( Bassariscus astutus ) ecol¬
ogy and behavior in central New Mexico, USA.
Western North American Naturalist 72:495-506.

Helgen, K. M., R. Kay, L. E. Helgen, M. T. N. Tsuchiya-
Jerep, C. M. Pinto, K. Koepfli, E. Eiirik, and J. E.

Maldonado. 2009. Taxonomic boundaries and
geographic distributions revealed by an integrative
systematic overview of the mountain coatis, Na-
suella (Carnivora: Procyonidae). Small Carnivore
Conservation 41:65-74.

Hintze, J. L. 1990. Number Cruncher Statistical Systems,
Version 5.3. Pacific Ease Co., Santa Monica,
California, 442 pp.

Hoffmeister, D. F. 1986. Mammals of Arizona. University
of Arizona Press, Tucson, xix + 602 pp.

Kennedy M. L., and S. L. Lindsay. 1984. Morphological
variation in the raccoon, Procyon lotor, and its
relationship to genic and environmental variation.
Journal of Mammalogy 65:195-205.

Kortlucke, S. M. 1973. Morphological variation in the
kinkajou, Potos flavus (Mammalia: Procyonidae),
in Middle America. Occasional Papers of Museum
of Natural History, University of Kansas 17:1-36.

Nowak, R. M. 1999. Walker’s Mammals of the World. Johns
Hopkins University Press, Baltimore 1:1-838.

Poglayen-Neuwall, I., andD. E. Toweill. 1988. Bassariscus
astutus. Mammalian Species 327:1-8

Ritke, M. E. 1990. Sexual dimorphism in the raccoon
(Procyon lotor). Morphological evidence for in-
trasexual selection. American Midland Naturalist
124:342-351.

Ritke, M. E., and M. L. Kennedy. 1993. Geographic varia¬
tion of sexual dimorphism in the raccoon Procyon
lotor. American Midland Naturalist 129:257-265.

Schmidly, D. J. 1984. The furbearers of Texas. Texas Parks
& Wildlife Dept., 111: xii + 55 pp.

Schmidly, D. J. 2004. The mammals of Texas. University
of Texas Press, Austin, 501 pp.

Stains, H. J. 1984. Chapter 15: Carnivores. Pp. 491-521
in Orders and families of Recent mammals of the
world (S. Anderson and J. K. Jones, Jr., eds.). John
Wiley & Sons, New York, xii + 686 pp.

Stangl, Jr., F. B., and E. M. Jones. 1987. An assessment
of geographic and seasonal biases in systematic
mammal collections from two Texas universities.
Texas Journal of Science 39:129-137.

Toweill, D. E., and D. B. Toweill. 1978. Growth and de¬
velopment of captive ringtails (Bassariscus astutus
flavus). Carnivore 1:46-53.

Verts, B. J., and L. N. Carraway. 1998. Land mammals of
Oregon. University of California Press, Berkeley,
xvi + 668 pp.

Stangl et al.—Cranial Variation in Bassariscus astutus

9

Addresses of authors:

Frederick B. Stangl, Jr.

Nicholas Lamar

Department of Biology

Department of Biology
Midwestern State University
Wichita Falls, Texas 76308

Midwestern State University
Wichita Falls, Texas 76308
frederick. stangl@mwsu. edu

Stephen Kasper

Sarah Henry-Langston

Department of Biology
Midwestern State University
Wichita Falls, Texas 76308

Lake Alan Henry Wildlife Mitigation Area
City of Lubbock
Lubbock, Texas 79457

Appendix

Following is a list of 152 specimens of Bassariscus astutus examined from the collections of Midwestern State
University (MWSU; n = 88) and the Museum of Texas Tech University (TTU; n = 64).

TEXAS (n = 146).—Bandera Co.: 18 mi NW Medina, 1 (TTU 92544). Brewster Co.: Black Gap Wildlife
Area, 1 (TTU 1381); BBNP, Harte Ranch Mt. Lodge, 1 (TTU 62985); 13 miN, 3 mi WMarathon, 1 (TTU29028);
13.4 mi N, 4 mi W Marathon, 1 (TTU 29026); 13.5 mi N, 2.5 mi W Marathon, 1 (TTU 29025); 13.75 mi N, .75
mi W Marathon, 1 (TTU 29024); 17.3 mi N, 0.6 mi E Marathon, 1 (TTU 22980); 17.9 mi N, 0.3 mi E Marathon,
1 (TTU 22981); 18 mi N, 3.0 mi E Marathon, 1 (TTU 22982). Burnet Co.: Burnet, 1 (MWSU 21871); 1.5 mi N,
3 mi WNaruna Community, 1 (TTU 59703). Clay Co.: 20 mi S Henrietta, 1 (MWSU 11040). Comanche Co.: 6
mi S Gorman, 1.5 mi E Hwy 6, 1 (TTU 43365). Crockett Co.: 9 mi S, 5 mi E Iraan, 1 (TTU 58509). Culberson
Co.: Guadalupe Mts., Lower McKittrick Canyon, 1 (TTU 25167); Guadalupe Mts., Upper Dog Canyon, 1 (TTU
32452); Sierra Diablo WMA, 1 (TTU 75781). Edwards Co.: 18 mi N Barksdale Eagles Nest Ranch, 1 (TTU
107712). Garza Co.: 16 mi S, 5 mi E Post, 2 (TTU 56563-4). Hood Co.: 4.2 mi N Granbury, 1 (MWSU 21769).
Howard Co.: Big Spring, 1 (MWSU 6049). Jack Co.: 9 mi N Graford, 1 (MWSU 10946). Jeff Davis Co.: Fort
Davis, 1 (MWSU 10945); 9 km N, 9.5 km E Fort Davis, 2 (TTU 32453, 32455); 9 mi NE Fort Davis, 2 (TTU
14063, 17409); 9.2 mi NE Fort Davis, 1 (TTU 17408). Kendall Co.: 2 mi N, 2 mi W Sisterdale, 1 (TTU 57961).
Kerr Co.: 14 mi W Hunt, 1 (MWSU 9808). Kimble Co.: 5 mi E Junction, 1 (MWSU 7507); 6 mi E Junction, 1
(MWSU 1378); 7 mi E Junction, 17 (MWSU 1767-9, 1777, 8785, 8788-9, 8791-2, 8794, 8797, 8800-1, 8803,
8806-7, 8811); 8 mi E Junction, 14 (MWSU 6473, 6484-6, 6488, 8779-80, 8784, 8786, 8795-6, 8798-9, 8809);
10 mi E Junction, 21 (MWSU 3388, 3704, 4077, 6469, 6474, 6476-83, 7499-506); 12 mi E Junction, 8 (MWSU
8778, 8781-2, 8787, 8790, 8793, 8805, 8808); Kimble Co., no specific locality, 3 (MWSU 1374, 3387, 6475);
1.6 mi S, 3 mi W Junction, 3 (TTU 23724-23726); 5 mi W Junction, 1 (TTU 39494); Texas Tech University Cen¬
ter at Junction, 1 ( TTU 71102); Walter Buck WMA, 1 (TTU 76655). McCulloch Co.: Near Brady, 1 (MWSU
16527). Menard Co.: 20 mi E Menard, 1 (MWSU 6050); 4 mi N, 8 mi W Menard, 1 (TTU 59704); 14 mi W, 2
mi S Menard, 1 (TTU 6661). Nolan Co.: 14 mi SE Sweetwater, 1 (TTU 6664). Palo Pinto Co.: 1 mi S Brad, 1
(MWSU 11512); 4 mi NNE Palo Pinto, 1 (MWSU 750); 10 mi W Graford, 1 (MWSU 8932); 11 mi W Graford,
1 (MWSU 13326); 15 mi W Graford, 1 (MWSU 18723); 17 mi SW Graford, Brazos River, 1 (MWSU 11513);
Possum Kingdom State Fish Hatchery, 1 (MWSU 13386); Santo, 1 (MWSU 747); Possum Kingdom Lake, 2
(TTU 38698, 38699). Pecos Co.: 5 mi S, 2 mi E Girvin, 1 (TTU 49075); 12.2 miN, 19.7 mi E Marathon, 1 (TTU

10

Occasional Papers, Museum of Texas Tech University

Appendix (cont.)

25161). Presidio Co.: 6.5 mi NW Plata, 1 (MWSU 8802); Big Bend Ranch State Natural Area, 2 (TTU 67579,
67580); Clay Miller Ranch, 2 (TTU 9755,11770); 10 mi WSW Valentine, Clay Miller Ranch, 1 (TTU 78532); 11
mi W Valentine, 4 (TTU 92744-92746, 92764). San Saba Co.: 5 mi E Cherokee, 1 (TTU 92786); 7 mi SE Bend,
1 (TTU 92787). Stephens Co.: 4 mi E Breckenridge, 1 (MWSU 8810). Sutton Co.: 4 mi S Sonora, Hwy 1691, 1
(TTU 9756); 35 mi E Sonora, 1 (TTU 35908). Terrell Co.: 20 mi S Sheffield, 1 (TTU 92796); 23 mi S Sheffield,
1 (TTU 92801). Tom Green Co.: 2 mi S San Angelo, 1 (TTU 92810). Travis Co.: 5 mi W Austin, Gaines Ranch,
1 (TTU 92820); 6 mi SW Leander, 1 (TTU 1671B); Travis Co., no specific locality, 1 (TTU 1671A). Val Verde
Co.: 3 miWComstock, 1 (MWSU7839); 12 miNComstock, 1 (MWSU 15459); 44 miNDelRio, 1 (TTU6663).
Wichita Co.: 1 mi W Wichita Falls, 1 (MWSU 751). Young Co.: Young Co., no specific locality, 1 (MWSU 5986).

MEXICO (n = 6).—Coahuila: 15 mi E Monclova, Gloria Mts., 2 (TTU 92458, 92459); 27 miNE Muzquiz,
Mariposa Ranch, 3 (TTU 92462-92464). Nuevo Leon: 1 kmN San Josecita, Zaragoza, 1 (TTU 57122).

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