Herpetologists' League

An Experimental Analysis of the Role of the Tail in Attaining High Running Speed in
Cnemidophorus sexlineatus (Reptilia: Squamata: Lacertilia)
Author(s): Royce E. Ballinger, Joseph W. Nietfeldt and James J. Krupa
Source: Herpetologica, Vol. 35, No. 2 (Jun., 1979), pp. 114-116
Published by: Allen Press on behalf of the Herpetologists' League
Stable URL: http://www.jstor.org/stable/3891774
Accessed: 04-03-2016 09:25 UTC

REFERENCES
Linked references are available on JSTOR for this article:
http://www.jstor.org/stable/3891774?seq=1&cid=pdf-reference#references_tab_contents

You may need to log in to JSTOR to access the linked references.

Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available at http://www.jstor.org/page/
info/about/policies/terms.jsp

JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content
in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship.
For more information about JSTOR, please contact support@jstor.org.

Herpetologists' League and Allen Press are collaborating with JSTOR to digitize, preserve and extend access to Herpetologica.

http://www.jstor.org

This content downloaded from 131.204.73.184 on Fri, 04 Mar 2016 09:25:16 UTC
All use subject to JSTOR Terms and Conditions
114 HERPETOLOGICA [Vol. 35, No. 2

Sceloporus undulatus: a study of the intraspe-

Alabama in Birmingham, Birmingham, AL

cific comparative demography of a lizard. Ecol-

35294, USA; Department of Biological Sci-

ogy 53:570-584.

ences, Illinois State University, Normal, IL

61761, USA; Biology Department, Wash-

Accepted: 13 June 1978

ington University, St. Louis, MO 63130,

Department of Biology, University of USA

Herpetologica, 35(2), 1979, 114-116

( 1979 by The Herpetologists' League

AN EXPERIMENTAL ANALYSIS OF THE ROLE

OF THE TAIL IN ATTAINING HIGH RUNNING

SPEED IN CNEMIDOPHORUS SEXLINEATUS

(REPTILIA: SQUAMATA: LACERTILIA)

RoycE E. BALLINGER, JOSEPH W. NIETFELDT, AND JAMES J. KRUPA

ABSTRACr: Time trials indicated that running speed of individual lizards was reduced by

an average of 36% by removing their tails.

Key words: Lacertilia; Locomotion; Tail

VITT et al. (1977) provided a general monly in open, grassy habitats. Its natural

theoretical framework within which the history was detailed by Fitch (1958), who

phenomenon of tail autotomy may be noted that C. sexlineatus tends to retain its

evaluated. Their broad categories of adap- tail as compared to skinks, anquids, and

tations include actively functional tails

most iguanids. These racerunners depend

used for climbing, swimming, balance,

on speed to escape predators (Fitch, 1958).

and defense, and passively functional The following experiment was conducted

(autotomous) tails used in the evasion of

to test the hypothesis that the tail is impor-

predators. The former category represents tant to attaining high running speeds.

a tail-retention adaptation whereas the

latter utilizes autotomy and thus implies a

MATERIALS AND METHODS

tail-loss adaptation. With few exceptions

The study was conducted at Cedar Point

(notably Ballinger, 1973; Clark, 1971; Cong-

Biological Station, Keystone, Nebraska, in

don et al., 1974; Vitt et al., 1977), tail autot-

the summer of 1977. Animals were collected

omy has received little experimental or

by noose or hand, held in a terrarium, and

analytical study, with adaptations being

provided with water and food ad libitum

inferred from tail-break frequencies re-

until used in the experiment. Lizards were

corded in life history studies or from obser-

forced to run in a long, wooden chamber

vations of behavior. In this study, we ex-

(6 m long, 10 cm wide, 20 cm high). Speed

perimentally analyze the function of the tail

of running was determined over a distance

in the six-lined racerunner, Cnemidophorus

of 3 m after it became apparent that the

sexlineatus.

The lizard Cnemidophorus sexlineatus is lizards could not be forced to maintain

widely distributed in the southeastern and maximum speeds consistently for the entire

central United States where it occurs com- 6-m length of the chamber. Running times

This content downloaded from 131.204.73.184 on Fri, 04 Mar 2016 09:25:16 UTC
All use subject to JSTOR Terms and Conditions
June 1979] HERPETOLOGICA 115

TABLE 1.-Mean values for length, weight, and time required to run 3 m for 10 Cnemidophorus sex-

lineatus.

Difference

Snout- Mass between

vent Length Body of tail Running time (s) Running time (s) shortest

length of tail mass removed with tail without tail running

Individual Sex (mm) (mm) (g) (g) (n; range) (n; range) times (s)

1 9 50 107 3.22 .23 1.63 (7; 1.5-1.8) 2.74 (5; 2.4-3.2) .9

2 9 54 84* 4.08 .38 1.55 (5; 1.2-1.95) 1.85 (2; 1.7-2.0) .5

3 9 62 151 6.60 .63 1.39 (4; 1.25-1.5) 2.44 (5; 1.9-3.0) .65

4 9 67 146 8.13 .55 1.26 (6; 1.1-1.35) 1.85 (4; 1.4-2.5) .3

5 9 73 151 9.54 .72 1.53 (9; 1.4-1.7) 2.13 (3; 1.7-2.5) .3

6 8 64 141 7.93 .69 1.40 (2; 1.4-1.4) 2.90 (3; 2.5-3.3) 1.1

7 9 68 153 14.33 1.03 2.28 (4; 2.0-2.4) 2.40 (1; 2.4) .4

8 e 53 112 4.49 .37 1.28 (2; 1.25-1.3) 2.03 (7; 1.7-2.5) .45

9 9 65 132 8.81 .98 1.95 (2; 1.7-2.2) 2.92 (9; 2.3-3.2) 1.4

10 9 54 128 4.78 .50 1.62 (4; 1.4-1.9) 2.72 (4; 2.0-3.4) .6

* See text for explanation.

were recorded with a stopwatch to the removed was 2.46 s compared to 1.57 s in

nearest 0.2 s. An enclosure at the end of intact animals. The loss of the tail resulted

the chamber served to prevent the lizards in a 36% decrease in speed. No relationship

from escaping, and a small mound (0.2 m

other than absence of tail explained the

high) in the center of the enclosure was

difference in speed, including sex, tempera-

constructed so that the lizards would not

ture, body weight, or body size.

slow down as they approached the enclo-

The manner in which the tail was used

sure. Thus, the lizards appeared to view

to increase speed was probably as a coun-

the end of the chamber as an escape route,

ter-balance mechanism to shift the center

but were chased to ensure maximum speed.

of gravity of an animal more directly on

Ten lizards were used in the study, and 2-9

the force exerted by the hind legs. Similar

trials were conducted on each lizard with

explanations for locomotion in bipedal liz-

its tail intact. Time (0.5-1 h) was per-

ards have been used by Snyder (1949,

mitted for rest between each trial. The tail

1952). The tail was raised off the ground

was then removed by hand, and additional

and held rigid in Cnemidophorus sexlinea-

trials were performed the following day ( s).

tus during running. Lizards attempting to

Unequal numbers of trials resulted because

run without the leverage afforded by the

some lizards refused to attempt to escape

tail were clumsy and often stumbled during

by running, perhaps because of their famil-

attempts to run. Furthermore, lizards with-

iarity with the chamber. Experiments were out tails often tumbled end over end near

the end of a run, perhaps because the liz-

conducted over the course of three weeks,

ards used the tail while stopping by swing-

but within a few days for any particular

ing it downward.

lizard. Body temperatures were taken

In Cnemidophorus sexlineatus, the elon-

before each trial, and only trials with lizards

gate tail, which comprises almost 70% of

having body temperatures in the preferred

the total body length (Table 1), probably

temperature range (35-410C; Carpenter,

serves as a counter-balance allowing high

1961; Fitch, 1956) were used.

running speeds. These lizards utilize speed

as the principal means of predator escape;

RESULTS AND DISCUSSION

therefore, caudal autotomy occurs less fre-

Table 1 summarizes the results of the

quently and less easily than in species utiliz-

experiment. The average time required for

ing tail-breakage rather than speed for

lizards to run 3 m after their tails were

escape from predators. Our experiments

This content downloaded from 131.204.73.184 on Fri, 04 Mar 2016 09:25:16 UTC
All use subject to JSTOR Terms and Conditions
116 HERPETOLOGICA [Vol. 35, No. 2

1974. Geckos: adaptive significance and ener-

suggest that natural selection has favored

getics of tail autotomy. Science 184:1379-1380.

tail-retention in Cnemidophorus sexlineatus.

FITCH, H. S. 1956. Temperature responses in

free-living amphibians and reptiles of northeast-

Acknowledgments.-We thank Dr. Brent Nickol,

ern Kansas. Univ. Kansas Pub. Mus. Nat. Hist.

Director, Cedar Point Biological Station, for assis-

8:417-476.
tance and use of facilities under his care, and

. 1958. Natural history of the six-lined

David Dempsey for assistance in collecting speci-

racerunner (Cnemidophorus sexlineatus). Univ.

mens.

Kansas Pub. Mus. Nat. Hist. 11:11-62.

SNYDER, R. C. 1949. Bipedal locomotion of the

LITERATURE CITED
lizard Basiliscus basiliscus. Copeia 1949:129-

137.

BALLINGER, R. E. 1973. Experimental evidence

1952. Quadrupedal and bipedal loco-

of the tail as a balancing organ in the lizard,

motion of lizards. Copeia 1952:64-70.

Anolis carolinensis. Herpetologica 29:65-66.

VITT, L. J., J. D. CONGDON, AND N. A. DICKSON.

CARPENTER, C. C. 1961. Temperature relation-

1977. Adaptive strategies and energetics of tail

ships of two Oklahoma lizards. Proc. Oklahoma

autotomy in lizards. Ecology 58:326-337.

Acad. Sci. 41:72-77.

CLARK, D. R. 1971. The strategy of tail autot- Accepted: 21 June 1978

omy in the ground skink, Lygosoma laterale. J.

School of Life Sciences, University of

Exp. Zool. 176:295-302.

Nebraska, Lincoln, NE 68588, USA

CONGDON, J. D., L. J. VIrr, AND W. W. KING.

Herpetologica, 35(2), 1979, 116-124

? 1979 by The Herpetologists' League

NOTES ON SYSTEMATICS OF HORNED LIZARDS

ALLIED TO PHRYNOSOMA ORBICULARE

(LACERTILIA: IGUANIDAE)

RICHARD R. MONTANUCCI

ABSTRACT: Two nominal subspecies of Phrynosoma orbiculare, P. o. bradti and P. o.

durangoensis Horowitz 1955, are considered synonyms. The name bradti is chosen to represent

populations from the Sierra Madre Occidental of Mexico. Phrynosoma boucardi Dumeril and

Bocourt (1870) is relegated to a subspecies of P. orbiculare based on intergradation with P. o.

cortezi in eastern Hidalgo. Alleged osteological differences between boucardi and orbiculare

are invalidated by a re-examination of available material. Phrynosoma o. boucardi is believed

not to be phylogenetically close to P. taurus and P. braconnieri. The name P. o. dugesi, once

problematical, is now applied to specimens from Jalisco which compare favorably with the

syntypes.

Key words: Lacertilia; Phrynosoma; Systematics

THE short-horned lizards Phrynosoma agreement persists concerning the patterns

douglassi and Phrynosoma orbiculare have of geographic variation and the subspecific

posed a taxonomic enigma for many years. limits within P. douglassi (Conant, 1975;

At one time the specific status of the two Reeve, 1952; Smith, 1946; Stebbins, 1966)

taxa was questioned (see Smith, 1946, and as well as within P. orbiculare (Davis, 1953;

references therein), although Smith and Horowitz, 1955; McDiarmid, 1963; T. Uz-

Taylor (1950) later felt that the two forms zell, personal communication). The work

should be retained as distinct species. Dis-

of Horowitz (1955) was an important con-

This content downloaded from 131.204.73.184 on Fri, 04 Mar 2016 09:25:16 UTC
All use subject to JSTOR Terms and Conditions