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Carpenter 
or lacertid, the male moves across the trunk 
or flank of the female and obtains a biting 
hold on this region while arching its body 
across the female and bringing its cloacal 
region into juxtaposition with hers before 
inserting a hemipenis (Fig. 16). A hindleg 
may be thrown across the base of the tail of 
the female as a holding mechanism. This 
hold is usually maintained throughout copu- 
lation in Cnemidophorus, but in the lacertid 
Acanthadactylus bosquianm once insertion is 
obtained the male may release his bite hold 
on the female. 
Another type of behavior that I believe to 
be ritualistic is the response of iguanid 
females which appear to be nonreceptive to 
courting males. This response has been seen 
in a number of iguanid genera, and I have 
called it female rejection behavior (Carpen- 
ter, 1962). The features of female rejection 
behavior are: caudal orientation toward the 
aproaching male, arched back, raised tail 
with tip pointing dorsally, and often a lower- 
ing of the head and inflated trunk, (Fig. 17). 
In this posture some species may sidlehop in 
a half circle about the male {Sceloporus 
undulatus, Tropidurus albemarlensis) . In 
Holbrookia maculata, such rejection behavior 
is further enhanced by exposure of the thigh 
which in this species has a contrasting yel- 
lowish color (Clarke, 1965). 
A most unusual type of ritualistic encoun- 
ter between males occurs in Vararms gilleni, 
a small monitor (Varanidae) from Australia 
Figure 16. The characteristic copulatory position 
of teiid and lacertid lizards involves the male’s 
biting and holding the flank or groin region of the 
female as he arches across her trunk — as here 
seen in a pair of the Six-lined Racerunner 
(CnemidophortLS sexlineatua — Teiidae) from Okla- 
homa. 
(Murphy and Mitchell, 1973; Carpenter, et 
al., 1976). Two males, when placed together, 
proceed to move along side of one another 
and with their legs flaying attempt to em- 
brace each other. Once they begin to embrace 
with belly to belly contact, they immediately 
flex their trunks laterally away from the 
substratum, and with simultaneous lateral 
flexion of the neck and tail, still embracing, 
rise from the substratum to form a tetrapod 
arch, the two snouts and the posterior tail 
regions forming the posts of the tetrapod 
(Fig. 18). Such actions are repeated many 
times. 
What I have presented here is a variety of 
behaviors in lizards related to social organi- 
zation which exhibit various degrees of ritu- 
alization. There are, of course, other aspects 
of social behavior and ritualistic actions 
which I have not discussed. We are just 
beginning to understand many aspects of 
lizard behavior. 
To gain some perspective on “where we 
have gone and where we might go” I have 
included a model (Fig. 19) that I made up 
a number of years ago that is centered 
around the ritualistic display-action-patterns 
exhibited by iguanid and agamid lizards. 
Without getting involved in all of the pos- 
sible relationships suggested by this model, 
I would like to point to some critical areas 
requiring investigation. Looking at the right 
upper part of this diagram we may first 
Figure 17. A female Lesser Earless Lizard 
{Holbrookia maculata — Iguanidae) may reject the 
courtship actions of the male by rising high on all 
four legs, arching her back, presenting caudally 
toward the male. This action presents a yellow 
mark on the posterior thigh region to the male. 
