Behavior and Neurology o1 Lizards 
N. Greenberg and P. D. MacLean, eds. 
NIMH. 1978. 
Ritualistic Social Behaviors in Lizards 
Charles C. Carpenter 
Department of Zoology 
University of Oklahoma 
SUMMARY. The social behavior of lizards falls into three main categories: (1) agonistic, (2) 
courtship and mating, and (3) parental care. Agonistic behaviors relate to aggression and sub- 
mission, territoriality, hierarchies, display, threat, fighting, and vocalizations. Courtship and 
mating behaviors include action sequences (both sexes) leading to copulation. 
A significant feature of lizard social interactions is their ritualistic nature. Ritualistic be- 
haviors appear to be more highly developed in those lizard families which exhibit social systems 
based on territoriality (Iguanidae, Agamidae, Chameleontidae, Gekkonidae). In less “territorial” 
lizards (Lacertidae, Teiidae, Scincidae, Varanidae, and Xantusiidae) there appears to be cor- 
respondingly less ritual. Little is known of the social interactions in other lizard families. 
DISPLAY BEHAVIOR 
Display behaviors may involve a variety of 
different types of movements (often time- 
specific), postural changes, color changes, 
and pattern accentuations, or the morphologi- 
cal elaboration of special structures. The 
interplay between male and female during 
courtship may relate to specific signals based 
on movements, postures, and other actions 
that are often different from agonistic be- 
haviors only in their social context. 
Representative examples of the above, 
primarily from iguanid and agamid lizards, 
exhibit the diversity of such actions and in- 
dicate the species-characteristic nature (ritu- 
alization) of these behaviors. Such behaviors 
offer bases for investigations of neural sub- 
strates of ritualized activity. 
For the present discussion, I shall define 
social behavior as those behaviors performed 
by an individual lizard which may be elicited 
by another individual or may have some 
effect on another individual (s), usually of the 
same species, but sometimes of another 
species. 
The social behaviors of lizards fall into 
three main categories — (1) agonistic, (2) 
courtship and mating, and (3) parental care. 
The approach to the study of social behavior 
of lizards, can be either through field or 
laboratory studies and not only involves ob- 
servations on the natural attributes of social 
structures, but also the experimental analysis 
of social interactions, the physiological bases 
of social status and actions, and the ritualis- 
tic characteristics of the behavior patterns 
performed. 
Parental care is little known in lizards and 
most of my remarks will deal with agonistic 
and courtship behavior. 
Agonistic behaviors include action involved 
in aggression, submission, territoriality, and 
hierarchy formation, and entail displays, 
threats, fighting, and certain vocalizations. 
Courtship and mating behaviors include those 
action sequences by one or both sexes which 
prepare for or lead to copulation. 
The lizard social systems for which ade- 
quate information is available range from 
those characterized by territoriality and/or 
hierarchies to those with incipient territorial 
development, to those showing intolerance 
between individuals that do not defend a 
particular area. 
Those lizard families which have been 
shown by field and laboratory studies to es- 
tablish and protect territories (Table 1) are 
the Iguanidae (Berry, 1974; Carpenter, 1962, 
1966, 1967a, 19676; Evans, 1951; Fitch, 
1940; Hunsaker, 1962; Jenssen, 1970; Mil-' 
stead, 1970; Noble and Bradley, 1933; Rand, 
