Ritualistic Behavior Lizards 
255 
1967; Stamps, 1973; Tinkle, 1967); Agami- 
dae (Blanc and Carpenter, 1969; Brattstrom, 
1971; Carpenter, Badham and Kimble, 1970; 
Harris, 1964; John, 1970; Pfeifer, 1962); 
Chameleontidae (Brain, 1961 ; Bustard, 1966; 
Parcher, 1974) ; and the Gekkonidae (Greer, 
1967; Kastle, 1964). Those families with 
some species showing some degree of terri- 
torial behavior are the Teiidae (Carpenter, 
1962; Hirth, 1963; Mueller, 1971); Lacerti- 
dae (Boag, 1973; Steward, 1965; Weber, 
1957); Scincidae (Fitch, 1954; Noble and 
Bradley, 1933) ; Varanidae (Pfeifer, 1959) ; 
and Xantusiidae (Lowe, 1948). 
The families for which little information 
is available on agonistic interactions or ter- 
ritoriality are the Anguidae, Cordvlidae, An- 
niellidae, Dibamidae, Gerrhosauridae, Helo- 
dermatidae, Lanthanotidae, Pygopodidae, 
and Xenosauridae. The dearth of information 
appears to be due to the secretive nature of 
the species in these families or to the lack of 
adequate studies. 
The territorial male performs some type of 
display to declare himself present to other 
males, to challenge and intimidate neighbors, 
or invading males. It demonstrates agonistic 
contact behavior such as biting, head butting, 
tail slapping, and chasing to assert its de- 
termination to defend an area, its territory. 
In some species females may exhibit terri- 
torial defense behavior, e.g., TJta stansburi- 
ana, (Tinkle, 1967). Uma exsul defend nest- 
ing burrows (Carpenter, 1967&), as do 
AmhlyrhynchuLS cristatus (Carpenter, 1966). 
The intensity of territorial actions may vary 
with the onset and waning of the breeding 
season. 
Among teiids, lacertids, the skinks, as well 
as the less known varanids and xantusiids, 
very few examples of defended areas have 
been recorded, although there is frequent 
overlapping of activity ranges. 
The relationships of females, juveniles, and 
hatchlings to the territorial and nonterri- 
torial systems are less understood. Mono- 
gamous male-female associations within a 
territory (Tinkle, 1967), as well as harems 
of females associated with one male, are 
recorded for iguanids (Carpenter, 1967). The 
relationships of mating systems to social 
structure, as dealt with by Noble and Bradley 
(1933), Tinkle (1969), Milstead (1970), and 
Hunsaker and Burrage (1969), need much 
more study before generalizations can be 
stated. 
RITUALISTIC ASPECTS OF BEHAVIOR 
The remainder of my remarks will be 
directed to the ritualistic nature of lizard 
behaviors. Each of these behaviors is made 
up of acts and/or act systems which involve 
specific motor patterns in their performance. 
An act is the most basic movement produced 
by a particular set of muscular actions (e.g., 
raising the head and lowering the head are 
separate acts). An act system is a series of 
acts performed in a particular sequence, 
either simultaneously and/or in a series (e.g., 
head bobbing, tail waving, striking, or bit- 
ing). Some acts may contribute to more than 
one act systems (See Russel, Mead, and 
Hayes, 1954, and Dilger, 1960, for further 
discussion of acts and act systems). 
In discussing these behaviors, I arbitrarily 
separate categories for the sake of examina- 
tion, but with the knowledge that some of 
these behaviors appear in a variety of con- 
texts and sequences. My remarks pertain 
primarily to iguanid and agamid lizards be- 
cause I am most familiar with them and be- 
cause their behavior is best known. 
Agonistic Behavior 
Threat 
Perhaps the simplest type of agonistic be- 
havior, as a type of aggression, is threat be- 
havior. Threat generally involves some type 
of postural change on the part of the per- 
former which in some way increases its ap- 
parent size and may present a new pattern 
or color to an adversary. Some examples of 
threat which appear quite spectacular are as 
follows : 
Ch^amydosaurus kingi (frilled lizard of 
Australia) is an agamid which, in aggressive 
