Springer-VerlagTokyohttp://www.springer.de101640289-07711439-5444Journal of EthologyJ EtholLifeSciences19210.

1007/s10164-006-0192-8

J Ethol (2006) 24:275–283 © Japan Ethological Society and Springer-Verlag Tokyo 2006
DOI 10.1007/s10164-006-0192-8

ARTICLE

Rajkumar S. Radder • Srinivas K. Saidapur •

Richard Shine • Bhagyashri A. Shanbhag

The language of lizards: interpreting the function of visual displays of the

Indian rock lizard, Psammophilus dorsalis (Agamidae)

Received: May 2, 2005 / Accepted: November 18, 2005 / Published online: April 5, 2006
Japan Ethological Society and Springer-Verlag
2006

Abstract The first step in understanding any communica- Key words Assertion display · Directed display · Lizards ·
tion system is to document signal diversity relative to the Reptiles · Social display
context of signalling (e.g. sex of the signaller and audience).
Observation of 30 free-ranging rock lizards (Psammophilus
dorsalis) on rock outcrops in southern India over a period
of 18 months revealed that these lizards produce a complex Introduction
array of ritualized signals involving push-ups (head-
bobbing), dorsal flattening, extension of the legs or gular Communication is an important component of all social
region, and tail-raising. Push-ups were performed by both interactions. Animals communicate with heterospecific
sexes, usually after moving from one location to another. organisms (competitors, predators, prey) and with conspe-
Push-ups were rarely accompanied by other postural mod- cific organisms (rivals, potential mates, social-group mem-
ifications, and seem to function as non-directed signals. bers) in complex ways, using a wide range of sensory
Dorsal flattening was elicited by birds flying overhead, and modalities. Some of the best-understood systems involve
seems to make the lizard less conspicuous to predators. stereotyped visual displays that sometimes convey very
There was, nonetheless, a strong sex difference in the fre- specific information, for example the dance of the honey-
quency of this behaviour, because the habitats used by bee (Dyer 2002) and dominance and appeasement postures
males (open rocks) exposed them to more birds. Males in mammals (Tomasello and Zuberbuehler 2002; Liebal
displayed to females by extending their gular folds and et al. 2004). Understanding the functional role of specific
arching their backs; other animals (e.g. squirrels, monkeys) displays poses formidable logistical problems, however,
also elicited the latter posture from both sexes. Leg exten- because of the complexity of signalling. For example, a sin-
sion was observed for both males and females, but in differ- gle signal might serve multiple functions, and/or a single
ent contexts – males in response to conspecifics, females in function might be accomplished by means of multiple
response to other animals. Females raised their tails in signals (Møller and Pomiankowski 1993; Johnstone 1995,
response to encountering a male. Thus, these lizards have a 1996, 1997; Brooks and Couldridge 1999). Also, the way in
complex repertoire of postures for predator evasion, for which a signal is expressed may vary with factors such as
interaction with other species and with conspecifics, and for the age, size, signalling environment, dominance, or physi-
communicating sex-specific social information about gender ological status of the sender or the receiver (Clutton-Brock
(tail-raise) or dominance status (gular extension, leg et al. 1982; Tokarz 1985; Snowdown 1990; Davies 1992;
extension). Sinervo and Lively 1996; Prum 1998; Leal 1999; Castellano
et al. 2000; Podos 2001; Baird et al. 2003; Whiting et al.
2003).
To facilitate the search for links between signal form and
R.S. Radder · S.K. Saidapur · B.A. Shanbhag function, many researchers have taken study systems into
Department of Zoology, Karnatak University, 580003 Dharwad, India the laboratory where stimuli can be controlled (Holberton
Fax +91-836-2448047
e-mail: bhagyashrishanbhag@gmail.com
et al. 1989; Evans 1991; Chappell et al. 1995; Evans and
Norris 1996). Although this method is well-suited to teasing
R.S. Radder · R. Shine
apart correlation from causation, an essential first step –
The School of Biological Sciences A08, University of Sydney, 2006
NSW, Australia especially for study organisms that have not attracted pre-
vious research – is to document the array of signals exhib-
R.S. Radder
The School of Biological Sciences A08, University of Sydney, 2006 ited under natural conditions. If such a study also examines
NSW, Australia the context of signal use (e.g. what ages/sexes of animals
276

perform these acts, and in response to which stimuli?), we
can frame preliminary hypotheses about the biological func-
tion of each signal type.
Diurnal territorial lizards are good animals for such a
study. Many taxa, particularly from the agamid and iguanid
lineages, select relatively open sites, where field observa-
tions are feasible, and perform stereotyped displays (for
example head-bobs, body-flattening, limb-waving, push-
ups) at high frequencies, and often in response to particular
(especially social) stimuli (Brattstrom 1971; Carpenter and
Ferguson 1977; Murphy et al. 1978; Gibbons 1979; Hasson
et al. 1989; Cooper and Greenberg 1992; Olsson 1992,1994;
Mori and Hikida 1994; Leal and Rodriguez-Robbes 1997;
Macedonia and Clark 2001; Watt and Joss 2003). Possible
functions of such displays include species recognition, mate
choice, defence of territories and attraction of mates
(Carpenter 1965; Stamps 1977; Martins 1991, 1993;
DeCourcy and Jenssen 1994; Leal and Fleishman 2002; Watt
and Joss 2003). Detailed studies, mostly on North American
species, have clarified the hormonal basis of morphological
features associated with these displays (notably, coloured
patches on males: e.g. Sinervo et al. 2000; Hews and Quinn
2003), and the ways in which variations of signal intensity
affect the responses of conspecifics (Leal 1999; Ord et al.
2001; Watt and Joss 2003). Phylogenetic analyses of lizards
have revealed extreme evolutionary lability in sex-specific
display colours (Wiens 1999; Quinn and Hews 2000), sug-
gesting that parallel studies on an array of taxa have the
potential to provide robust (phylogenetically independent)
tests of hypotheses on the functional significance of
independently-evolved, but often highly convergent, signal-
ling systems.
Our study provides such a set of data. Psammophilus
dorsalis is an Indian agamid (dragon) lizard restricted to
rocky habitats. The species exhibits strong sexual dimor-
phism and dichromatism (males are larger than females,
and develop much brighter colours during the breeding sea-
son). Although the communication systems of P. dorsalis
have never been studied previously, our preliminary obser-
vations revealed that these lizards have a complex reper-
toire of distinctive, highly ritualized display postures. This
study was undertaken to:
males are larger than females (average snout–vent lengths
(SVL) 102 and 72 mm respectively, mass 40 and 25 g).
Females and juvenile males are mottled brown dorsally and
pale ventrally, and thus are well-camouflaged in their rocky
habitat, whereas breeding adult males develop more spec-
tacular coloration – brilliant yellow or orange stripe mid-
dorsally and generally jet-black laterally and ventrally
(Fig. 1). Adult males are territorial, and engage in physical
battles with each other during some parts of the year (per-
sonal observation). Three sites, near the village of Hampi
(15°18′ N 75°2′ E) in Karnataka state, India, were selected
for the study. This area is hot all year, with the ambient
temperature between 0700 and 1900 h ranging from 20 to
47°C over the annual cycle. Each study site was ~0.5 ha in
extent and ~1 km from the other sites. All were hilly areas,
with the substrate comprising bare rocks and historical
stone ruins. Psammophilus dorsalis are common at all three
sites (>90 lizards per ha, Radder et al. 2005). We captured
thirty adult lizards (14 males, SVL > 90 mm and 16 females,
SVL > 65 mm) in late November 2001, marked them by toe-
clipping and then released them at their respective capture
sites. The toe clips were identifiable through binoculars
from a distance of 5–7 m; lizards had no overt reaction to
our presence at this distance. The lizards were observed at
regular intervals from December 2001 to July 2003 (total of
10 observation periods, each 3–5 days in duration).

1 document the array of visual signals made by these liz-

ards; and
2 quantify the relative incidence of different types of dis-
play as a function of the identity of the displaying lizard,
and the context (e.g. audience) of the display.

Materials and methods

Species and study site

Psammophilus dorsalis is a medium-sized diurnal terrestrial

lizard usually found on rocky hills in southern India, breed-
ing from April to August (Sarkar and Shivanandappa 1989;
Shanbhag 2002). The dorsal body colouration of the lizards Fig. 1. Indian rock lizards, Psammophilus dorsalis: (a) an adult male
varies between sexes and seasons (unpublished data). Adult in a push-up display; (b) an adult female in a tail-raise display

Each focal (marked) lizard was observed for 30 min con-
tinuously on two occasions during each observation period:
once in the morning (0800–1230 h) and once in the after-
noon (1600–1900 h), the main activity periods for these liz-
ards (Radder et al. 2005). The focal time was divided into
six consecutive blocks of 5 min each (Grant and Dunham
1988). The order of observation was randomized for a given
individual and sex during each visit to the study site. The
following behaviour was recorded whenever a focal lizard
performed a display:
1 form of the display, including body posture and the num-
ber of head bobs performed; and
2 social context – presence of a conspecific male or female
lizard or other animal (monkey, squirrel, or bird) within
a 10 m radius of the displaying individual.
We cannot be certain we ascertained the stimulus elicit-
ing the display on every occasion; for example, the stimulus
animal may have been hidden from us, or the presence of
another animal may have been irrelevant to the display; in
a few instances we may have mistakenly classified stimulus
animals as females when they were, in fact, small adult
(female-coloured) males (see above).
Tests for reliability and data analyses
We assessed the reliability of our data in two ways. First,
two observers simultaneously recorded data on the same
focal individuals to ensure inter-observer reliability. There
was no disagreement between two observers on measures
of body posture and only minor (<5%) discrepancies in the
number and type of head bobs. Second, we videotaped liz-
ards (80 min) to test the reliability of the data recorded by
an observer using binoculars compared with that seen with
the naked eye. Comparison of data from video clips and
direct recording revealed only minor discrepancies (<2%).
For statistical analysis we combined records from the two
30 min observation periods on each focal animal during a
given visit to yield frequencies per hour per survey period.
Data were kept separate for each individual, to avoid sta-
tistical problems from data pooling (Leger and Didrichson
1994). Thus, our statistical tests are based on single value
per individual as the unit of replication, and mean values
cited in the text are based on overall means with each ani-
mal contributing a single value only. Because many of the
display rates were highly non-normal in distribution (often
with many zero values), we used non-parametric tests to
evaluate statistical significance. An alpha level of 0.05 was
used to determine statistical significance.
To compare males and females in terms of the total num-
bers of displays that they performed, we used Mann–
Whitney U test. To compare the sexes with regard to the
proportions of each type of display we calculated numbers
out of the total sample of directed displays (i.e. excluding
simple push-ups – see below) by each lizard and, again,
analysed these data with Mann–Whitney U test. To examine
the responses of males and females to different kinds of
277
stimulus (conspecifics, heterospecifics, etc.) we calculated
the number of each display type performed by an individual
lizard as a proportion of all displays directed to that stimu-
lus type. This procedure was designed to correct for differ-
ent exposure of individual lizards to different stimuli. We
looked at the effect of stimulus type on the proportion of
displays of each type separately within each sex, using the
Kruskal–Wallis test. Data were analysed using the software
Statview 5 on an Apple Macintosh G5 Computer.
Results
Types of display
Simple push-ups or head bobs
Usually performed with forelimbs straightened such that
the anterior body of the lizard was slightly raised from the
substrate, this display consisted of rapid (1–2 per second)
vertical displacements of the head and trunk while the feet
remained immobile on the substrate (Fig. 1a). Two distinct
types of head bob were performed by these lizards – single
and double. A single head bob consisted of a single cycle of
raising the head ~1 cm above the initial position and then
lowering it again. Double head-bobs were identical except
that the head was raised and lowered twice in quick succes-
sion. Both single and double head-bobs usually took less
than 2 s to complete.
Complex push-ups with display body postures
These displays commenced as described above, but with the
addition of one of the following body postures immediately
after the simple push-up (“modifiers”: Jenssen 1977):
Tail raising The distal half of the tail was lifted vertically
and held pointing upwards, and also occasionally moved
from side to side (Fig. 1b). Lizards often remained in this
tail-raised position for 1–2 min.
Gular extension A small patch of brightly-coloured skin at
the throat was extruded forwards and downwards, then
quickly withdrawn (within 1–2 s) two or three times in quick
succession.
Leg extension The lizards simultaneously extended and
straightened either the forelimbs, or both forelimbs and
hind limbs. When only forelimbs were used, these were bent
and straightened to produce an up-and-down motion of the
forebody. When both fore and hind limbs were used in the
display, the simultaneous extension and retraction of all
four limbs produced up-and-down motion of the entire
body.
Arching of back The spine was bent upward to form an
arch, with the highest point of the curve at the middle of
the torso; lizards in this posture typically remained motion-
less for 1–2 min. This display caused a striking change in the
overall appearance of the displayer.
278

60 6
female

50 male
5
% of displays per lizard

40

# of tail-raises
4
30

3
20

10 2

0
tail-raise gular-extend leg-extend arch-back dorsal flatten 1

Fig. 2. Sex differences between the relative frequency of different

types of “directed” displays by rock lizards. These proportions are
mean values (and standard errors) based on numbers of each display
type as a proportion of the total number of displays recorded for each
individual lizard. These data are based on a total of 87 displays by 16
female lizards, and 208 displays by 14 males
Dorsal flattening The lizard crouched down so that its ven-
tral surface was in direct contact with the substrate, and
flattened its body dorsoventrally such that the entire ventral
surface was in contact with the substrate.
Sex differences in frequency and form of displays
Males were significantly larger in SVL than females (means
98.8 ± 1.97 and 75.3 ± 1.75 mm, F1,28 = 79.79; P < 0.0001) and
performed more displays (means 14.86 ± 1.27 and 5.44 ±
0.68, Mann–Whitney U = 6.50; P < 0.0001; n = 14 males and
16 females). Importantly, the two sexes also used different
habitats, and thus performed their displays in different con-
texts. Adult males selected very high rocky perches (mean
5.07 ± 0.24 m above ground level, n = 14) whereas females
were seen mostly close to the ground (mean = 1.25 ± 0.14 m,
n = 16; comparing the sexes, Mann–Whitney U = 0.00;
n = 28; P < 0.0001).
Push-ups
Both male and female P. dorsalis performed push-up dis-
plays, and these were by far the most common type of
display seen in our lizards (means of 94.4% of displays by
females (n = 1480/1568), 85.0% by males (n = 1168/1376):
comparing the sexes, for total numbers, Chi-square test,
df = 1; χ2 = 22.50; P < 0.0001). Within these totals, the sexes
also differed in the relative numbers of single-bob and
double-bob displays. Females performed mostly single-
bobs (55.1% of total bob displays) whereas males usually
double-bobbed (single-bobs were 44.3% of total bob
displays; test on proportions, Chi-square test, df = 1;
χ2 = 32.86; P < 0.0001).
Tail raising This posture was exhibited by females only
(n = 42 displays by females; Fig. 2). A female’s body size
strongly affected display frequency, with larger females dis-
0
60 65 70 75 80 85 90
female snout-vent length (mm)
Fig. 3. Effect of female body size (SVL) on the number of tail-raise
displays the lizard performed during our observation periods. Each
point represents data for a single female rock lizard. See text for details
of statistical analysis of these data
playing the tail-raise posture more often than did smaller
females (r = 0.62; P < 0.01; Fig. 3).
Gular extension Gular extension was observed for males
only and amounted to 24% of total directed displays by this
sex (n = 47 displays; Fig. 2).
Arching of back Both sexes performed this display. Males
did so more often than females (Fig. 2; Mann–Whitney
U = 54.00; P < 0.015, n = 32 and 18 displays for males and
females, respectively) but this difference was because of the
overall higher display rate of males. When calculated as a
proportion of all directed displays by each lizard, the sex
difference fell well short of significance (Mann–Whitney
U = 90.00; P = 0.36).
Leg extension Leg extensions were performed more often
by males than by females, although the difference was only
marginally significant in terms of absolute numbers (Mann–
Whitney U = 65.50; P = 0.049, n = 39 and 25 displays for
males and females, respectively) and not even close to sig-
nificance in terms of the proportions of directed displays by
each animal (Mann–Whitney U = 102.00; P = 0.68).
Dorsal flattening Dorsal flattening was performed mostly
by males (Fig. 2; absolute numbers – Mann–Whitney U =
1.00; P < 0.0001; proportions – Mann–Whitney U = 10.00;
P < 0.0001; n = 90 and 2 displays for males and females
respectively).
Directed and non-directed displays
Our data suggest a primary dichotomy between simple
push-ups and more complex displays. “Non-directed” dis-
plays (i.e. those for which no other animal was seen within
10 m radius of the displaying lizard) almost invariably con-
sisted of push-ups only, with no additional posturing on
>99% of occasions. In contrast, directed displays were
 279

invariably accompanied by at least one of the above five
body postures associated with push-ups.
Push-ups also differed from the other displays in their
relationship to movement by the lizard performing the dis-
play. Most (89%) non-directed displays occurred immedi-
ately after the lizard had moved from one location to
another, whereas this was true of only approximately half
(48%, = 142/296) of the directed displays. Most of these
latter cases (92/142, = 65%), however, occurred immedi-
ately after some movement in response to other lizards or
animals in the vicinity of the displayer. The remaining 52%
of “directed” push-up displays (154/296, or less than one-
quarter of all push-ups) were produced by sedentary
lizards.
Social context of directed displays
The displays occurred in different social contexts. We focus
on “directed” displays in which another animal was present
within 10 m radius and, hence, may have been the stimulus
eliciting that display. These potential stimuli were of three
main types: conspecific males, conspecific females, or other
species (monkeys, squirrels, or birds). Importantly, males
and females differed in the numbers of displays they exhib-
ited in response to these different types of stimulus
(Fig. 4a). For example, the proportion of dorsal-flatten
displays was higher in males than females in response to
birds (Mann–Whitney U = 7.00; P < 0.0001; n = 83 and 1 dis-
plays for males and females, respectively) and squirrels

Fig. 4. Effect of social context (a) total number of displays (d) leg-extend
and sex on the relative frequency
7 100
of different types of “directed”
display by rock lizards. These
proportions are mean values (and
6 female
number of displays

80
standard errors) based on 5 male

% of displays
numbers of each display type as a
proportion of the total number of 60
4
displays recorded for each
individual lizard in response to 3
40
each type of stimulus. These data
are based on a total of 87 displays 2
by 16 female lizards, and 208 20
1
displays by 14 males
0 0
female male monkey squirrel bird female male monkey squirrel bird
lizard lizard lizard lizard

(b) tail-raise (e) arch-back

100 100

80 80
% of displays
% of displays

60 60

40 40

20 20

0 0
female male monkey squirrel bird female male monkey squirrel bird
lizard lizard lizard lizard

(c) gular-extend (f) dorsal flatten

80 100

60 75
% of displays
% of displays

40 50

20 25

0 0
female male monkey squirrel bird female male monkey squirrel bird
lizard lizard lizard lizard
280
(Mann–Whitney U = 22.50; P < 0.02; n = 7 for males and 1
for females), and the proportion of gular-extension displays
was higher towards conspecific males (Mann–Whitney
U = 22.50; P < 0.0001) than to conspecific females (Mann–
Whitney U = 6.00; P < 0.02).
Simple push-ups These displays were non-directed
Tail raising This body posture was typically exhibited by
females in the presence of male conspecifics (n = 39
records) and, much more rarely, to female conspecifics
(n = 3). Because females displayed so rarely to other
females (Fig. 4a), however, the latter cases actually consti-
tuted a high proportion of records in this category (Fig. 4b).
The proportion of tail-raise displays differed according to
the stimulus to which the female lizard was exposed
(Kruskal–Wallis H = 16.26; df = 4; P < 0.0001).
Gular extension Gular extension was observed only in
males, and was directed both to conspecific males and
females but not to heterospecific animals (Fig. 4c; Kruskal–
Wallis H = 33.14; df = 4; P < 0.0001).
Leg extension This posture was seen in both males and
females during directed displays, but the social context dif-
fered between sexes. Most leg-extension displays by males
occurred in the presence of conspecifics, especially females
(Fig. 4d). In contrast, female leg-extension displays were
recorded primarily in response to squirrels (Fig. 4d). For
both sexes, the response was elicited non-randomly by dif-
ferent stimuli (males, Kruskal–Wallis H = 25.11; df = 4;
P < 0.002; females, Kruskal–Wallis H = 27.63; df = 4;
P < 0.001).
Arching of back Both the sexes displayed arched-back pos-
tures, but in different contexts. Males arched their back on
encountering animals such as monkeys or squirrels (Fig. 4e)
whereas females arched their back to conspecific males
(Fig. 4e, f). For both sexes, the incidence of the arched-back
display was affected by the stimulus involved (males,
Kruskal–Wallis H = 35.3; df = 4; P < 0.0001; females,
Kruskal–Wallis H = 21.14; df = 4; P < 0.0001).
Dorsal flattening Males employed this body posture during
directed displays when birds flew above them or (less often)
in response to nearby squirrels (Fig. 4f; Kruskal–Wallis
H = 45.54; df = 4; P < 0.0001). In females, dorsal flattening
was observed on two occasions only.
Discussion
The spectacular sexual dichromatism of Indian rock lizards
is accompanied by equally spectacular sex divergence in
ritualized visual displays. For both sexes most displays are
simple push-ups with head-bobbing, as seen for a wide
diversity of lizard species from other continents and other
phylogenetic lineages (Jenssen 1977; McMann 1993;
Martins 1991, 1993; Martins and Lamont 1998; Watt and
Joss 2003; Martins et al. 2004). These movements are usu-
ally performed when the lizards are solitary, and thus func-
tion as non-directed displays; we discuss their possible
significance below. Importantly, however, the sexes diverge
even in this simplest of displays, with males usually per-
forming a more intense version (i.e. double rather than
single head-bobs). All the other display postures of P. dor-
salis are more complex, and performed in specific social
and ecological contexts. That specificity provides strong
clues to the biological function of each display, and the
reasons why the sexes usually differ in the form and fre-
quency of each posture. Below, we review this evidence to
generate hypotheses on the functional significance of these
components of the visual-communication system in Indian
rock lizards.
Field and laboratory studies have been performed on the
social behaviour of many lizard species, although attention
has been disproportionately focussed on a relatively small
number of major phylogenetic lineages. For example, sev-
eral phrynosomatid species from North America have
become “model organisms” within this research field (e.g.
see Martins 1991, 1993; Ord and Blumstein 2002; Fox et al.
2003). To the best of our knowledge, our study is the first
analysis of social postures within any Indian lizard species;
it is also one of the first detailed field-based studies of any
agamid, except for distantly-related Australian and Malay-
sian taxa, for example Chlamydosaurus (Shine 1990),
Ctenophorus (Gibbons 1979; Olsson 1995), and Draco
(Mori and Hikida 1994). Unsurprisingly, then, some of the
specific display postures exhibited by P. dorsalis differ from
those previously described for other lizard species. There is,
nonetheless, broad similarity, perhaps enforced by the
broad morphological similarities among many diurnal aga-
mid and iguanid taxa (Melville et al. 2001).
Most previous studies on agamid lizard behaviour have
focussed on behavioural postures and action patterns asso-
ciated with displays, fighting, courtship and mating (e.g.
Carpenter et al. 1970; Brattstrom 1971; Murphy et al. 1978;
Gibbons 1979; Shine 1990; Mori and Hikida 1994; Watt and
Joss 2003) and less attention has been paid to the functional
and contextual use of pushup displays (but see Shine 1990
and Watt and Joss 2003). Further, many of these studies
have relied on staged encounters (e.g. Gibbons 1979; Watt
and Joss 2003), rather than displays performed in response
to natural encounters either with conspecifics or with other
species (for example predators). Thus, our data set is
unusual in these respects and reveals a surprisingly complex
and context-dependent array of behavioural postures in
these Indian lizards.
The most frequent display by P. dorsalis, as by many
other lizard species, was a simple push-up with head-
bobbing. Because this display was performed so frequently,
by both sexes, including juvenile P. dorsalis (personal obser-
vations), in the absence of any other overt stimuli (other
animals), it seems unlikely to convey important social infor-
mation about attributes such as the sex, body size, or dom-
inance status of the individual performing the display. In
this respect, P. dorsalis differs from of many other lizards
including the Australian agamid lizard Amphibolurus
muricatus (e.g. Watt and Joss 2003). Displays performed
spontaneously in the absence of obvious opportunity for
visual social stimuli from nearby conspecifics and without
the accompaniment of movements and body postures have

been categorized as “assertion” displays in Sceloporus and
several anoline species (Carpenter and Ferguson 1977).
Assertion display frequency, number of locomotory pauses,
and distance moved are correlated in Sceloporus and
several anoline species. In a few populations different
individuals exhibit different cadences of the assertion dis-
plays and individuals within a population may also exhibit
more than one cadence (Carpenter and Ferguson 1977). As
push-up displays are often conspicuous and energetically
costly, the movement may serve non-social functions: for
example, rapid vertical motion of the lizard’s head may
enhance distance perception by use of parallax. Alterna-
tively, lizards may use this simply to broadcast their pres-
ence and status in the new place, because these displays
often occur immediately after a lizard changes its position
(Dugan 1982).
The function of directed displays can be inferred with
more confidence, especially in when there is strong sex
divergence in expression of particular postures. For rock
lizards, one posture (gular extension) was performed by
males only, and one (tail-raise) by females only (Fig. 2). At
one level explanation of this sex-specificity is trivial –
morphological differences between the sexes make each
one incapable of performing the display characteristic of the
other. Thus, female rock lizards lack the enlarged coloured
dewlap needed for gular extension and males have greatly
distended tail bases (to accommodate the paired hemipenes
and protective muscles) that might hinder an adult male’s
attempts to lift his tailbase from the substrate. Although
sexual dimorphism in tailbase thickness is widespread in
squamate reptiles, it is particularly well-developed in Indian
rock lizards. That is, male P. dorsalis have relatively thick
tailbases compared not only with conspecific females but
also with males of most other lizard species. Male P. dorsalis
also lose their tails more frequently than do females
(unpublished data), further reducing the ability of males to
perform “tail-raise” displays. We speculate that selection
may have favoured these sex-specific displays because, for
both displays, the posture offers an “honest” signal about
the performing lizard’s gender.
Tail-raising behaviour in phrynosomatid lizards has been
attributed a role in courtship interactions and female rejec-
tion behaviour (e.g. Martins 1991, 1993). However, tail-
raising by female P. dorsalis seems unlikely to function
either in courtship interactions or in female rejection behav-
iour, because females raised their tails frequently during
non-breeding periods also. A previous analysis of tail-
coiling behaviour in agamid lizards (Ctenophorus decresii
species complex) advocated a role for species recognition
in this display (Gibbons 1979), but a sex-identification role
for tail-raising (as we suggest for P. dorsalis) has been
reported in studies on other organisms. For example,
monogamous butterfly fish use a tail-up display to recognize
their partners, apparently to facilitate sex recognition and,
thus, to reduce the probability of a “mistaken” attack by the
partner (Yabuta 2002). Similarly, magpie birds use tail dis-
plays during both intersexual and intrasexual encounters,
and multiple elements of this display provide information
about the identity of the signaller with regard to age and
281
sex (Blanco and Puente 2002). In birds, tails play an impor-
tant role in aerial locomotion and, thus, there may be a
conflict between adaptations for signalling and adaptations
for aerodynamic effectiveness. Such tradeoffs limit the
extent of tail modification for signalling, and help to main-
tain an “honest” link between tail attributes and mate qual-
ity (e.g. Andersson 1982, 1986), as we have suggested above
for the swollen tailbases of male rock lizards. Professional
herpetologists frequently assign sex to captured squamates
simply on the basis of this gross difference (Fitch 1987), and
Indian rock lizards may do the same.
Another display with a strong sex difference in expres-
sion was “dorsal-flatten”: males exhibited this posture much
more often than did females (Fig. 2). The posture clearly
makes a lizard less visible to a predator flying overhead,
and, indeed, was elicited mostly by birds (and, occasionally,
by squirrels leaping over the lizards: Fig. 4). Similar flatten-
ing against the substrate has been reported in many other
lizard species, and has been widely interpreted as an anti-
predator (defensive) tactic (Heatwole 1970; Brattstorm
1971; Leal and Rodriguez-Robbes 1997). Why, then, do we
see such a strong sex difference in the frequency of this non-
social display among rock lizards? The answer may be hab-
itat differences between males and females that expose
them to over-flying predators to different extents. Selection
of open high perches by males, presumably to enhance ter-
ritory surveillance, means they are frequently in open
exposed habitats and in turn, frequently encounter poten-
tially dangerous birds in conditions where crypsis via dorsal
flattening offers an effective response.
Why, then, did female lizards typically respond to birds
by back-arching rather than dorsal flattening? Females are
usually at or close to ground level where they are partially
hidden from flying birds, and any birds they do encounter
are likely to be walking on the ground rather than flying.
Thus, the important issue is crypsis from a side view rather
than from above. We have recorded several instances of
predation by birds on these lizards at our study sites
(Radder et al. 2006). Alternatively, P. dorsalis may use this
back-arching as a pursuit-deterrent signal to potential pred-
ators, and/or simply to deter predators by increasing the
apparent size of the displayer. In other lizards the same
posture may serve as a predator-deterrent signal and for
social interactions (Leal and Rodriguez-Robbes 1997).
Last, and most puzzling, leg-extension displays were per-
formed by both sexes but in different contexts. Males leg-
extended to conspecifics, especially females, suggesting that
the display asserts dominance. Females leg-extended prima-
rily to other species, however, especially squirrels.
Although this study is only an initial step in understand-
ing the communication system of P. dorsalis, our data are
useful not only to suggest biological functions for most
displays, but also to provide robust support for some of
these ideas. One notable result is that sex-based divergence
in display postures and frequencies can arise for many rea-
sons and can convey several different kinds of information.
For example, the raised-tail posture may provide informa-
tion about a female’s receptivity to males; the gular-
extension posture provides information about social factors
282
(presumably territory ownership) in males; and the leg-
extension posture may provide different kinds of infor-
mation depending upon whether it is performed by a male
or a female. Yet other differences between the sexes in
display frequencies are secondary consequences of other
behavioural differences between the sexes, notably habitat-
selection, such that males encounter different kinds of pred-
ator (Fig. 4), in different situations, than do conspecific
females.
Diurnal lizards, with their highly ritualized, complex, and
context-dependent visual signals, are ideal models to exam-
ine the evolution of communication systems in animals (Fox
et al. 2003; Marler 2003). Our preliminary field studies on
P. dorsalis suggest that this species may be very suitable for
further research, for example experimental presentation of
stimuli to tease apart correlation from causation. Such stud-
ies could test the hypotheses we have generated on the basis
of observational data, and ultimately enable better under-
standing of the ways in which these animals convey
information to each other as part of their social interactions.
In turn, such understanding based on a study organism that
is far removed both geographically and phylogenetically
from the usual “model species” within this field of research
has the potential to significantly enhance the generality of
our ideas and conclusions about social communication in
animals.
Although reptiles have traditionally been regarded as
exhibiting only relatively simple, “primitive” social behav-
iour compared with the more “advanced” endothermic ver-
tebrates, recent research has strongly challenged this view.
There is increasing evidence of phenomena such as long-
term monogamy (Bull 1988), complex kin-based sociality
(O’Connor and Shine 2003) and individual recognition
(Bull et al. 1994) in lizards, and surprisingly high similarity
in traits such as ritualistic social displays and basal ganglia
structures across diverse amniote species including lizards
and humans (Baxter 2003). Our study supports and extends
earlier work on other lizard taxa by showing that these
animals have a complex repertoire of context-dependent
displays. Interestingly, the “audience effect” in P. dorsalis
involves different responses not only to conspecific males
and females, and to different species (birds, squirrels, and
monkeys), but also depends upon the location of the
“audience”. Is it reasonable, then, to refer to the rock-lizard
display system as a “language”? The complexity of any com-
munication system is limited by the number and diversity
of components available, and the ways in which those com-
ponents can be combined. In this respect, the effects of
signaller sex and audience context on frequencies of differ-
ent display types in P. dorsalis suggest that these postures
may convey much information. Although conventional
usage might deem it absurd to conclude that lizards have a
language, the rock lizards would qualify under Snowdon’s
(1990) definition (“any rule-based system that generates
predictable sequences of behavior”). Our data show that
the push-up displays of P. dorsalis have a distinct structure
with explicit rules; for example, the type of display is gov-
erned by sex and size of the producer, the type of receiver
(audience effect), and the microhabitat where the display is
performed. Thus, according to Snowdon’s definition, P. dor-
salis does indeed have a language.
Acknowledgements This work was supported by a Council of Scien-
tific and Industrial Research’s (CSIR, India) Associateship awarded to
RSR, Special Assistance Programme (SAP-II) University Grant Com-
mission (UGC, India) sanctioned to Department of Zoology Karnatak
University Dharwad and by the Australian Research Council (to RS).
Authors thank Professor R.Gadagkar, Center for Ecological Sciences,
Indian Institute of Science, Bangalore for his keen interest in this study.
We thank Principal Chief Conservator of Forests, Bangalore for per-
mitting us to study rock lizards.
References
Andersson M (1982) Female choice selects for extreme tail length in a
widowbird. Nature 299:818–820
Andersson M (1986) Evolution of condition-dependent sex ornaments
and mating preferences: sexual selection based on viability differ-
ences. Evolution 40:804–816
Baird TA, Timanus DK, Sloan CL (2003) Intra- and intersexual varia-
tion in social behavior: Effect of ontogeny, phenotype, resources and
season. In: Fox SF, McCoy JK, Baird TA (eds) Lizard social behav-
ior. The John Hopkins University Press, Baltimore, pp 7–46
Baxter LR Jr (2003) Basal ganglia systems in ritualistic social displays:
reptiles and humans; function and illness. Physiol Behav 79:451–460
Blanco G, Puente JDL (2002) Multiple elements of the black-billed
magpie’s tail correlate with variable honest information on quality
in different age/sex classes. Anim Behav 63:217–225
Brattstrom BH (1971) Social and thermoregulatory behavior of the
bearded dragon, Amphibolurus barbatus. Copeia 1971:484–497
Brooks DR, Couldridge V (1999) Multiple sexual ornaments coevolve
with multiple mating preferences. Am Nat 154:37–45
Bull CM (1988) Mate fidelity in an Australian lizard Trachydosaurus
rugosus (Scincidae). Behav Ecol Sociobiol 23:45–49
Bull CM, Doherty M, Schultz LR, Pamula Y (1994) Recognition of
offspring by females of the Australian skink, Tiliqua rugosa. J Her-
petol 28:117–120
Carpenter CC (1965) Aggression and social structure in iguanid lizards.
In: Milstead WW (ed) Lizard ecology: a symposium. University of
Missouri Press, Columbia, pp 87–105
Carpenter CC, Ferguson GW (1977) Variation and evolution of stereo-
typed behavior in reptiles. In: Gans C, Tinkle DW (eds) Biology of
reptilia: ecology and behavior, A, vol 7. Academic, New York, pp
335–554
Carpenter CC, Badham JA, Kimble B (1970) Behavior patterns of
three species of Amphibolurus (Agamidae). Copeia 1970:497–505
Castellano S, Giacoma C, Dujsebayeva T (2000) Morphometric and
advertisement call geographic variation in polyploid green toads.
Biol J Linn Soc 70:341–360
Chappell MA, Zuk M, Kwan TH, Johnsen TS (1995) Energy cost of
an avian vocal display: crowing in red jungle fowl. Anim Behav
49:255–257
Clutton-Brock TH, Guiness FE, Albon SD (1982) Red deer: behavior
of two sexes. University of Chicago Press, Chicago
Cooper WE, Greenberg N (1992) Reptilian coloration and behavior.
In: Gans C, Crews D (eds) Biology of reptilia, physiology e hor-
mones, brain and behavior, vol 18. University of Chicago Press,
Chicago, pp 298–422
Davies NB (1992) Dunnock behavior and social evolution. Oxford
University Press, Oxford
DeCourcy KR, Jenssen TA (1994) Structure and use of male headbob
signals by the lizard Anolis carolinensis. Anim Behav 47:251–262
Dugan B (1982) A field study of the headbob displays of male green
iguanas (Iguana iguana): variation in form and context. Anim Behav
30:327–338
Dyer FC (2002) The biology of the dance language. Annu Rev Entomol
47:917–949
Evans MR (1991) The size of adornments of male scarlet-tufted mala-
chite sunbirds varies with environmental conditions as predicted by
handicap theories. Anim Behav 42:797–803

Evans MR, Norris K (1996) The importance of carotenoids in signaling
during aggressive interactions between male firemouth ciclids
(Cichlasoma meeki). Behav Ecol 7:1–6
Fitch HS (1987) Collecting and life history techniques. In: Siegel RA,
Collins JT, Novak SS (eds) Snakes ecology and evolutionary biology.
Macmillan, New York, pp 143–164
Fox SF, McCoy JK, Baird TA (2003) The evolutionary study of social
behavior and the role of lizards as model organisms. In: Fox SF,
McCoy JK, Baird TA (eds) Lizard social behavior. John Hopkins
University Press, Baltimore, pp xi–xiv
Gibbons JRH (1979) The hind leg pushup displays of the Amphibolu-
rus decresii species complex (Lacertilia: Agamidae). Copeia 1979:29–
40
Grant BW, Dunham AE (1988) Thermally imposed time constraints on
the activity of the dessert lizard, Sceloporus merriami. Ecology
69:167–176
Hasson O, Hibbard R, Ceballos G (1989) The pursuit deterrent func-
tion of tail-wagging in the zebra-tailed lizard (Callisaurus dra-
conoides). Can J Zool 67:1205 –1209
Heatwole H (1970) Thermal ecology of the desert dragon, Amphibolu-
rus inermis. Ecol Monog 40:425–457
Hews DK, Quinn VS (2003) Endocrinology of species differences in
sexually dichromatic signals: Using organization and activation
model in a phylogenetic frame work. In: Fox SF, McCoy JK, Baird
TA (eds) Lizard social behavior. John Hopkins University Press,
Baltimore, pp 253–277
Holberton RL, Able KP, Wingfield JC (1989) Status signalling in dark-
eyed juncos, Junco hyemalis: plumage manipulations and hormonal
correlates of dominance. Anim Behav 37:681–689
Jenssen TA (1977) Evolution of anoline lizard display behavior. Am
Zool 17:203–215
Johnstone RA (1995) Honest advertisement of multiple qualities using
multiple signals. J Theor Biol 177:87–94
Johnstone RA (1996) Multiple displays in animal communications:
backup signals and multiple messages. Philos Trans Roy Soc Lond B
531:329–338
Johnstone RA (1997) The evolution of animal signals. In: Krebs JR,
Davies NB (eds) Behavioural ecology: an evolutionary approach, 4
edn. Blackwell Scientific, Oxford, pp 55–178
Leal M (1999) Honest signalling during prey-predator interactions in
the lizard Anolis cristatellus. Anim Behav 58:521–526
Leal M, Rodriguez-Robles JA (1997) Signalling displays during
predator-prey interactions in a Puerto Rican anole, Anolis cristatel-
lus. Anim Behav 54:1147–1154
Leal M, Fleishman LJ (2002) Evidence for habitat partitioning based
on adaptation to environmental light in a pair of sympatric lizard
species. Proc Roy Soc Lond B 269:351–359
Leger DW, Didrichson IA (1994) An assessment of data pooling and
some alternatives. Anim Behav 48:823–832
Liebal K, Pika S, Tomasello M (2004) Social communication in sia-
mangs (Symphalangus syndactylus): use of gestures and facial
expression. Primates 45:41–57
Macedonia JM, Clark DL (2001) Headbob display analysis of the
Grand Cayman anole, Anolis conspersus. J Herpetol 35:300–310
Marler P (2003) Variations among individuals: introduction. In: Fox SF,
McCoy JK, Baird TA (eds) Lizard social behavior. The John Hopkins
University Press, Baltimore, pp 3–6
Martins EP (1991) Individual and sex differences in the use of the
push-up display by the sagebrush lizard, Sceloporus graciosus. Anim
Behav 41:403–416
Martins EP (1993) Contextual use of the push-up display by the sage-
brush lizard, Sceloporus graciosus. Anim Behav 45:25–36
Martins EP, Lamont J (1998) Estimating ancestral states of a commu-
nicative display: a comparative study of Cyclura rock iguanas. Anim
Behav 55:1685–1706
Martins EP, Labra A, Halloy M, Thompson JT (2004) Large-scale
patterns of signal evolution: an interspecific study of Liolaemus liz-
ard headbob displays. Anim Behav 68:453–463
McMann S (1993) Contextual signalling and the structure of dyadic
encounters in Anolis carolinensis. Anim Behav 46:657–668
Melville J, Schulte II JA, Larson A (2001) A molecular phylogenetic
study of ecological diversification in the Australian lizard genus
Ctenophorus. J Exp Zool 291:339–353
Møller AP, Pomiankowski A (1993) Why have birds got multiple sexual
ornaments? Behav Ecol Sociobiol 32:167–176
283
Mori A, Hikida T (1994) Field observations on the social behavior of
the flying lizard, Draco Volans sumatranus, in Borneo. Copeia
1994:124–130
Murphy JB, Lamoreaux WE, Carpenter CC (1978) Threatening behav-
iour in the angle-headed dragon, Goniocephalus dilophus (Reptilia,
Lacertilia, Agamidae). J Herpetol 12:455–460
O’Connor D, Shine R (2003) Lizards in “nuclear families”: a novel
reptilian social system in Egernia saxatilis. Mole Ecol 12:743–752
Olsson M (1992) Contest success in relation to size and residency in
male sand lizards, Lacerta agilis. Anim Behav 44:386–388
Olsson M (1994) Nuptial coloration in the sand lizard, Lacerta agilis:
an intra-sexually selected cue to fighting ability. Anim Behav 48:607–
613
Olsson M (1995) Forced copulation and costly female resistance behav-
ior in the Lake Eyre dragon, Ctenophorus maculosus. Herpetologica
51:19–24
Ord TJ, Blumstein DT (2002) Size constraints and the evolution of
display complexity: why do large lizards have simple displays? Biol
J Linn Soc 76:145–161
Ord TJ, Blumstein DT, Evans CS (2001) Intrasexual selection predicts
the evolution of signal complexity in lizards. Proc Roy Soc Lond B
268:737–744
Podos J (2001) Correlated evolution of morphology and vocal signal
structure in Darwin’s finches. Nature 409:185–188
Prum RO (1998) Sexual selection and the evolution of mechanical
sound production in manakins (Aves: Pipridae). Anim Behav
55:977–994
Quinn VS, Hews DK (2000) Signals and behavioral responses are not
coupled in males: aggression affected by replacement of an evolu-
tionary lost color signal. Proc R Soc Lond B 267:755–758
Radder RS, Saidapur SK, Shanbhag BA (2005) Population density,
microhabitat use and activity pattern of the Indian rock lizard, Psam-
mophilus dorsalis (Agamidae). Curr Sci 89:560–565
Radder RS, Saidapur SK, Shanbhag BA (2006) Big boys on the top:
effects of body size, sex and reproductive state on perching behav-
iour in the tropical rock dragon, Psammophilus dorsalis. Anim Biol
(in press)
Sarkar HBD, Shivanandappa T (1989) Reproductive cycles of reptiles.
In: Saidapur SK (ed) Reproductive cycles of Indian vertebrates.
Allied Publishers Limited, New Delhi, pp 225–272
Shanbhag BA (2002) Reproductive biology of Indian reptiles. Proc
Indian Natl Sci Acad B 68:497–528
Shine R (1990) Function and evolution of the frill of the frill neck
lizard, Chlamydosaurus kingii (Sauria: Agamidae). Biol J Linn Soc
40:11–20
Sinervo B, Lively CM (1996) The rock-paper-scissor game and the
evolution of alternative male strategies. Nature 380:240–243
Sinervo B, Miles DB, Frankino WA, Klukowski M, DeNardo DF
(2000) Testosterone, endurance, and Darwinian fitness: natural and
sexual selection on the physiological bases of alternative male
behaviors in side blotched lizards. Horm Behav 38:222–233
Snowdon CT (1990) Language capacities of non-human animals. Year-
book of Phys Anthro 33:215–243
Stamps JA (1977) Social behavior and spacing patterns in lizards. In:
Gans C, Tinkle DW (eds) Biology of reptilia: ecology and behavior,
a, vol 7. Academic, New York, pp 335–354
Tokarz RR (1985) Body size as a factor determining dominance in
staged agonistic encounters between male brown anoles (Anolis
sagrei). Anim Behav 49:661–669
Tomasello M, Zuberbuehler K (2002) Primate vocal and gestural
communication. In: Bekoff M, Allen CS, Burghardt G (eds) The
cognitive animal: empirical and theoretical perspectives on animal
cognition. MIT Press, Cambridge, pp 293–329
Watt MJ, Joss JMP (2003) Structure and function of visual displays
produced by male Jacky dragon, Amphibolurus muricatus, during
social interactions. Brain Behav Evol 61:172–183
Whiting MJ, Nagy KA, Bateman PW (2003) Evolution and mainte-
nance of social status-signaling badges: experimental manipulations
in lizards. In: Fox SF, McCoy JK, Baird TA (eds) Lizard social behav-
ior. The John Hopkins University Press, Baltimore, pp 7–46
Wiens JJ (1999) Phylogenetic evidence for multiple losses of a sexually
selected character in phrynosomatid lizards. Proc Roy Soc Lond B
266:1529–1535
Yabuta S (2002) Uncertainty in partner recognition and the tail-up
display in monogamous butterfly fish. Anim Behav 63:165–173