Behavioural Ecology of The Peninsula Dragon Lizard, Ctenophorus Fionni

BEHAVIOURAL ECOLOGY OF
THE PENINSULA DRAGON LIZARD

CTENOPHORUS FIONNI
Greg Johnston
School of Biological Sciences

Flinders University of South Australia
A thesis submitted for the degree of Doctor of Philosophy
March 1997
"As a child, if the family turned their backs for an instant, he slipped
away to entertain himself in solitary pastimes that provoked pitiless
ridicule: he gathered stones from the river and polished them for the
pleasure of seeing their colors shine; he scouted the dismal landscape
looking for dry leaves to arrange in artistic compositions; he was
moved to tears by a sunset, wanting to capture it forever in a line of
poetry or in a painting he could imagine but felt incapable of
realizing. Only his mother accepted his peculiarities, seeing them not
as signs of perversions but as evidence of a soul that was different.•
Isabel Allende
Of Love and Shadows
For Barbara Johnston (nee Baker)

AUTHOR'S DECLARATION
I certify that this thesis does not incorporate without

acknowledgement any material previously submitted for a degree or
diploma in any university; and that to the best of my knowledge and
belief it does not contain any material previously published or
written by another person except where due reference is made in the
text.
G.R. Johnston
©This thesis is copyright. Apart from any fair dealing for the purpose of
private study, research, criticism, or review no part may be reproduced by any
process without written permission from the author.
SUPERVISOR'S DECLARATION
I believe that this thesis is properly presented and conforms to the

specifications of thesis presentation in the University and is prima
facie worthy of examination.
C.M . Bull
'
A mature male Peninsula Dragon, Ctenophorus fionni

(photo: John Cornish)
CONTENTS
Title Page
Dedication
Declarations
Contents
Acknowledgments
SUMMARY
1. Introduction .......................................................... . 1
2. Sexual Dimorphism ........... ................... ... ......... ... .... 37
3. Reproduction . . ... . .. . .. . . . .. . . .. . .. .. . .. . .. . ... .. . . .. .. . .. .. . . . . .. . . . 70
4. Growth and Survivorship: Proximate causes of SexuaJ

.
S1ze D'1morp h'ism .,............................................... .. 96
5. Ecological Differences between Males and Females ...

128
6. ].\,fating System and the Opportunity for Sexual

Selection . . . . . . . . . . . . . . . . . . . . .. . . . . . .. . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . 17 5
7. Targets of Sexual Selection in Males ........................ 210
8. Mate Choice or Male Competition ? . .. . ... .. .. .. . . . .. ... .. . 240
9. Synthesis ............................................................... 258
References ....................................................................... 268

Appendices
ACKNOWLEDGEMENTS
This project was formally supervised by Associate Pr'ofessor C.
Michael Bull, and was funded by an Australian Postgraduate
Research Scholarship, a Flinders University Scholatship and an Ethel
Mary Read Award from the Royal Zoological Society of New South
Wales.
Mats Olsson unwittingly provided the inspiration for this project,
long ago, in a little caravan, somewhere near Lake Eyre.
Errol, Elizabeth & Mark Carmody allowed me to work on their
property. Andrew & Vicki Gohl provided welcome showers,
conversation, meals in Cowell whenever I got sick of staring at
bloody lizards during field trips. The Whyalla Amatuer Reptile
Keepers Association, Ben Baghurst, Anne Brow, Paula Dempsey,
Richard Evans, Rachel Furbank, Michelle Hale, Brett Johnston, Colin
Kranz, Cra,ig Martins, Paul Mazdon, and Elizabeth Yeatman helped
me in the field.
At Flinders University Russ Baudinette, Mike Bull, Peter Daenke,
Dick Edyveane, Mike and Tracy Freake, Sally Fryar, Tabatha Hall,
Jon Havenhand, Peter Hudson, Mark Jerisevic, Greg Kirby, Louise
Lawson, Duncan Mackay, Jim Mitchell, Yvonne Pamula, Rolf Prager,
Ken Sanderson, Grant Scott, Rod Wells, Bruce White, Sue Willmot
and Tatia Zubrinich contributed to the project in various ways.
Thanks guys.
Grant Fleming and Wendy Sysouphat provided office and laboratory
facilities at the Whyalla Campus of the University of South Australia.
Jim & Karen Scriven at the Whyalla Fauna Park for laughs and cages.
Colin Kranz helped in many ways to establish the basic reproductive
biology of Peninsula Dragons.
Barbara and Ron Johnston (alias Mum & Dad), and Craig and Linda
Martins for keeping open houses to me and miscellaneous hangers
on.
Mike Bull ,read and passed comments on the entire thesis. Paula
Dempsey and Jeremy Roberston read several chapters each. Paula
also provided support in many other ways.

SUMMARY
This thesis aimed to explain the evolution of sexually dimorphic

traits in an Australian agamid lizard, the Peninsula Dragon
(Ctenophorus fionni). The study investigated (1) the extent to
which sampling or demographic artefacts, fecundity selection,
natural selection and sexual selection can explain the dimorphic
traits in this species, and (2) the relative importance of inter- vs
intrasexual selection acting upon traits which were subject to
sexual selection.
Adult male and female Peninsula Dragons differ from each other
in body size, shape and colouration. Male and female Peninsula
Dragons did not differ in any measured trait before they become
mature, but upon reaching maturity males and females diverge
from a common juvenile pattern. Males tend to grow to a larger
size than females and exhibit an increase in relative growth.
Males develop larger heads and generally darker dorsal
colouration that females. Adult males also develop unique, sex-
limited traits, such as a yellow throat. These patterns suggest
that selection for sexually dimorphic traits is acting mostly upon
adults, and on adult males in particular, in this species.
Peninsula Dragons mate and reproduce during the austral spring.

Most females produced a single clutch of eggs per year, but some
females in some populations produced multiple clutches. Clutch size
varied between two and six. There was no relationship between
female body size and clutch size, so there is no mechanism by which
fecundity selection might act on female body size.
The pattern of sexual size dimorphism in Peninsula Dragons appears

to be largely a result of differences in growth pattern between the
sexes. Males grow to a larger asymptotic size than females, but the
full extent of the size difference was not realised due to high
mortality among large males. This is presumably a result of indirect
costs of reproduction (such as defending a home range) bourn by
males.
There is no evidence for the hypothesis that sexually dimorphic traits

have evolved as a result of ecological differences between the sexes.
No differences could be found between male and female Peninsula
Dragons in diet, activity times, or thermal preferences. Differences in
thermal biology and perch use in the field appear to be a secondary
effect of increased vigilance by territorial males for potential mates
and rivals. These relationships support the argument that ecological
differences may be the result, rather than a cause, of differences
between the sexes.
The mating system of Peninsula Dragons is best described as

resource defence polygyny. Older males defend home ranges
containing rock crevices, which appear to be used as refuges from
climatic extremes and predators. Many younger, mature males live
'within older males' home ranges. For the most part only older,
territory holding males pair (and presumably mate) with one or more
females. Males spend more time seeking mates than do females and
are able to reproduce at a higher rate than females. Males show no
parental care and exhibit greater variance in reproductive success
than females. These factors combine to provide an opportunity for
sexual selection to act on males. The circumstances of the mating
system permit the action of both intrasexual and intersexual
selection.
A selection analysis identified body size and head width as

independent targets of sexual selection in one cohort. Head length
was identified as a target of weak sexual selection in another
cohort. Pairing between males and females was used as the
measure of fitness. Pairing was interpreted as mate guarding
behaviour by males and was used as an index of mating success.
Body size and head size were both important (but independent)
correlates of territorial status in the field, and also whether a
male won or lost in staged fights. These f'mdings suggest that
large body size and large head size in male Peninsula Dragons can
be explained adequately by intrasexual selection. There was no
direct evidence that female mate choice was acting on body size
or shape in the selection analysis in the field. Furthermore, there
was no clear evidence that females were choosing among males in
lab oratory mate choice trials.
An experimental manipulation in the field showed that there was

no difference in pairing success between males painted to
resemble females, and normally coloured males. However, males
painted to resemble females lost twice as much weight as control
males. The home ranges of the males painted to resemble females
reduced in size, whereas the home range area of control males did
not change. Thus, the bright colouration of male Peninsula
Dragons appears to be important for home range defence, and
may function in sex recognition among males. Colour does not
appear to be a cue for intersexual selection.
In conclusion, sexual selection is the only hypothesis that appears to

explain the existence of sexually dimorphic traits in Peninsula
Dragons. Large body size, large head size and the characteristic male
colouration were all important for home range defence, and thus
provided males with access to mates. While intersexual selection
appears to be unimportant in this species, intrasexual selection is an
important force acting to promote sexually dimorphic traits in male
Peninsula Dragons.
CHAPTER 1
INTRODUCTION
Explaining the evolution of sexual dimorphic traits has. been a major

endeavour of evolutionary biologists since Darwin (1871) invoked
his theory of sexual selection to explain the phenomenon. This thesis
is concerned with the evolution of sexually dimorphic traits in the
Peninsula Dragon lizard ( Ctenophorus ftomli). I describe the sexual
differences, and ask why male and female Peninsula Dragons differ in
size, shape and colouration, and what selective pressures may have
given rise to these differences.
Sexually dimorphic traits are those which differ between males and
females. In some species sexual dimorphism results from traits which
occur in one sex, but not the other. The antlers of male red deer
( Cervus elephas) or the train of male peacocks (Pavo cristatus) are
examples of such sex-limited traits. Alternatively, both sexes may
possess a trait, but differ in the degree to which it is expressed.
Chapter 1 INTRODUCTION
Examples are sexual differences in body size in Elephant Seals

(Mirounga leonina) or tail length in widow birds (Euplectes progne).
The extent to which sexually dimorphic traits require a separate

evolutionary explanation to non-dimorphic traits is a major area of
controversy (Cronin 1991). In part this controversy has been fed by
the tendency to regard the phenomenon of sexual dimorphism as the
subject requiring explanation, rather that seeking explanations for
individual traits. It is important to recognise that organisms consist
of an array of traits, and each trait may be subject to a variety of
different selective pressures. Each trait must be regarded separately
because each may require a separate explanation. Sexual dimorphism
itself is not subject to selection, rather selection acts on particular
dimorphic traits. Many different sources of selection acting on
different components of fitness may contribute to the evolution of
each sexually dimorphic trait.
EVOLUTION, COMPONENTS OF FITNESS AND

SOURCES OF SELECTION
Evolution refers to changes in gene frequencies between generations.

Gene frequencies may change through random processes such as the
founder effect (Mayr 1963) or genetic drift (Wright 1931; Kimura
1983). Alternatively, evolution may occur under the influence of
selection (Darwin & Wallace 1858).
2
Chapter I INTRODUCTION
The founder effect and genetic drift can explain random (non
adaptive) differences among isolated populations. But these
stochastic processes cannot give rise to consistent differences
between the sexes within populations of diploid species because
genes from males and females must continually recombine through
sexual reproduction.
Selection occurs if there is a relationship between the expression of a

trait and the fitness of an organism (Endler 1986). For selection to
act on a trait, the trait must vary among individuals in a population.
However, selection on phenotypic traits can only give rise to an
evolutionary response if variation in the trait is heritable (Lande &
Arnold 1983). An individual's fitness, in this context, is the number
of offspring which survive and reproduce (Endler 1986).
Traits which show an evolutionary response to selection may be

subject to selection from one or more sources. These sources of
selection may be viewed as a hierarchy. Each source has a direct
fitness consequence for the organism, and each source has a clear
relationship with other sources of selection. Survival is a prime
determinant of fitness, and is the first source of selection. If an
organism survives to maturity, its fitness depends on its mating
success, the second source of selection. The number of offspring
produced per mating, or fecundity, may then have a further influence
on fitness. This is the third source of selection. Variation in fitness
due to differential survival is referred to here as natural selection.
3
Variation in fitness due to differential mating success is referred to

as sexual selecti0,n. Variation in fitness due to the number of
offspring produced per mating is referred to as fecundity selection.
A key reason for distinguishing between these three types of

selection is that they may be antagonistic in their effects on a trait.
For example sexually selected traits confer benefits in terms of
mating success, but may also impose costs in terms of survival
(Darwin 1871). An example is provided by the Tungara Frog
(Physa/aemus pustu/osus) in which males make themselves more
attractive to females by increasing the complexity of their calls, but
in so doing increase the likelihood of being detected and eaten by
predatory bats (Ryan 1985). However, in other cases, the different
types of selection may complement each other. For example, the
large claws of male fiddler crabs (Uca pugilator) are both attractive
to females and afford males some protection from predation by ibises
(Bildstein, McDowell & Brisben 1989).
To understand the evolution of any trait completely it is necessary to

understand all of the sources of selection acting on that trait (Arak
1988; Halliday 1992). An observed response to selection may be a
compromise between several selective forces. Body size provides a
good example because it represents a compromise between
conflicting selective pressures. Small size may optimise energetic
efficiency, and thus survival in both sexes. On the other hand, large
size in males may provide access to mates, and thus enhance mating
success. Large size in females may allow them to carry more
4
offspring, and offer a fecundity advantage. Thus body size may be a

compromise between all of these different sources of selection.
Traits may also show a response to selection acting on other
correlated traits.
Formal selection analyses using the approach developed by Arnold

and Wade (Wade & Arnold 1980; Arnold & Wade 1984a,b) set out to
identify the intensity, direction and form of selection arising from
different components of fitness, and due to different episodes of
selection. Although the practice of attributing different episodes of
selection to different kinds of selection has been questioned
(Halliday 1992), this approach seems to have some heuristic value.
Estimates of selection coefficients which can be assigned to different
episodes of selection according to their effects on different
components of fitness, permit one to assess the relative importance
of different evolutionary forces in natural populations (Endler 1986).·
Measures of phenotypic selection are mosi: useful when combined

with a thorough understanding of the biology of the organism being
studied. For example, temporal differences in selection may be
important in identifying ecological conditions under which particular
traits are adaptive (e.g. Schluter & Smith 1986). Such differences
might be ignored if one did not have the appropriate background
information on the natural history of an organism. When it is
necessary to conduct observations or experiments in the laboratory,
it is important to interpret the results in the light of conditions in the
field. For example, studies of the South African frog (Hyperolius
5
Chapter l INTRODUCTION
marmoratus) have shown: a clear preference among females for male

calls with particular properties in the laboratory (Dyson & Passmore
l 988a,b ), but in large field populations females mate
randomly(Telford & Dyson 1988). As it turns out, females are only
choosy when there are less than eight males in a chorus (Dyson
1989).
EXPLANATIONS FOR SEXUALLY DIMORPHIC

TRAITS
Sexual selection is the pre-eminent explanation used for sexually

dimorphic traits (Andersson 1994). Scant attention has been paid to
alternative explanations (Hedrick & Temeles 1989), despite at least
one book purporting to investigate them (Bradbuty & Andersson
1987). The following paragraphs briefly outline three alternative
explanations for sexually dimorphic traits. The major ideas
associated with sexual selection are then summarised in more-detail.
Artefacts of Growth and Survivorship
Sexual differences in growth and survivorship may offer a proximat_e

explanation for differences in body size between males and females
(Shine 1990; Stamps 1993). This is especially true in organisms
6
which continue to grow after reaching maturity. Prolonged growth

following maturity means that differences in adult survivorship may
have a profound effect on the size distribution of each sex. If
survivorship has a small or inconsistent relationship with fitness,
differences in survivorship, and thus body size, may not be a direct
result of selection.
Similarly, if growth is highly plastic and shows low heritability,

sexual differences in body size may occur in a population due to
local environmental conditions. Thus sexual size dimorphism need
not reflect past selection on body size. Nor need it reflect a genetic
response to current selection on body size. It may simply be a
physiological response to an environmental constraint on growth or a
trade-off between life-history traits (Stamps 1993 ).
Natural Selection
Selection due to differential mortality may favour different

phenotypic optima in each sex. This may occur in at least three
different ways. Firstly, the use of different food types by the two
sexes (niche-divergence) may lead to differences in the morphology
of trophic structures (bill or jaws) (Selander 1966; Schoener 1967),
or differences in body size (Hedrick & Temeles 1989; Shine 1989).
These morphological differences may become adaptive by reducing
intraspecific competition for food resources (Slatkin 1984).
7
Secondly, energetic differences between males and females may lead

to different body sizes in each sex (Schluter & Smith 1986; Sandell
1989). For example, large body size in one sex may be attributed to
combat for feeding territories. Small body size may be attributed to
situations in which one sex does most of the hunting, and in which
prey capture is enhanced by agility (e.g. raptorial birds: Andersson &
Norberg 198l;see also Ghiselin 1974 and Vollrath & Parker 1992).
Finally, colouration may be a response to predation pressure. The

dull colouration of female birds may be a response to the increased
risk of predation when they are incubating eggs (Wallace 1891). On
the other hand, the bright colouration of males may be aposematic,
informing predators that these potential prey are poisonous ( e.g.
Dumbacher, Beehler, Spande, Garraffo & Daly 1992) or otherwise
unprofitable (Baker & Parker 1979; Gotmark 1992).
Fecundity selection
Where fecundity shows a relationship with female body size,

fecundity selection may act to increase or decrease female body size.
In some birds fecundity is maximised by females who mature and
breed early. Thus females who invest in reproduction, rather than

growth (ie. are smaller) may be favoured by fecundity selection
(Price 1984; Schluter & Smith 1986. In many other organisms
8
fecundity selection favours large female size because clutch size is

positively related to female body size (Hedrick & Temeles 1989;
Shine 1988).
Sexual Selection
Sexual selection, the mechanism most frequently used to explain

sexual dimorphism, arises from differential mating success among
individuals in a population (Darwin 1871; Arnold 1983). s.exual
selection may work in two ways: (1) by favouring traits in one sex
which attract the other sex (inter sexual selection or mate choice), or
(2) by favouring the ability among members of one sex to compete
with each other for mates (intrasexual selection or competition for
mates) (Darwin 1871).
Intersexual Selection
Mate choice, in its broadest context, occurs when members of one

sex are more likely to mate with certain members of the opposite sex
than others (Kirkpatrick & Ryan 1991; Halliday 1992). However, this
definition ignores the important distinction between passive
attraction and active mate preference (Parker 1983). Passive
attraction occurs where males are chosen more often because they
9
are more easily detected by females. Active mate preference occurs

where males are chosen after several have been detected and
inspected. The need to attract a mate requires that an effective signal
(behavioural, morphological, acoustic or olfactory) be. present to
allow mate recognition. Such signals may be relatively simple, and
need not communicate a great deal of information. On the other
hand, active mate preference relies upon signals that may
communicate complicated information about the 'quality' of a
potential mate. The complexity of active mate choice suggests that
there must be some adaptive advantage, for the chooser, in making a
choice (Parker 1983).
There is some evidence that the two forms of mate choice may act on
'
different traits in some organisms (e.g. Gibson 1996). Indeed mate
choice may be a two stage process, involving initial passive
attraction to a location with ,one .or more males, and subsequent
active preference by the female among those males. Much 'copying'
behaviour reported in female mate choice studies may be the result of
a tendency for females to aggregate around a male with the most
effective distance signal, rather than imitation of the mating
decisions of other females (McComb & Clutton-Brock 1994). The
simultaneous action of passive attraction and active mate preference
may explain why many species have multiple ornaments or signals
(Mfller & Pomiankowski.1993; Gibson 1996).
Female mate choice (in the broad sense) has been demonstrated to
occur in a variety of species (Searcy 1982; Bateson 1983; Thornhill
10
& Alcock 1983; Endler 1986). Empirical research on mate choice has
been largely concerned with identifying traits that distinguish
preferred from rejected males. The relative importance of passive
attraction and active choice in sexual selection is not clear for many
groups of organisms.
A primary question in sexual selection studies is whether the choosy

sex benefits from being choosy? (Maynard Smith 1987; Borgia 1987).
The advantages of passive mate attraction are clear enough - both
partners gain an appropriate, conspecific mate. The advantages of
active mate choice are somewhat more complex. Active mate choice
for a trait may be arbitrary, and carry no clear adaptive advantage
(Fisher 1930, 1958). Alternatively, the chosen trait may carry some
advantage to the choosy sex (Zahavi 1975, 1977; Hamilton & Zuk
1982). Active mate choice may offer short-term or long-term
benefits. A choosy female may benefit in the short-term if mating
with a particular male increases her fecundity, her chances of
survival, or the survival of her young. This may occur if males
provision the female (e.g. Thornhill 1980), or care for her offspring
(e.g. Nisbet 1973). She may benefit in the long-term if she gains
some genetic advantage for her offspring. There is a huge literature
on the idea that females choose mates for their genetic quality. This
literature is ably reviewed by Andersson (1994) and will not be
discussed here.
Many cases of mate choice involve female discrimination among

males on the basis of resources offered to the female or her offspring
11
(Thornhill 1976, 1980; Rutowski 1982; Petrie 1983a; Gwynne 1984;

Alatalo, Lundberg & Glynn 1986). Other studies have shown that
females discriminate strictly on the basis of male traits (Searcy 1982;
Arnold 1983 ). Male traits that have been shown to be subject to mate
choice include visual cues (Andersson 1982; Sigmund 1983; Houde
1987), auditory cues (Gibson & Bradbury 1985; Ryan 1985; Hedrick
1986; Loffredo & Borgia 1986), and olfactory cues (Huck & Banks
1982; Moore & Breed 1986; Shapiro & Dewsbury 1986; Moore
1988).
Once mate choice occurs in a species, any gene causing a preference

for a trait in a female will be passed to her offspring. Any heritable
basis for the male's mating advantage will also be passed onto his
offspring, resulting in a positive feedback between preference and
attractiveness. This may give rise to a genetic covariance between
the chosen male trait and female preference for the trait and result in
a runaway process (Fisher 1930, 1958; Lande 1981). This process
may lead to extreme enhancement of male ornaments by sexual
selection, until opposing natural selection pressures act against
further enhancement of the trait (Fisher 1930).
An intriguing idea put forward recently is that once female choice

has evolved, it may be taken advantage ofby males (Ryan 1990;
Ryan et al. 1990; Ryan & Rand 1990). Male traits may evolve
through 'sensory exploitation' by the males of pre-existing female
preferences. Like Fisher's runaway process, the sensory exploitation
hypothesis offers a mechanism by which extreme male ornaments may
12
evolve under the influence of mate choice. But neither explains why
female choice might evolve initially. There is considerabl~ empiricaI
evidence for Fisher's runaway process (Andersson 1994), but there is
little support for the sensory exploitation hypothesis.
lntrasexual selection
Intrasexual selection refers to variance in mating success resulting

from fighting and other forms of combat over access to mates. Traits
which allow an individual to win a fight for access to mates, or which
minimise the costs of fighting for access to mates may be favoured by
intrasexual selection. These traits might include weapons, which
confer an advantage by allowing individuals to win contests and gain
access (directly or indirectly) to females. The antlers of red deer
(Cervus elephas) (Clutton-Brock, Guiness & Albon 1982), or the
thumb spur.s of gladiator frogs (Hyla rosenbergi) (Kluge 1981) are
examples of weapons.
Traits which minimis(Jthe costs of fighting may confer an advantage

on contestants by allowing them to avoid unnecessary escalation of
risky contests. This may be achieved by badges of status in situations
where individuals seldom meet more than once (Rohwer 1982). A
badge of status usually takes the form of a colour patch, but may be
just body size. It provides information to potential competitors about
the. fighting ability of the individual 'wearing' the badge. This
13
information would permit individuals to avoid many potentially

costly fights. Alternatively, traits which allow individual recognition
may be advantageous where individuals meet repeatedly. The variable
head pattern of Turnstones (Arenaria interpres) permits territorial
neighbours to recognise each other (Whitfield 1986), and thus avoid
unnecessary costly fights.
Summary of explanations for sexually dimorphic traits
Most sexually dimorphic traits could be explained by several of the

hypotheses outlined above. It is important not to assume that sexual
selection is a universal explanation for sexually dimorphic traits.
Many of the explanations for sexually dimorphic traits only refer to
sexual differences in body size (e.g. fecundity selection, proximate
effects of growth and survival), and do not address other dimorphic
traits. Other explanations may explain sexually dimorphic colouration
( e.g. the unprofitable prey hypothesis, sexual selection), or
differences in body shape ( e.g. energetic differences, sexual
selection).
14
SEXUAL DIMORPHISM AND SQUAMATE REPTILES
The squamate reptiles are divided into two major lineages (Estes, de
Queiroz & Gauthier 1988) (Figure 1.1). The Iguania are primarily
visually oriented, sit-and-wait insectivores, in which males usually
defend most or all of their home-ranges. This lineage includes the
iguanid lizards of the Americas and the Chamaeleons and Dragon
lizards of the Old World. The other lineage, the Scleroglossa, are
primarily olfactorily oriented, active foragers, in which males do not
normally defend territories. This lineage includes all non-iguanian
lizards, and snakes.
Sexual dimorphism is widespread, and takes many forms in both

lineages of squamate reptiles. Many species show sexual differences
in body size (Fiteh 1981). The degree of sexual size dimorphism
varies from a Cuban population of the Teiid lizard Amieva auberi, in
which the body length of males is 59% that of females (Schwartz
1970), through to the iguanian lizard Ano/is ferreus, in which the
body length of males is 183% that of females (Roughgarden 1995).
Sexual differences in body shape are also widespread among

squamates. These may include relatively subtle differences in the
lengths of appendages and head size (Cooper & Vitt 1989), through
to extreme development of weapons, such as the horns of males of
some Chamaeleons (Darwin 1871). In addition to these fixed sexual
differences in shape, some species are able to affect temporary
15
differences in body shape. Males of many iguanians have distinctively

patterned, extendible dewlaps on their throats, which they display
during courtship and territorial disputes (Noble & Bradley 1933;
Smith 193 5; Mori & Hikida 1994 ). Females have smalle; less
distinctive dewlaps. The patagia of flying dragons (Draco spp.) differ
between the sexes in size and colour, and are extended to affect
temporary shape changes during social encounters (Smith 1935; Mori
& Hikida 1994).
The sexes differ in colour in many squamates. In snakes these

differences are generally subtle (Shine & Madsen 1994), but can be
pronounced among lizards (Cooper & Greenberg 1992). Males are
usually the brighter coloured sex among lizards. The brightest
colours are usually restricted to the ventral or ventro-lateral
surfaces, or extensible skin folds ( e.g. dewlaps). Presumably this
allows the use of colour as a social signal while avoiding costs of
predation from dorsal patterns (Gibbons & Lillywhite 1981).
Selection for dimorphic traits in squamates
As with other groups of animals, there has been controversy over the
relative roles of sexual selection versus other explanations for
sexually dimorphic traits in reptiles. The following is a brief review
of the major findings with respect to the alternative explanations for
dimorphic traits, and sexual selection as they relate to the Squamata.
16
Chapter l . INTRODUCTION
Alternatives to Sexual Selection
Artefacts of growth & survivorship: Squamate reptiles show

indeterminate growth, much of which occurs after they reach
maturity (Andrews 1982). Within species, individuals show
considerable variation in growth rate and in their final (asymptotic)
size (Halliday & Verrell 1988), This variation in growth pattern
suggests that environmental constraints on growth, and sexual
differences in survivorship may act as proximate determinants of
sexual size dimorphism (SSD) (Stamps 1993). Sexual size
dimorphism may appear larger if young males suffer higher mortality
than young females, leaving mostly older larger males in a population
(Stamps, 1995). Higher mortality among older, larger females than
among older males may similarly increase apparent SSD (Watkins
1996). Patterns of SSD due to proximate factors may not be
temporally stable, and might be thought of as resulting from sampling
error. Variation in food availability may effect the growth rates of
·· males and females differently, and these different responses may
explain spatial and temporal variation in SSD among populations
(Andrews & Stamps 1994; Stamps, Krishnan & Andrews 1994;
Forsman 1991; Madsen & Shine 1993). Few data are available on the
heritability of growth parameters in reptiles (cf Baverstock 1972). So
it is not presently possible to determine whether selection acting on
growth pattern is likely to give rise to an evolutionary response, or
to remain a proximate source of variation in body size.
17
Natural selection: Niche-divergence and consistent patterns of

differential mortality due to predation have been invoked to explain
sexually dimorphic traits in squamates. Sexual differences in
morphology are accompanied by differences in ecology in several
species of lizards and snakes (Shine & Lambeck 1985; Shine 1986,
1989; Houston & Shine 1993; Perez-Mellado & De La Riva 1993)
leading to the suggestion that the dimorphic traits may have evolved
through niche-divergence.
Consistent patterns of differential mortality between males and

females is widespread among squamate reptiles (e.g. Dunham 1981;
Watkins 1996). These differences may occur due to different costs of
reproduction, or differential exposure to predation, and may offer an
ultimate explanation for behavioural and morphological differences
between the sexes. Generally the brighter markings of males cause·
them to be more prone to predators, but there are some exceptions.
For example, male adders (Vipera berus) are more brightly patterned
than females (Prestt 1971 ). The distinctive colour pattern of males
may (paradoxically) make them less visible to predators due to a
flicker-fusion effect when they are moving in search of mates (Shine
& Madsen 1994). There is some evidence that brightly coloured male
Lacerta agilis may be advertising to predators that they are

unprofitable prey, and therefore suffer lower mortality due to
predation than females (Olsson 1993a).
Fecundity selection: A positive relationship between fecundity and

body size among female squamates is commonplace (Shine 1988b ).
18
The logic that such a fecundity advantage may result in selection for
larger body size is compelling, but the idea has remained difficult to
test (Shine 1988b ). This may be because fecundity does not refer
only to, the number of offspring produced, but also to offspring size.
A trade-off between offspring number and offspring size (Stearns
1992) may confound any evolutionary relationship between clutch
size and maternal body size.
Sexual selection
Lizards have been of historical importance in the study of sexual

selection. Research done during the 1930's and 1940's on lizards
(Noble & Bradley 1933; Noble 1934; Greenberg & Noble 1944) was
among the first empirical research on sexual selection. These studies
stand out against a dearth of research on sexual selection prior to the
1970's (Campbell 1972; Andersson 1994).
Despite the long history, a recent review (Andersson 1994) has

shown that few reptiles have been well studied with respect to sexual
selection. Of the 3000 species of squamate reptiles, Andersson could
find published evidence for sexual selection acting on specific traits
for only five species. Since Andersson's review, several studies have
identified traits which are subject to sexual selection in other lizard
species. These traits (from Andersson's review and later studies)
include body size (Ruby 1981; Andren 1986; Madsen 1988; Hews
19
1990), head size (Hews 1990; Cooper & Vitt 1993; Bull & Pamula
1996), colouration (Noble 1934; Crews 1975; Sigmund 1983;
Ellingt~n 1993), display frequency (Greenberg & Noble 1944; Ruby
1981), and territory size (Trivers 1976). The paucity of sexual
selection studies on reptiles is surprising, given the widespread
differences in body size, shape and colouration (Stamps 1983), and
the importance of lizards to modern ecological studies (Huey,
()f
Schoener & Pianka 1983). This may reflect the difficult~observing
lizard behaviour relative to other taxa.
The relative importance of intra sexual selection and 'inter sexual

selection, is a point of contr-oversy in the Squamate reptiles, as in
other organisms (Bradbury & Davies 1987; Halliday 1992). There is
considerable anecdotal and comparative evidence that male-male
combat is important among squamate reptiles, and various traits have
been shown to be advantageous in this context (e.g. body size: Shine
1978, 1994; Carothers 1984; Tokarz, 1985; Olsson 1992, Shine &
Fitzgerald 1995; colouration: Werner 1978, Thompson & Moore
1991, Olsson 1994a,b; sprint speed: Garland, Hankins & Huey 1990).
However, few of these traits have been shown to confer an advantage
in terms of mating success, and therefore, to be subject to sexual
selection.
Two recent reviews have disagreed as to the prevalence of female

mate choice in lizards. On the one hand, Olsson & Madsen (1995)
have argued that mate choice is rare. On the other hand, Tokarz
(1995) argues that mate choice. is commonplace among lizards. This
20
apparent variance of opinion may be due to the use of different

criteria for the presence of mate choice. Tokarz (1995) expressly
regards any case in which females are more likely to mate with
certain males than others as evidence for mate choice, following
Kirkpatrick & Ryan (1991) and Halliday (1992). Thus he includes
instances of both passive attraction, and active preference (sensu
Parker 1983) in his review. Olsson & Madsen (1995) use a more
exclusive definition, which requires females to show active mate
preference among conspecific males. Taking both reviews together,
there is considerable evidence for passive mate attraction among,
lizards (see Tokarz 1995), but little evidence for active mate
preference among them (Olsson & Madsen 1995). This is, perhaps,
not surprising because male squamates have never been observed to
feed females, or to exhibit parental behaviour (Shine 1988a).
Although females could still gain advantages for their offspring by
choosing males with 'better' genes.
Active female mate preference has been convincingly shown in only

two species of lizard. Female Broad-headed Skinks (Eumeces
/aticeps), when given a choice, prefer males with larger heads, but
do not distinguish between malison the basis of head colouration
(Cooper & Vitt 1993). Female geckoes (Gonatodes albogularis)
when given a choice between similary sized males, prefer males with
brightly coloured heads (Ellington 1993). Active male mate choice
has also been shown in lizards. Males may show an active preference
for unfamiliar (Tokarz 1992), or large, more fecund females (Olsson
1993c). There is intriguing evidence that Sleepy Lizards, Tiliqua
21
rugosa, mate with the same partners for up to eight successive years
(Bull 1988, 1994), but there is no experimental evidence to show
that this is a result of active mate preference by either sex. The lack
of evidence for active mate preference in squamate reptiles contrasts
with the situation in many other taxa in which female mate choice has
been reported frequently (Andersson 1994). It is not clear whether
this is a real difference. It may simply rer'1ect
,/
the difficulty of
observing courtship and mating activities in lizards.
Squamate mating systems vary from monogamy (Tinkle 1967; Jenssen

1970; Bull 1988, 1994), through polygyny (Blair 1960; Harris 1964;
Rand 1967; Manzur & Fuentes 1979; Ruby 1981; Hews 1990;
Wikelksi, Carbone & Trillmich 1996), to promiscuity (Madsen et al.
1992; Olsson et al. 1994). The temporal and spatial distribution of
receptive females is a primary determinant of the mating system and
the intensity of sexual selection in lizards (Stamps 1983), as in many
other taxa (Emlen & Oring 1977).
Female squamates appear to distribute themselves with respect to

resources, rather than males (Manzur & Fuentes 1979; McCluskey et.
al l 990a,b; Hews 1993). The disappearance of a male and his
replacement by another male rarely leads to females moving from
their home ranges (Berry 1974; Madsen & Loman 1987; Hews 1993;
but see Harris 1964). Consequently, reproductive success among
male squamates depends largely on (1) the number of females living
in a male's home range, (2) the probability of fathering those females'
offspring, and (3) the fecundity of each female (Stamps 1983). Male
22
lizards may increase their reproductive success by arranging their .

home ranges to overlap with female home ranges, or by increasing
the size of their home range to encompass more than one female
(Stamps 1983). They may also defend their home ranges
(territoriality), or guard females, so that they ensure exclusive
breeding rights to females (Stamps 1983). Territoriality appears to
be ancestral to all squamate reptiles, and appears to have been lost a
number of times in the non-iguanian squamates (Martins 1994).
The probability of paternity is commonly assumed to be high in

territorial species (Berty 1974; Trivers 1976; Ruby 1981), even
though DNA fingerprinting has shown similar assumptions in birds to
be unfounded (eg. Westneat 1987; Sherman & Morton 1988). If
tei:ritoriality secured a male's paternity, then the numbe.r of females
in a male's home range multiplied by each female's fecundity should
give a reasonable estimate of a male's reproductive success (Stamps
1983). However, males of many non-territorial species overlap their
home ranges. Even in territorial species, 'sneaker' males may
sequester themselves within the home ranges of territorial males. In
these cases a male is less certain of fathering all of the offspring of a
female with whom he has mated. Males of some non territorial
scincids, lacertids and teiids (Carpenter & Ferguson 1977; Olsson
1993b), and even some territorial iguanians (Ruby 1981) 'tend'
females. This mate guarding behaviour may be another method by
which males increase the probability of paternity.
23
Multiple mating by female squamates may promote sperm

competition, and increase a female's fitness (Smith 1984; Madsen et
al. 1992). Females who mate with multiple males gave birth to more
viable offspring than those who mated with fewer male partners
(Madsen et al. 1992; Olsson et al. 1994).
/
'
THE STUDY ORGANISM
The Peninsula Dragon ( Ctenophorus fionni) is a member of the C.

decresii species group (Houston 1974) and is endemic to the Eyre
Peninsula in South Australia (Figure 1.2). All members of this group
are medium-sized(::,; 96 mm snout-vent length) diurnal, rock dwelling
lizards with dorsoventrally depressed bodies. Males are much larger
than females, have larger heads, and are more colourful than females
(Houston 1974, 1978). Members of this group are commonly referred
to a Rock Dragons. It is one of several of informally recognised
species groups (Houston 1974, 1978; Storr, Smith & Johnstone
1983) within the Australian genus Ctenophorus. Australian Dragon
Lizards are a distinctive clade within the Agamidae (Moody 1980).
Within the lguania (Figure 1.1 ), the Agamidae ( dragon lizards) and
the Chamaeleontidae are recognised as the sister lineage (the
Acrodonta) to the Iguanidae (Estes, de Queiroz & Gauthier 1988;
Frost & Etheridge 1989). This is contrary to a recent proposal that
the Iguanidae are paraphyletic, including the Agamidae-
24
Chamaeleontidae clade, and which divided the Iguanidae into several

families and regarded the Agamidae as a subfamily (the Agaminae) of
the Chamaeleontidae (Frost & Etheridge 1989). I have not adopted
Frost & Etheridge's (1989) proposal because: (1) their study was
based mostly on morphological variation in Iguanids, and did not
take full account of the variation shown by Agamids and
Chamaeleontids, and (2) recent unpublished molecular data suggest
that the Agamidae warrant familial status (Hutchinson pers. com.
1996). In this thesis I use the term lguania to refer to the Agamidae,
Chamaeleontidae, and Iguanidae collectively. The term Iguanidae
refers to all new world iguanians (i.e. Crotaphytidae, Iguanidae,
Phrynosomatidae, and Tropiduridae of Frost and Etheridge 1989).
The Peninsula Dragon Lizard (Ctenophorus fionni) was described

from a single female collected from Port Lincoln in South Australia
by Frederick Wood-Jones (Proctor 1923). Since then this species has
received scant attention in the literature. A study on the water
physiology of C. fionni from Wintinna in northern South Australia
(Warburg 1966) was, in fact, based on the closely related, but then
undescribed, C. tjantjalka (Johnston 1992). Several gut parasites
have been described from. C. fionni, including Maxvachonia hrygooi
for which C. fionni (albeit misidentified as C. decresii) is the type
host (Mawson 1972).
The Peninsula Dragon occurs in a series of more or less isolated

populations that are restricted to suitable rock outcrops. There is
considerable variation among these populations in male colour and
25
body size (Houston 1974, 1978), but females are more dully coloured
than males and do not show the same degree of geographic variation
in colour (Houston 1974) .. Some of the variation in colour between
populations appears to be related to substrate colour matching
(Gibbons & Lillywhite 1981). Male Peninsula Dragons defend
territories with an unusual hind limb pushup display (Gibbons 1979).
The evolution of sexually dimorphic traits in the Peninsula Dragon is

interesting for a number of reasons. First, there have been no
detailed studies of the evolution of sexual dimorphic traits in any
Agamid, and such a stud·y would provide an important comparison
with the closely related Iguanids, and other squamates. Secondly, a
study of the Peninsula Dragon may contribute to the long-running
debate over the relative importance of various sources of selection
(mortality, sexual and fecundity) in the evolution of sexually
,dimorphic traits. Finally, such a study would contribute to the debate
over the relative importance of intra· and intersexual selection in
sexual selection.
STUDY SITES
The major part of this study was conducted at Midgee Rocks (33
25'8, 137 OS'E) (Figure 1.2 &1.3). This site consists ofa series of
isolated outcrops of the middle proterozoic (1450-1500 Ma BP)
coarse-grained Charleston Granite (Parker, Fanning & Flint 1985) at
26
an altitude of 120 m. The outcrops are separated by longitudinal

dunes of siliceous sand (the Moornaba Sand) which were deposited
between 16 and 25 thousand years BP (Ludbrook 1980). The granite
outcrops vary in area from less than one hectare to six hectares. My
main study site was an outcrop with an area of 3. 5 hectares (Figure
1.3).
There are three main types of vegetation at Midgee Rocks. The

deeper sands near the crests of the dunes support a mixed Eucalyptus
incrassata-E. socialis low mallee, with emergent Hakea
francissiana. The shallower sands surrounding the granite outcrops
support a Melaleuca uncinnata closed shrubland. The outcrops
themselves support an Acacia rigiophylla shrubland. All three
vegetation types have a dense understorey of Spinifex (Triodia
irritans). The area was last burnt in the early 1970's (E. Carmody,
personal communication, 1991). The blackened stumps of native
pines (Callitris collumellaris) are a prominent feature around the
bases of the outcrops, suggesting that the area was once open
woodland.
Lizards for this study were also collected from three other locations.
Mount Young (33 06'S, 137 29'E; Figure 1.2) is a large (30 ha.)
isolated, steep-sided quartzite outcrop of the Proterozoic Moonabie
Formation (Parker et al. 1985). It rises from sea level to an altitude
of 136 m and has an Allocasuarina stricta sparse woodland on the
crest. A dense hummock grassland of Triodia irritans covers the
slopes. The base of Mount Young is surrounded by Holocene mud
27
flats inhabited by a low shrubland of Halosarcia spp. These mud flats

are unsuitable for Peninsula Dragons.
Siam Station (32 33'S, 136 43'E; Figure 1.2) is located near the
eastern margin of the Gawler Ranges. It is an area of rolling hills of
outcropping proterozoic Yardea Dacite interspersed with Holocene
dunes (Parker et al. 1985). The entire area is below 200 min
altitude. Many of the rounded outcrops are bare, but areas of skeletal
soils support an open shrubland of Acacia tarcoolensis, with
localised dense patches of Triodia irritans hummock grassland.
Iron Duke (33 !S'S, 137 07'E; Figure 1.2) is the southern point of
the Middleback Ranges. This range is an extensive (many square
kilometres) area of outcropping iron stone of the early proterozoic
Middleback Gronp (Parker et al. 1985). The vegetation in the
Middleback Ranges is complex, and varies from Eucalyptus oleosa-E.
brachycalyx dense mallee, through Eremophila-Beyeria-Alyxia
shrubland, to open Triodia irritans hummock grassland. The
environment and herpetofauna of the Middleback Ranges have be·en
described in some detail elsewhere (Johnston 1982).
All four of these localities are within one hundred kilometres of

Whyalla (Figure 1.2), and share a similar climate. Whyalla has a
Mediterranean climate with a mean annual rainfall of 270 mm, twenty
percent of which falls in the austral summer (Figure 1.4). Records
taken at Whyalla provide one of the longest continuous records of
rainfall and daily temperatures in the area.
28
APPROACH & ORGANISATION OF THIS THESIS
In this thesis I have been deliberately didactic in my approach and

have attempted tests between exclusive alternative explanations
(Platt 1964) for the evolution of a particular trait wherever possible.
However, where it is apparent that multiple processes may be
working simultaneously, I have attempted to determine the relative
contributions (Quinn &, Dunham 1983) of these processes.
This chapter has outlined the major current hypotheses explaining the
origins of sexually dimorphic traits, summarised the available
information on the Peninsula Dragon, and described my study sites.
My intention has been to provide the necessary background to the
rest of this .thesis. In the rest of this thesis I test each of the major
ideas with respect to sexually dimorphic traits in the Peninsula
Dragon lizard, Ctenophorus fionni.
Chapter two describes the sexual dimorphism in Peninsula Dragons.

This description is based on ideas derived from the study of
heterochrony, relative timing and rates of developmental processes.
This approach was chosen because it has the potential to identify
differences in ontogeny which may have given rise to sexually
dimorphic traits (e.g. Hews 1996). This information may also suggest'
the evolutionary mechanisms which may have given rise to dimorphic
traits.
29
Chapter three outlines the reproductive biology of Peninsula

Dragons. This has not been described before, and provides a basis for
the design of several experiments reported' in later chapters. Data on
the relationship between fecundity and female size are also used to
assess the relevance of fecundity selection in the evolution of female
size in Peninsula Dragons.
Chapter four tests the idea that differential mortality between males
and females may serve as a proximate mechanism giving rise to
differences in the size distribution between the sexes. The chapter
tests this hypothesis by describing patterns of growth and
survivorship in male and female Peninsula Dragons.
Chapter five assesses the hypothesis that ecological differences

between the sexes may have given rise to the morphological
differences between them. The chapter is divided into four sections,
each of which deals with one of four ecological niche axes thought to
be important for reptiles (food, space, temperature and time; Pianka
1973). My approach has been to determine whether there is a
difference between the sexes in their realised niches in the field.
Then, if a difference was found, I determine whether there was a
difference in fundamental niche. A difference between the sexes in
the fundamental niche was taken as evidence that natural selection
may have favoured niche-divergence along that particular niche axis.
Correlations between resource use and morphology were then used to
test hypotheses about the relationship between the ecological and
morphological differences between the sexes. If no difference was
30
found between the fundamental niches of the sexes, but the realised
niches differed, this was taken as evidence that the ecological
differences between the sexes were the result (rather than the cause)
of morphological differences between the sexes. In these cases the
pattern of resource use relative to seasonal changes in reproductive
state and territorial behaviour, or responses to experimental
manipulations of the social environment were used to examine the
basis for the ecological differences between male and females.
Chapters six, seven and eight collectively examine the hypothesis

that sexual selection has given rise to the sexually dimorphic traits in
Peninsula Dragons. Chapter six describes the mating system in a
population of Peninsula Dragons and estimates the potential for
sexual selection in this population. Chapter seven identifies the
morphometric traits which may be subject to sexual selection. It also
examines whether intrasexual selection is adequate to explain the
patterns of sexual selection, or whether intersexual selection (mate
choice) must also be invoked as an explanation. In Chapter eight I
report an experimental manipulation of the colour pattern in male
Peninsula Dragons in the field. This experiment was designed to test
whether mate choice or male competition explain the sexual
differences in colouration.
Thus chapters two to five describe the sexual dimorphism in

Peninsula Dragons, provide necessary background information, and
test alternative hypotheses regarding the origin of the dimorphism.
Chapters six to eight test the hypothesis that sexual selection has
31
given rise to the dimorphism. Chapter nine provides a synthesis of

the results in terms of the various hypotheses on the origin of
sexually dimorphic traits, and suggests future lines of research.
32
:!l ~
=
:-g
::, "'
§ Ql
ctl
~ Ql 32
c:
Ql
E Iiiiii
Ql
X :::t: >
Scinooklea
u
Varanoidea
Gekkota
Anguimorpha
lguania · I I I
I I I Scincomorpha
I I I
I I I
I I I
Scleroglossa
Squamata
I
FIGURE 1.1: Phylogeny of the Squamate reptiles (After Estes et al.

1988). The Dragon Lizards (Agamidae) are asterisked.
33
134° 136°
l•
!
II .
I
i
ii
•
• f.';m Station * •
I
~
• •
• Iron Duke•
p WHYALLA
• Mount Young
• •
Mldgee Rocl<, '*
KILOMETRES
100 200
•
0
FIGURE 1.2: The distribution of the Peninsula Dragon lizard

(Ctenophorus fionni). Each dot represents a locality record based the
Australian Museum and the South Australian Museum collections.
The asterisks represent sites which were visited as part of this study.
34
FIGURE 1.3 : Photograph of the main study site at Midgee Rocks ,

South Australia (see Figure 1.2).
35
30
,.,
t:,.j
25
30
20 !~
--e=.. ,.,
-
25 15
~
-J
...:i 10
--
-
20
<
r..
0
(""j
-<
z
c:i:::
15
5
10
J FMAMJJA SOND
MONTH
FIGURE 1.4: Climate in Whyalla, South Australia (see Figure 1.2).

The histogram shows mean monthly rainfall (mm). Triangles
represent mean monthly minimum (T) and maximum(&) temperatures
("C).
36
CHAPTER2
SEXUAL DIMORPHISM
Sexual dimorphism is widespread among vertebrates; taking the form of sexual

differences in body size, body shape and colouration (Darwin 1871; Andersson
1994). These dimorphisms may arise from a variety of proximate and ultimate
causes. For example, differences in body size may arise from non-heritable
differences in survivorship between the sexes (Dunham 1981). Ahematively,
body shape may be modified by sexual selection to produce weapons used in
contests or ornaments to attract mates (Darwin 1871; Andersson 1994). Body
size and shape may also be modified by fecundity selection arising from a
fecundity advantage associated with large female size (Darwin 1871; Shine
1988b). Different energetic or ecological requirements of males and females
may also give rise to, or at least amplify, morphological differences between
the sexes (Darwin 1871; Selander 1966; Slatkin 1984).
The form of a particular sexual dimorphism may suggest which one, or

combination, of these mechanisms applies. For instance short-term temporal
Chapter2 SEXUAL DIMORPHISM
variation in sexual size dimorphism may indicate that an apparent dimorphism

results from differences in survival between the sexes, and is an artefact of
population structure (Dunham 1981). The presence of weapons in one sex
implies that intrasexual selection may be important (eg. Silverman & Dunbar
1980) whereas the presence of ornaments may suggest the action of mate
choice (eg. Andersson 1982). Sexual differences in jaw structure may arise
through intraspecific niche partitioning, but equally might arise through
intrasexual selection ifjaws are used as weapons in contests (Shine 1989) or
intersexual selection ifjaw shape influences mate choice.
The form of sexually dimorphic traits in adults must also be understood in

tenns of ontogeny. Shape usually changes in response to changes in absolute
body size, as occurs during growth. The study of changes in shape, or the
relative size of traits, is referred to as allometry. The pattern of trait
allometries may also suggest the manner by which the dimorphism is produced
ontogenetically (Vitt & Cooper 1985; Livezay & Humphries 1986; Patton &
Brylsltl 1987; McNamara 1995; Hews 1996). An understanding of ontogenetic
processes may, in turn, lead to inferences about the form of selection acting on
each sex (Schoener et al. 1982; Vitt 1983; Carothers '1984; Vitt & Cooper
1985; Berglund et al. 1986; McGillivray & Johnston 1987; Zeh 1987; Arak
1988; Cooper & Vitt 1989; Vial & Stewart 1989) and whether selection is
acting on immature or adult stages (Shine 1990). Thus allometric differences
between sexes within a species can be critical in both ultimate and proximate
analyses of the evolution of sexual dimorphism.
The flightless steamer ducks (Trachyeres spp.) of South America provide a

good example. Flightlessness in birds has generally been explained as a
response to reduced predation in species living on islands which lack predators
38
Chapt~r 2 SEXUAL DIMORPHISM
(eg. Lowe 1928). However, an allometric analysis of a mixed population of

flighted and flightless Trachyeres patachonicus showed that flightlessness in
this group results from an increase in body size with no commensurate
increase in wing area of flightless individuals, rather than through a reduction
in wing size, as would be predicted by the reduced predation hypothesis
(Livezay & Humphrey 1986). Increased body size in flightless steamer ducks
is advantageous in territorial defense of food resources and young (Livezay &
Humphrey 1985). Flightlessness in steamer ducks appears not to be related to
reduced predation pressure, but instead seems to be a result of selection for
larger· body size, a conclusion based on an allometric analysis, which could not
so easily have been arrived at by· other means.
Although a general description of sexual dimoi:phism in Ctenophorus ftonni is

available (Houston 1974; see Figures 2.1 & 2.2), there are no integrated
accounts of the extent of sexual dimoi:phism in body .size, shape and colour for
this species. Indeed, theff! are few detailed descriptions of sexual dimoi:phism
in any lizards (Vitt & Cooper 1985; Olsson 1992; Hews 1996), despite their
historical importance in the study of sexual dimoi:phism (eg. Schoener 1967;
Trivers 1976). This chapter gives a detailed description of sexual dimoi:phism
in the Peninsula Dragon lizard, paying particular attention to the ontogenetic
processes which give rise to it. This accota1t provides a basis for the rest of
this thesis, which explores the alternative explanations for sexual dimoi:phism
in C. ftonni and other lizards.
39
METHODS
Data were collected from lizards during a four year study (1991-1994) of
Ctenophorosfionni at Midgee Rocks on Eyre Peninsula (Figure 1.2 & 1.3).
Li2:ards were sexed by probing (Schaefer 1934), or by everting the hemipenes
with gentle pressure at the base of the tail Ihards were classed as adults if
their SVL was greater than 50mm, and as juveniles if their SVL was less than
50mm (see Chapter 3). Sex could not be determined reliably for many
juveniles.
Body Size and Shape
The description of sexual dimorphism in body size and shape was based
entirely on measurements taken from lizards at Midgee Rocks (N = 381).
Individual lizards were captured, measured, permanently marked by toe
clipping (Fenner 1979), and returned to the site of capture within 12 hours.
The measurements (see Figure 2.3) were snout-vent length (SVL), depth of
the head immediately behind the eye (HD), maximum head width across the
tympani (HW), distance from the tip of the snout to the anterior margin ofthe
tympanum (HL), forelimb length (FLL), hindlimb length (HLL), tail length
(TL) and body weight (Mass). All linear measurements were made with
vernier calipers to the nearest 0.1mm. Lizards were weighed to the nearest
0.5g on Salter 50g scales. All measurements were highly repeatable. Not all
traits were measured on all individuals, so sample sizes differ among traits.
40
Chapter 2 SEXUAL DIMORPHISM
, Only measurements taken at the first time each lizard was captured are
reported here.
The null hypothesis that adult males and females did not differ in overall body
shape was tested with a Discriminant Function Analysis using a subset of
lizards fur which all measurements were available ( 128 adult males and 92
adult females). A discriminant function ~a multivariate technique which
maximises between groups variation with respect to within groups variation to
maximise correct classification ofindividuals. among predetermined groups
(Reyment, Blackith & Campbell 1984). Data from all years. were pooled for
this ana]ysis and all measurements were log10-transformed to ensure
multivariate normality. The discriminant function derived from the initial
analysis of adults was then used to classify 116 unsexedjuveniles in a post hoc
analysis. This allowed an overall comparison of the shape ofjuvenile lizards
with the shape of adult males and females. The null hypothes{s that adult
males and adult females did not differ was also tested for each trait
individually with oneway analyses of variance, based on those lizards admitted
to the discriminant function analysis.
Ontogenetic variation was examined by fitting measurements of each trait to

the logarithmic form of the allometric equation Y = bSVL0 (Huxley 1932),
where Y is the value of the trait being examined, SVL is used as a measure of
size, a is the allometric coefficient (slope) and bis they-intercept. Allometric
relationships were calculate<i forjuveniles, adult males and adult females
separately. Initial attempts to analyse juvenile males and females separately
were hampered by the low number ofjuveniles that could be assigned a sex.
Analysis of residuals indicated that the log10-transformed data were normally
distri'buted in each case and among-group variances were homogeneous. An
41
Cbapter2 SEXUAL DIMORPl:IlSM
analysis of covariance (ANCOVA) was performed on log 10-transformed

values for each trait, with log 10SVL as the covariate and sex/age-class as
treatments. These analyses were used to test the null hypothesis that the slopes
did not differ between groups. A significant SVL by sex/age-class interaction
term in the ANCOVA indicates that at least two groups differed in their
allometric coefficients. Where differences were found, post-hoc pairwise tests
were used to determined which slopes differed.
Colour
Qualltative descriptions of colour variation due to sex and age are based on
the lizards studied at Midgee Rocks. When each lizard was first captured its
dorsal colour was scored from one to five with increasing scores indicating
increasing degree ofmelanisation (see Figure 2.4). Colour photographs were
also taken of the dorsal and ventral surfaces of males the first time they were
captured in the spring of each year of the study. All photographs were taken
with an Olympus OM-4ti SLR camera and a Zuiko 50mm macro lens on a
matte black background, at a distance of350mm, with flash at f22 using

Kodachrome-64 slide film. These provided information on the variation in
individual male colour pattern with age, and were used to measure the size of
the black chest patch in 59 males.
Chest patch area (mm2) was estimated by tracing the ventral profile of a lizard
from photographs onto blank paper, and then tracing in the ventral area that
was covered with black pigment (see Figure 2.2). These two profiles were cut
from the sheet of paper and weighed. The weights were then compared to the
42
Chapter 2 SEXUAL DIMORPHlSM
weight of standard squares of paper ofknown area to give an estimate of the

the total area (cm2) of the ventral surface, and the area of the black chest
patch. The area of the chest patch, expressed as a proportion of the total
ventral area, was regressed against the male's SVL to test the null hypothesis
that the chest patch does not increase in relative size with increased body size.
Quantitative measurements of reflectance were taken from three adult males

and three adult females from Mount Young (Figure 1.2) with an Evans
Electroselenium spectrophotometer. This instrument expresses light reflected
from a surface as a proportion of light reflected by a magnesium carbonate
blank (Benford, Lloyd and Scharz 1948). Lizards from Mount Young were
used because specimens from Midgee Rocks were not available when I had
access to the spectrophotometer, and the two populations are very similar in
colour. Reflectance was measured at nine wavelengths over the visible
spectrum; 425µm., 445µm, 465µm, 485µm, 515µm, 545µm, 595µm, 655µm,
and 685µm. All measurements were taken middorsally so as to measure the
general background colour of the body. The measurement port was placed
over the same area of the lizard for measurements at each wavelength to
minimire errors due to changes in the range of angles of incidence of the light
beam on the scales, and due to variation in back pattern. Peninsula Dragons
show little variation in reflectance due to variations in body temperature,
administration of ACTII or mild electric shock (Sharrad 1969; Gibbons &
Lillywhite 1981; personal observations), so a single set ofreflectance
measurements was taken from each individual lizard. None of the lizards used
in this study sloughed their skin during the ten days preceding or succeeding
reflectance measurements. All measurements were taken between 0900h and
I600h during spring when the lizards were active, and had body temperatures
of30-38°C. The spectrophotometer was calb'brated prior to every
43
measurement. A Wilcoxon's signed-ranks tests (T, Sokal and Rohlf 1981)

based on the mean reflectance measurements at each wavelength was used to
test for differences in reflectance between the sexes.
RESULTS
Body Size and Shape
The pooled measurements of all C. flonni caught at Midgee Rocks indicate

that average SVL for males (SVL = 75.9 ± 9.5 mm) was greater than that for
females (SVL = 69.5 ± 7.44 mm) (Figure 2.5). Based on these average SVLs
males were 9% larger than females.
Discriminant function analysis of eight measurements using males and females

as a priori groups resulted.in a significant discriminant function (Wilk's 11. =
0.486, P < 0.01) (Table 2.1) which correctly classified 89.1% of adult lizards.
The same discriminant function classified 97.4% ofunsexed juveniles as
females (Figure 2. 6). Males were significantly larger than females for all eight
measurements (Table 2.2).. Head width and hindlimb length had the highest
correlations with the discriminant function, whereas SVL and Mass had the
lowest. This indicates that the major differences between males and females
were in relative head size and relative length of the appendages. Despite the
significant sexual size dimorphism, shape, rather than overall size was the
major difference between the sexes.
44
A comparison between allometric coefficients for adult male, adult females

and juvenile Peninsula Dragons shows three patterns of variation in allometry
(Figures 2. 7-2.9). First, males and females have a higher allometric coefficient
for Mass than juveniles, and males differ from females only in that they grow
to a larger body size (Figure 2.7, Table 2.3). Secondly, for all three head
measurements, males showed higher allometric coefficients than females and
juveniles, but females showed lower allometric coefficients than juveniles for
two two of the head measurements (Figures 2.8, Table 2.3). Thirdly, for limb
and tail measurements, adults of both sexes showed lower allometric
coefficients than juveniles, but males had higher allometric coefficients than
females (Figures 2.9, Table 2.3). Figures 2.7, 2.8 and 2.9 give one graphical
example of each of these patterns of allometry. Full allometric statistics for all
measurements are given in Table 2.3.
Colour
All juveniles and adult female Peninsula Dragons are very similar in colour and
pattern. Dorsally they may vary from slate-grey to red-brown (Figure 2.1).
This backgrouod colour matches the colour of the rocks on which the lizards
live. The entire dorsal surface is stippled or mottled with black to varying
degrees. These markings tend to form an irregular dark dorsolateral band in
older females. The dorsum of the body is covered with small regular to large
irregularly shaped white patches, which may be spread evenly or tend to form
transverse rows across the back. Ventral surfaces are white to grey, being
darker if the dorsum is also darker (Figure 2.2). The throat has a pattern of
grey lines which may vary in individuals from completely covering the throat
45
Chapter2 SEXUAL DIMORPIDSM
with a variegated network to a few irregular lines along the lower jaw. There
is a distinctive dark-edged pale patch behind and about the same size as the
eye.
Adult males are pale grey to dark grey dorsally (Figure 2.1 ). The lateral
surfitces, through to the entire dorsal surfitce ofthe body, may be black. The
dorsum of the body is covered with small regular to large irregular white or
yellow patches, which may be spread evenly or tend to form transverse rows
across the back. Ventral surfitces are white apart from a black chest patch
which extends from the gu1ar fold onto the anterior and ventral surfitces of the
upper arm, and back along the midline to at least haffi.vay along the body
(Figure 2.2 & 2.10). This chest patch varies between 7 and 23 percent of the
total ventral surfitce of the body (x= 14.6%, SD= 2.6%) (Figure 2.10), but
there is no relationship between body size and the proportion of the ventral
surfitce with black pigment (F1.ss = 0.000, P = 0.984; Figure 2.11). The throat
is usually yellow, but may occasionally be white, and has a network of grey-
.blue Jines over the whole gu1ar surfitce. These Jines may vary from a
variegated network to a series of distinct lines. There is a wide pale line
running from below the eye, under the tympanum and onto the scapular fold,
which is always bordered by black.
Measurements of Ctenophorus flormi from Mmmt Young illustrate the

differences in dorsal reflectance between the sexes (Figure 2.12). Both sexes
show an increase in reflectance at greater wavelengths, but adult females were
significantly more reflectant than adult males ( T = 2.68, P = 0.007).
Juvenile males begin to develop adult colouration once they reach about
45mm SVL. First, a grey wash develops on the chest. This eventually deepens
46
Cbapter2 SEXUAL DIMORPHISM
in colour and expands to form the black chest patch. Shortly after the grey
wash appears on the chest, the gular fold becomes yellow. This yellow
expands anteriorly until it covers the entire throat. While this is happening the
1k<-
darker reticulations and lines develop on.(throat. Some time after these
distinctive male ventral patterns have appeared (usually at the end of their first
summer) the dorsum begins to darken (Figure 2.4 & 2.13). fuitially black
pigment is restricted to irregular stippling over the entire dorsal surlace
(melanisation score of I). Then, as more melanin is laid down, a series of dark
blotches appear along the dorsolateral line (melanisation score 2). These
blotches eventually coalesce to form a more or less continuous black
dorsolateral line, which is only broken by the white patches which cover the
whole dorsal surlace (melanisation score 3). As the background dorsal colour
changes from the juvenile brown to grey the dorsolateral line expands to cover
the entire lateral surlace, leaving only a grey vertebral line (melauisation score
4). Ultimately melanin is also laid down over the vertebrae, so that the black
now covers the entire dorsal surlace of the body, except for the white areas
remaining from the juvenile colour pattern (melanisation score 5). At the same
time the dorsal surfaces of the head, legs and tail lose the juvenile brown
colour and become grey.
Melanin is laid down on the dorsum of both male and female Peninsula
Dragons as they grow (Figure 2.13). All liurds scored for the degree of
melanisation as hatchlings had a melanisation score of l. Males then gradually
progressed through all levels ofmelanisation outlined above, so that all adult
males had achieved a score of at least 4 and most continued to darken and
achieve a score of 5. No females were given scores of greater than three, so
that only males achieved scores of 4 and 5 (Figure 2.13). Among liurds
assigned a score of 2, males were significantly smaller (x = 52.9, SD= 5.71, N
47
= 20) than females (x = 68.0, SD= 4. 74, N = 57) (F1,1s = 133.69, P = 0.000).
Similarly, among lizards assigned a score of 3, males were significantly smaller
(x = 61.6, SD= 2.76, N = 22) than females (x = 74.7, SD= 4.32, N = 47) (U
= 5, P = 0.0000). No valid comparison was possible between males and
females given scores of 1 because many lizards in this category could not be
sexed accurately.
DISCUSSION
Sexual dimorphism in adults
Adult male and female Peninsula Dragons differ in several respects. Males
grow to a larger body size, develop relatively larger heads, and show more
extensive melanisation on the dorsum than feniales. Adult males also develop
yellow throat colouration and a black chest patch, which are absent in adult
females. Immature male and female Peninsula Dragons are very similar to each
other in size, shape and colour.
The degree of sexual size dimol]lhism (SSD = mean male SVL/mean female
SVL = 1.07) in this study is well within the range of values reported
previously for Peninsula Dragons (SSD = 1.02 - 1.14; Houston 1974). It is
also within the values reported previously for other agamid lizards (SSD =
0.86 - 1.36; Stamps 1983). Large male body size in lizards is widely regarded
as a result of selection for increased size associated with male combat (Stamps
1983; Ruby 1981; Tokarz 1985; Hews 1990; Olsson 1992). However,
estimates of SSD must be inteI]Jreted cautiously. Male-biased SSD may be
48
explained as a result of intrasexual selection, but may also be due to

differences in suzvivorship between the sexes (Dunham 1981; Stamps 1993).
For example, the average SVL of males may be larger ifboth sexes grow at
i
the same rate but males live longer. This possib~ is examined in Chapter 3.
In addition to the clear differences in body size between male and female
Peninsula Dragons, the discriminant function analysis of body measurements
showed that adults of each sex differ from each other in shape. Of all the
measurements, HW and HLL have the highest correlations with the
discriminant function, suggesting that the major differences between males and
femalesare in the relative size of the head and relative length of the hindlimbs:
HW also has a very large discriminant function coefficient (of similar
magnitude as SVL and MaS!l; Table 2.1 ), emphasising that relative head size is
a particularly important difference between the sexes. Previous studies have
shown that the relatively larger head of male lizards may be used as a weapon
during male combat (Carothers 1984), and that a relatively large head may
confer an advantage among competing conspecific male lizards of similar body
size (Cooper & Vitt 1989; Hews 1990). The large head in male lizards may
also be a means of emphasising signal colours on the head and throat for
intrasexual and/or intersexual communication (Hasson 1989; Tokarz 1995), or
may be a means of partitioning food resources between males and females
(Schoener 1967). HLL is known to be.an important determinant oflocomotor
performance (Garland 1985) and high locomotor performance is correlated
with social dominance in some lizards (Garland, Hankins & Huey 1990). HLL
may be particularly important in C. fio1mi and other rock dwelling species of

Ctenophorus because they perform a hind leg pushup display during male
contests (Gibbons 1979). The hypothesis that sexual differences in head size
may be a means of partitioning resources among the sexes is investigated in
49
Chapter 5. The hypothesis that the differences in head size and HLL between
the sexes are the result of male-male competition is examined in Chapter 7.
Although the sexual d.iffurences in body size and shape are clear enough, adult
male and female Peninsula Dragons differ most obviously in colour. Adult
male C. flonni have a darker dorsum than the females, and also distinctive,
bright colours on the head. Both are features of other territorial lizard species.
Males also have black chest patches which are absent in females.
Distinctive dorsal body colour occurs in males of many lizards and appears to
be a signal of social status (Werner 1978; Madsen & Loman 1987; Zucker
1989, 1994). These colours may change depending on the social status of
males of some species (Madsen & Loman 1987; Zucker 1989, 1995), but
darkening of the dorsum in C. ftonni is irreversible. For Peninsula Dragons,
this may be a nonmodifiable signal of a male's social status, similar to the dark
plumage of male red-winged blackbirds (Roskaft & Rohwer 1987).

Alternatively, the dark colour may play a role in thermoregulation by affecting
the rate of heat exchange with the environment (Norris 1967). While this is a
plaust11le function of the dorsal colouration;the black chest of males is unlikely
to be important in thermoregulation because it is not exposed during
heliothermic basking. During thlgmothermic basking, contact between the
ventral surface of the lizard and the substrate is more likely to be important
than the reflective properties of the lizard. fudividual differences in the extent
ofblack pigment are known to play an important role in the energetics and
behavioural ecology ofheliotherms (Kingsolver I 985).
The bright colours on the head and throat of male territorial lizards have been
interpreted as signals involved in species recognition (eg. Williams & Rand
50
1977; Losos 1985), mate recognition (Greenberg & Noble 1944; Sigmund
1983; Tokarz 1995) or recognition of riva}s (Werner 1978; Madsen & Loman
se)('ve;I oti'~n!.,ces ,.,.. _
1987). The function of the.·/.. ·colouration.. .:,f Peninsula Dragons is
investigated in Chapter 8.
Ontogeny of the Sexual Dimorphisms
Separate analysis of allometry in juvenile males and juvenile females would

have allowed a direct test of the assumption that juveniles of both sexes show
a common allometric relationship. Unfortunately, the small number of
juveniles for which sex could be determined precluded this analysis.
Nevertheless, the similarity in the magnitude of the coefficients of
determination (R2) between all juvenile and adult female regressions (Table
2.3) suggests that juveniles of both sexes do show the same allometric
relationships. Ifjuvenile males and females showed different allometric
relationships, but were analysed jointly, one would expect smaller values ofR2
associated with the regression ofjuvenile lizards for each trait. On this
assumption, both male and female adults diverge from the common juvenile
pattern of allometry for almost all traits. This suggests that any selection
acting on sexual differences in body shape is affecting adults only (contra
Shine 1990). A similar finding has been reported for at least one other lizard
(Hews 1996). The one exception to this pattern in C. ftonni is that females and
juveniles do not differ in their allometric coefficients for HD.
When different patterns of relative growth are based on static data (i.e. where
repeated measures of relative size are not available over the life time of
51
individuals), they must be viewed with some caution. Males and females may
exhibit a common ontogenetic change in the relative size of a trait, and may
differ in survivorship, perhaps due to sexual differences in the costs of
reproduction. If this were the case, then the difference in survivorship may
give rise to differences in size between the sexes. In tum, this would lead to
differences in relative size of the trait. These 'apparent differences in body size
and allometry would therefore be artefacts of the sexual differences in
survivorship, rather than true differences between the sexes (Dunham 1981).
In this study, the differences in allometry are not likely to be artefacts because
male and female body sizes overlap broadly. Although this may not be true if
selection were acting directly on relative, rather than absolute, trait sizes.
Males diverge from the juvenile relationship between SVL and Mass through
'time' hypermorphosis (Shea 1983), in which size increase occurs either
because sexual maturity is delayed and the period of relatively rapid juvenile
growth is extended, or because growth continues over a prolonged adult
period. In the three head size traits males diverge from the common juvenile
shape by 'rate' hypermorphosis (Shea 1983), in which the relative size ofa
trait increases and thus extends beyond the normal developmental trajectory.
Males exht'bit a lower allometric coefficient than juveniles for all limb and tail
measurements, suggesting they are peramorphic for these body measurements.
Females, on the other hand, show a lower allometric coefficients than juveniles
for all body measurements except HD.
As with body size and shape, the dorsal colour pattern of adult male Peninsula
Dragons develops from a common female and juvenile colour pattern (Figure
2.13). Subadult males resemble larger, older females in developing broad black
dorsolateral lines. Then as the males grow, the brown parts of the dorsum
52
become saturated with melanin. Thus the dorsal darkening in males appears to
be the resuh ofhypermorphosis in the degree ofmelanism The lack of
distinctive colouration in juvenile male C. fionni may allow them to avoid
intraspecific aggression since adult male colouration releases aggression in
many lizards (Cooper and Greenberg 1992) or to avoid predation because the
paler dorsal colour blends with the substrate (Booth 1990). Whether adult
male colouration incurs increased predation is examined in Chapter 4. The role
of female-like colouration in juvenile males is examined in Chapter 6.
Describing male Peninsula Dragons as peramorphic (extending beyond the

juvenile form), and females as paramorphic (resembling the juvenile form),
does not imply that either sex has been subject to evohttionary heterochrony.
Here the terms simply refer to the intraspecific differences in the patterns of
relative growth during ontogeny. Phylogenetic analysis of ontogenetic changes
in shape between several species of Ctenophorus, and at least one ontgroup,
would be necessary before cladistic directionality can be ascribed to
heterochronic traits (Fink 1982).
In summary, adult male and female Peninsula· Dragons differ from each other
in body size, shape and colouration. There is no evidence that the sexes differ
in any trait before they become mature, but upon reaching maturity males and
females diverge from a common juvenile pattern. Males tend to grow to a
larger size than females and exhibit an increase in relative growth, whereas
females exhibit a decrease in relative growth. Adult males also develop unique,
sex-limited traits. These patterns suggest that selection is acting mostly upon
adults, and on adult males in particular.
53
Chapter2 SEXUAL DlMORPIDSM
Trait coefficient correlation

logSnout-vent Length -40.723 0.309
logMass C:0.233 0.397
logHead Length 1_ ••• 046
('c.'"4 0.483
logHead Depth :}6.415 0.319
logHead Width ·.,29.986 0.622
logForelimb Length ·\.5.904 0.497
logHindlimb Length '"'15.092
L,.,, . 0.572
logTail Length ::;~5.216 0.437
constant l1-7.793
TABLE 2.1: Unstandardised discriminant function coefficients and pooled-

within-groups correlations with discriminant function based on eight
measurements of adult male and female Ctenophorus fionni from Midgee Rocks.
See Figure 2.6. . . '
54
Males (N=l28} Females N=92)
Variable x(SD) x{SD) F-statistic
Snout-vent length (mm) 76.2(9.ll) 70.6(6.09) 26.3345***
Body mass (g) 19.8(6.13) 14.6(3.53) 54.3564***
Head length (mm) 18.7(2.45) 16.5(1.28) 62.0728***
Head width (mm) 19.7(2.70) 16.8(1.05) 95.8303***
Head depth (mm) 8.8(1.33) 8.0(0.76) 28.5734***
"'
V, Forelimb. length (mm) 30.5(3.19) 27.6(1.73) 62.7945***
Hind.limb length (mm) 63.8(7.26) 56.0(4.17) 87.4777***
Tail length (mm) 142.5(6.13) 127.1(11.21) 49.9688***
TABLE 2.2: Comparison of measurements from sexually mature male and female
Ctenophoros fionni used in a discriminant :function analysis. N = sample size, x = mean,
SD= standard deviation,***= P<0.001.
Juveniles Adult Females Adult Males F-values
Trait N a(SE) R• N a(SE) R• N a(SE) R• jvf jvm fun
Head Length (mm) 121 0.86(0.042) 0.77 107 O. 72(0.0S3) 0.63 147 l.01(0.036) 0.85 051.48••• 021.01••• 10.09••
Head Width (mm) 120 0.94(0.043) 0.79 107 0.63(0.044) 0.65 · 147 1.03(0.041) 0.81 092.ss••• 032.96*** 18.J9•••
Head Depth (mm) 121 0.74(0.069) 0.49 107 0. 76(0.082) OAS 145 1.17(0.046) 0.82 000.59 007.3t•• 16.23**•
Forelimb Length (mm) 119 0.89(0.032) 0.117 !OS O.SS(0.045) 0,59 147 0,81(0.026) 0.87 163.32*** 151.13*** 14.81***
"'°' Hindlimb Length (mm)· 119 0.92(0.079) O.S4 105 O.S7(0.063) 0.44 147 0.8S(O.Q38) 0.77 039.71••• 036.8!*** 06.74*
Tail Length (mm) 116 1.16(0.046) 0,84 !OS 0,78(0.066) 0.60 147 1.00(0.034) 0.88 096.2S*** 122.26**• 05.56*
Mass(g). 121 3.03(0.097) 0.119 105 2.117(0.090) 0.91 147 2.92(0.049) 0.96 090.11*•• 198.87*** 00.77
TABLE 2.3: Sample sizes (N), allometric coefficients (a), coefficients of detennination (R') and statistical comparisons of aDometric coefficients for
seven traits among juvenile, adult female and adult male Cterwphorus jionnl. ADometric coefficients were compared using Analysis of Covariance to test
for equality of slopes on a log-log plot of the trait against Snout-vent Length (mm). • P < 0.05; •• P < 0.01, ••• P < 0.001.
Chapter 2 SEXUAL DIMORPlilSM
FIGURE 2.1 : Dorsolateral view of.male (above) and female (below)

CtenophoniS fionni.
57
FIGURE 2.2: Ventral view of male (above) and female (below) Ctenophorus
fionni .
58
/\
tHW
'
....J
~-FLL--1
~
FIGURE 2.3: Measurements taken from Ctermphorus ftanni.
59
Chapter2 SEXUAL DIMORPIDSM
'.
:".'
,,
'
4
FIGURE 2.4: Scoring system used to classify the degree of melanisation of
the dorsal surface ofCtenophorusjionni.
60
..JO
:V1ALES
30 N = 152
20
10
>-
u
zUl
::, 0
Cl 20
:.:.J
~
:.:. FEMALES
15 N= 114
10
0
50 60 70 80 90
SNOUT-VENT LENGTH (nun)
FIGURE 2.5: Size frequencies of male and female Ctenophorus fionni

measured at Midgee Rocks between 1991 and 1994. Sexual maturity occurs
between 50 and 55mm SVL in both sexes.
61
0
-6 0 6
DISCRIMINANT FUNCTION I
FIGURE 2.6: Frequency histogram of discriminant function scores for adult
male (hollow), adult female (hatched), and juvenile (solid) Ctenophorus fionni
from Midgee Rocks.
62
30
-
~
CJ
1--
:x:
(!) 20 0
iii
s:
10
.........
0 .•~-~-.--..-
50 60 70 80 90
SNOUT-VENT LENGTH (mm)
FIGURE 2.7: Relationship between Mass and Snout-vent Length in male(•),

female ( 0) andjuvenile (•) Ctenophorus fionni from Midgee Rocks.
Allometric coefficients for these groups are given and compared in Table 2.3.
63
Chapter2 SEXUAL D!MORPIDSM
25,~---------------------~--,
. ......
5,.J-.---~---~---~---~---i------,
30 40. 50 60 70
FIGURE 2.8: Relationship between head width and Snout-vent Length in

male (•), female (0) and juvenile(•) Ctenophoros fionni from Midgee
Rocks. Allometric coefficients fur these groups are given and compared in
Table 2.3.
64
70
--- --- --
SNOUT-VENT LENGTH {mm)
FIGURE Z.9: Relationship between Hindlimb Length and Snout-vent Length

in male ( • ), female (0) and juvenile (•) Ctenophorus jionni from "Midgee
Rocks. Allometric coefficients for these groups are given and compared in
Table 2.3.
65
Cbapter2 SEXUAL DIMORPHISM
A B C
FIGURE 1.10: Variation in the extent of the black pigment on the ventral
surface of adult male Ctenophorus jionni at Midgee Rocks. The three
diagrams represent tracings of the extent of black pigment taken from
photographs and representing males with the smallest (A), average (B) and
largest 'chest patches'. Black pigment covered 7.1%, 14.6 % and 23.2% ofthe
total ventral surface of these individuals, respectively.
66
Chapter 2 SEXUAL DlMORPHlSM
0.3
~
CG
-< •
..J
~
f- 0.2
z
t.:;J • ••
> •
..J
• •
gf- 0.1
••
••
•
•
0.(1 L..-..~~---1..~~~-'--~~~'-----~~----'
50 60 70 80 90
FIGURE 2.11: Relationship between the proportion of the ventral surface

that is covered with black pigment and SVL among 59 male Ctenophorus
fionni at Midgee Rocks. The line is a linear regression with the equation Y =
0.148 - 0.000X, indicating that the size of the black chest patch does not vary
with size (F1,sa = 0.00, P = 0.984).
67
20
..-.. 15
'cf:.
'-'
(.IJ
u
z
<: 10
E-
u
(.IJ
...J
µ;..
(.IJ
a: 5
0
400 500 600 700
WAVELENGTH (µm)
· FIGURE 2.12: Reflectance over the human visual spectrum of dorsal skin of
male (0) and female(•) Ctenophorusfionni.
68
5 ...
0
~ 4 .....,,,,, • •
u
J. ~ OJ!
r./J
0::: 3 Ill, L, c£ffrul 0
;:)
0
...J
0
u 2 -- ••• • J., ~H!l1~ o
l ~.dllid1iRWao o •
30 40 50 60 70 80 90
FIGURE 2.13: Size distnlmtion of male(•) and female (0) Ctenophorus

fionni from Midgee Rocks with different dorsal color scores. The colour
scores are based on Figure 2.4.
69
CHAPTER3
REPRODUCTION
Dragon lizards show considerable variation in their reproductive

biology. They may be sexual or parthenogenetic (Hall 1970),
oviparous or ovoviviparous (Greer 1989). They may lay clutches of
one or two eggs continuously throughout the year (Inger &
Greenberg 1966; Allison 1982), or lay clutches of 2-35 eggs in a
strongly seasonal pattern (Greer 1989).
Most Australian dragon lizards have highly seasonal reproductive

cycles. They undergo vitellogenesis in August and September, mate
and lay eggs later in September and October, and hatch between ·
January and March(Bradshaw 1981; James & Shine 1985). This
means that reproduction tends to occur synchronously during spring
and su=er in southern Australia and during 'the wet' in the tropical
north of the country. There are several exceptions to this pattern. At
least one tropical species (Chelosania brunnea) breeds from July to
Chapter 3 REPRODUCTION
September, during the tropical dry (Pengilley 1982; James & Shine
1985). Females of a temperate species, Ctenophorus caudicinctus,
are gravid during February and March (in late summer and
autumn)(Storr 1967; Bradshaw 1986; Bradshaw, Saint Girons &
Bradshaw 1991). Pogona minor breed between September and March
in the southern part of their range (Davidge 1980), and breed
between July and September in the northern part of their range
(Bradshaw 1981). In high altitude areas of New South Wales
Amphibolurus nobbi mate and lay eggs near hibernacula during
spring and autumn, just after and just before a winter period of
inactivity (Witten 1974).
Many smaller Australian dragon lizards are sexually mature at less

than one year of age (Tyler 1960; Storr 1965, 1967; Pianka 1971;
Bradshaw 1971, 1981; Cogger 1978; Baverstock 1979; Witten &
Coventry 1984) and appear to survive and breed for only one year
(Cogger 1978; Pianka 1971; Haverstock 1979; Bradshaw 1981,
1986). In some of these species females may lay up to three clutches
in one breeding season, depending on environmental conditions, but a
single clutch is usual for many (Bradshaw 1981 ). Thus many of these
'annual' species are essentially semelparous. Other, generally larger,
dragon lizards may mature at ages between two and five years (Hay
1972; Bradshaw 1971; Davidge 1980) and are iteroparous. They lay
between one and three clutches per year over many years (Badham
1971; Mitchell 1973; Bradshaw 1981; Griffiths 1994 ).
71
This variation in reproductive and life-history traits among dragon

lizards emphasises the need to collect data on each individual
species, rather than relying on inference from patterns of related
species. This chapter presents data on reproduction in Ctenophorus
fionni for the first time. The data reported here were collected
incidentally. Therefore· they are patchy, and do not present a
complete picture of the reproductive biology of this species.
Nevertheless, they are important, and are presented here for three
reasons. First, they provide information on the reproductive cycles in
this species, which allowed behavioural observations and field
experiments to be planned for appropriate times of the year.
Secondly, they provide information on the age at maturity, frequency
of reproduction and clutch sizes. These parameters are useful for
estimating the potential for sexual selection (Sutherland 1985 ).
Lastly, they allow an assessment of whether fecundity selection may
influence female size in Peninsula Dragons. Fecundity selection can
only act in species which exhibit a positive relationship between
fecundity and female body size (Darwin 1871; Shine 1988b).
METHODS
The study of reproductive cycles was based on 120 Ctenophorus

fionni collected from Iron Duke (Figure 1.2) and lodged in the South
Australian Museum. This collection was made by the author between
April 1985 and April 1986 just prior to Iron Duke being cleared for
an extensive open cut mine. Over this period the area was visited in
72
most months. During each visit lizards (N 4-14 per visit) were
collected by ha~d, and killed by freezing within 12 hours of capture.
/II
They were fixedi{10% formalin and stored in 70% alcohol. These
specimens also form the basis for a study of diet in C. fionni
(Chapter 5).
Iron Duke is the southernmost hill in the Middleback Ranges and lies
12 km N ofMidgee Rocks, my main study site. The nearest
meteorological station to both Iron Duke and Midgee Rocks is at
Whyalla (Figure 1.2). The population of Peninsula Dragons which
inhabit the rocky outcrops at Iron Duke is completely isolated from
the one at Midgee Rocks by several kilometres of unsuitable sand
dune country. The vegetation at Iron Duke is Eucalyptus oleosa-E.
brachycalyx mallee woodland. A more detailed description of the
Iron Duke area is provided by Johnston (1982).
For each lizard snout-vent length (SVL) was measured and the
reproductive tract was inspected through a small, lateral incision into
the peritoneal cavity. In females the number of vitellogenic follicles
and corpora lutea were counted for each ovary. The maximum
diameter of the largest follicle was measured and any oviductal eggs
were counted. The presence ofvitellogenic follicles or corpora lutea
was taken as an indication of reproductive maturity. In males the
maximum width (w) and length (/) of the right testis were measured
and used to calculate the testicular volume (V) in ml using the
formula:
V = 4/3i.(/2w/2).
73
A gonadosomatic index (GSI) was calculated as the ratio between

testis volume and SVL. The epididymides were classified as
translucent ( empty) or opaque ( containing sperm). The presence of
opaque epididymides was used to assess reproductive maturity among
males. Male lizards were pooled into collections from spring
(August-October), summer (November-January), autumn (February-
April) and winter (May-July) to allow statistical analysis of the
testicular cycle. A oneway analysis of variance was used to test the
null hypothesis that the GSI of mature males did not differ between
these four seasons.
Observations of the reproductive cycle based on the specimens from

Iron Duke were supplemented with observations made at two other
locations which were each visited several times between 1991 and
1994. A mark-recapture study at Midgee Rocks between 1991 and
1994 provided additional information on the frequency of
reproduction. Gravid females were also collected in October and
November from Siam Station (Figure 1.2) and Midgee Rocks in 1992
to determine clutch size. These females were taken to the laboratory,
where they were housed individually in clear plastic containers
(250mm x 200mm x 100mm) with flywire lids and overhead,
broadspectrum heat lamps. Dampened paper towelling was placed on
the floor of the containers to prevent dehydration of any eggs. Each
,9.t,~d'1 gmvid
Afemale was given weekly intraperitoneal injections of Oxytocin ,
(HERIOT AGVET, Rowville, Victoria) at a dosage of 6.0ml/kg until
they layed. For each female, SVL was recorded along with clutch
size, egg lengths, egg widths and egg weights. All lengths were
74
measured with vernier callipers to the nearest 0.1mm. Weights were

measured to the nearest 0. lg with Mettler top loading electronic
balances. Not all measurements were taken from all clutches, so
sample sizes varied for different statistical analyses. After
oviposition most females were returned to the site from which they
were captured.
A nonparametric ANOVA showed that clutch sizes did not differ

between females from Iron Duke, Midgee Rocks and Siam Station.
Thus these data were pooled. Linear, regression was then used to test
the null hypothesis that female SVL and clutch size were not related.
Eggs were placed into moist vermiculite in 500 ml plastic containers,

covered with a layer of damp tissue paper and with cling-wrap plastic
stretched across the top of the container following Johnston (1979).
The eggs were then incubated in controlled temperature water baths
at either 24 C, 28 C, or 32 C. The SVL (mm) and body weight (g)
of each hatchling were measured using he same equipment used to
measure the eggs.
Additional data on interclutch intervals were gathered using three

females from Mount Young (Figure 1.2) in September 1991. They
were housed in an outside enclosure (1.5m x 3.0m) in Whyalla with a
single male. The walls of the enclosure were 0. 7m high sheet metal
buried into the sand. The enclosure was covered with wire netting
and the lizards were kept on a natural sand substrate. Blocks of
sandstone were placed in two piles at each end of the enclosure to
provide shelter. Water and insects (mealworms, crickets &
75
cockroaches) were made available ail libitum in an attempt to

provide optimal conditions in which the females might lay multiple
clutches. These lizards were checked ,daily to determine whether they
had laid eggs. Oviductal eggs distended the body wall of gravid
females. When a female had laid this was clearly apparent due to her
sudden emaciated appearance.
Observations of reproductive behaviour in the field were niade at

Midgee Rocks in most months between August and February in each
year from 1991 to 1994.
RESULTS
Size and Age at Maturity
The smallest female from Iron Duke with vitellogenic follicles or

corpora lutea had an SVL of 60mm. The largest immature female had·· .
'
an SVL of 53mm. The smallest male with opaque epididymides had
an-SVL of 55mm and the largest male to have transparent
epididymides had an SVL of 51mm. At Midgee Rocks the smallest
male to have the characteristic chest patch of adult males had an
SVL of was 55mm (Figure 2.11). Males at Midgee Rocks were darker
than females by the time they reach an SVL of 50mm (Figure 2.13).
Thus Peninsula Dragons from both of these localities appear to reach
maturity at similar sizes. Lizards caught at Midgee .Rocks reached
50-60mm, the minimum size at maturity, by April when they were
76
about 3 month old. This pattern has been confirmed by formal growth
models for both sexes (Chapter 4).
Most Peninsula Dragons at Midgee Rocks appear not to breed until

their second spring breeding season, despite having achieved a size at
which they would be reproductively mature in their first breeding
season. Two subadult males were observed to pair(and presumably
mate -see below) with females after older, larger, territory-holding
males who had been consorting with the females died (see Chapter
6). No females at Midgee Rocks were found to be gravid prior to
entering their second spring, when they were 20 months old.
Reproductive Cycle
Enlarged, vitellogenic ovarian follicles were present in females from

Iron Duke in September and November. Oviductal eggs were present
in November (2/8 females) and December ( 4/6 females). Gravid
females were found at Midgee Rocks, Siam Station and Mount Young
in September, October and November in all years of the study.
Gravid females were also found in December 1992 and January 1993
at Mount Young (Figure 3.1).
Among mature males from Iron Duke the GSI was significantly
different between seasons (FJ,58 = 22.25, P < 0.001). The spring
mean was significantly higher than the means -in all other sea sons
(Scheffe P < 0.0S)(Figure 3.1). The testes were enlarged with
77
obvious seminiferous tubules during August and September. In all

other months the testes were flaccid and seminiferous tubules were
not visible on gross dissection.
Frequency of Reproduction
Among the twenty adult females collected from August to December

at Iron Duke most (85%) showed either enlarged, vitellogenic
follicles or oviductal eggs, indicating that they would pro duce a
clutch of eggs, or corpora lutea, indicating that they had already
produced a clutch.
Repeated captures of adult females within and among years at

Midgee Rocks showed that the proportion of females found gravid at
least once in a breeding season ranged from 19/30 (63%) in 1991 to
1/15 (7%) in 1994. Most mature females (43/45 = 96%) at Midgee
Rocks were recorded to produce one clutch in any one year, implying
that multiple clutches were abnormal. The two females who produced
two clutches in a season were recorded gravid in late September and
again in late November. Most females (41/45 = 91%) were recorded
gravid in only one year during the study. The four females who were
gravid in more than one year were gravid in two successive years. No
females were recorded gravid in more than two breeding seasons in
succession.
78
Chapter J REPRODUCTION
All three of the females kept in Whyalla mated and produced

clutches. Two of them produced two clutches in the 1991 breeding
season. One female laid her first clutch on 26 November 1991 and
laid a second clutch on 4 and 5 January 1992. The second female laid
her first clutch on 9 December 1991 and laid her second clutch on 20
January 1992. Thus the minimum period between the production of
two clutches was 39 days. The third female laid her clutch on 30
November 1991.
Clutch Size, Incubation and Hatchlings
There was no significant variation in clutch size between females

collected at Iron Duke (N = 20), Siam Station (N = 9), and Midgee
Rocks (N = 7)(Kruskal-Wallis ANOVA, r,. 2 = 1.872, P = 0.599) so
clutches were pooled for the following analysis. Clutch size varied
from 2 to 6 (x = 4.24, SD= 1.130, N = 36), and showed no
significant relationship to maternal SVL (F 1 35 = 0. 99, P =
0.158)(Figure 3.2).
The majority of clutches incubated in water baths failed to hatch due

to fungal infections. The few eggs that did hatch showed that the
incubation period varied according to the temperature at which the
eggs were kept. Five eggs incubated at 24 C took 145 days to hatch,
whereas three eggs incubated at 32 C took 57 days to hatch. A
further 13 eggs, incubated at fluctuating temperatures between 28 C
79
and 30 C, took between 50 and 87 days to hatch. dimensions of eggs

and hatchlings are given in Table 3.1.
Peninsula dragons lay their eggs in burrows constructed in soil

beneath rocks. At Midgee rocks a gravid female (#46) was observed
digging into a soil-filled crevice on 30 October 1991 (Figure 3.3).
Two hatched, dried Peninsula Dragon eggs were found beneath a
granite boulder in February 1993 (Figure 3.4). Hatchlings appeared
in the Midgee Rocks population in January and February of each year
(Figure 3.5).
Pairing and Mating Behaviour
Although mating was not observed directly during this study, two
interesting aspects of mating behaviour were observed at Midgee
Rocks. First, females were observed to reject the sexual advances of
some males. Second, adults were found in heterosexual pairs in
spring.
On 1 September 1991, a pair of adults was observed from late

morning (11:30) to mid afternoon (14:00) at Midgee Rocks. During
this time the male (#37) spent most of his time perched on top of a
low granite boulder, about 0. 7m above the surrounding flat soil. The
female (#95) spent most of her time stationary not far from the
entrance to a crevice in the same boulder, 0.3m from the male. The
crevice was just wide enough for the female's body. Every 15
80
minutes, or so, the male dashed at the female and grabbed her on the
back of the neck with his jaws. She invariably responded by
retreating quickly into the nearby crevice. This dislodged the ardent
male, who rolled, untidily down the rock. He would then bob his
head several times before returning to the top of the rock. These
head-bobs did not seem to be directed toward anything in particular.
The female remained concealed in the crevice for several minutes
before she would move cautiously out into the sun again. This
sequence of events continued for the entire period of observation.
On another occasion, on 3 October 1991 at Midgee Rocks, an adult

female (#48) was observed perched on top of a low rock on a large
expanse of flat granite in mid afternoon. She had been stationary for
115 minutes before an adult male (#150) moved towards her from
some vegetation about Sm away. The male crossed the flat area in a
series of short(± 1.0 m) sprints, punctuated by vigorous headbobs.
When the male was 0.2 m from the female he stopped and
headbobbed toward her continuously for 5-10 seconds. He moved
0.05-0.1 m around her between each headbob, until he was behind
her. He then rushed at the female and attempted to grab her on the
back of the neck. At this point the female moved so that the male
was rolled onto the ground. She raised her tail in an arc and
extended her hind legs, so her cloaca was exposed (Figure 3.6). The
male, after regaining his composure, headbobbed again, then moved
toward the female and licked her cloaca. He then stopped
headbobbing, appeared to lose all interest in the female and moved
away from her after about 15 seconds. The female held her posture
until the male moved off. When the female was captured, 20 minutes
81
after the male had left her, she was found to contain palpable
oviductal eggs.
Adult C. fionni were found in heterosexual pairs during August,

September and October. Paired lizards were found within 0.5 m of
each other while active, or in bodily contact within a crevice when
inactive. Forty five pairs found during the study were recorded
together for l-10 days (x = 2.2, SD= 2. 78). All females found with a
male for more than one day had palpable ovarian follicles when they
were found in a pair, but not at the end of the consort ship. These
females were assumed to have ovulated during the partnership.
DISCUSSION
The results presented here show that Peninsula Dragons normally

breed between August and November. Males have enlarged testes in
August and September. Males and females were found in pairs from
August to October, when mating is assumed to occur. Females were
normally gravid between September and November, and normally
produced a single clutch of between two and .six (average 4.2) eggs.
The eggs were laid in burrows in soil beneath rocks which females
were observed digging. The eggs incubated over the summer and
hatchlings appeared in the field in January and February. Thus
Peninsula Dragons are similar in their timing of reproduction to the
majority of other Australian dragon lizards (Greer 1989).
82
Like other dragon lizards, Peninsula Dragons are able to respond to

appropriate environmental conditions by extending their reproductive
season. The Central Netted Dragon, Ctenophorus nuchalis, may
produce a single clutch of eggs in dry years, but may produce
clutches continuously from October to March in wet years (Bradshaw
et al. 1991). During the wet summer of 1992-1993 gravid Peninsula
Dragons were found until January at mount Young, but no gravid
females were found after November at Midgee Rocks in the same
year. Thus different populations of C. fionni respond differently in
the same year, possibly due to local conditions of food and water
availability. The majority of lizards at Midgee Rocks produced a
single clutch whereas lizards at Mount Young were probably able to
produce two or three clutches.
Two females Peninsula Dragons from Mount Young that were held in
captivity under conditions of excess food availabilityduring the
spring and summer of 1991-1992 produced multiple clutches. Wild
females at Mount Young produced no more than a single clutch in
the same year. This suggests that food availability may limit the
ability of females to produce multiple clutches, at least at Mount
Young.
The reproductive cycles of male and female C. fionni coincide so that

the testes are producing sperm at the same time as mating and
ovulation occur. Thus sperm storage, which occurs in some lizards at
similar latitudes (Smyth & Smith 1968), is unnecessary in Peninsula
Dragons. However, adult males did maintain opaque epididymides
throughout the year, and, in so far as this indicates the presence of
83
spermatozoa, it appears that males do store sperm. This may be

advantageous if females producing a second clutch mate between
clutches. However I do not know whether this occurs. No obvious
sexual activity was observed in the field after November. Sperm
storage in the female reproductive tract has been ·reported in many
lizards, including iguanids (Cuellar 1966), but I did not determine
whether it occurs in C. fionni.
Even though they may mature within four months of hatching, most
Peninsula Dragons at Midgee Rocks appear not to reproduce until
their second spring, when they are over 18 mo old. This delayed
reproduction may not occur in all populations because several gravid
females collected at Siam Station were of a size suggestive of their
being in their first season. This assumes that all populations studied
have the same growth rate, and age at maturity.
Females at Midgee Rocks may reproduce in two consecutive years,

although most captured in more than one year were found gravid
only once during this study. The period of about 40 days between
clutches limits the ability of females to produce multiple clutches in
most years. Peninsula Dragons are known to survive for at least six
years in captivity (personatobservations). Thus, despite its relatively
small size, C. fionni does not appear to be an annual species, unlike
many other Australian dragon lizards (Storr 1967; Pianka 1971;
Cogger 1978; Baverstock 1979; Bradshaw 1981, 1986).
The observations that females are capable of rejecting (and do reject)

the sexual advances of some males suggests that they may be capable
84
of mate choice. The posture which the gravid female used to present
her cloaca to the male (Figure 3.6) has been documented as a
'rejection'' posture in several other agamid and iguanid lizards
(Carpenter & Ferguson 1977). It may be that this behaviour indicates
that a female is not receptive, rather than exhibiting active mate
choice. The observation of the female who avoided mating by running
into a crevice is difficult to explain without further information about
her reproductive state at the time. The rejection posture may be a
means that females have developed to avoid the costs associated with
unwanted matings (see Olsson 1995).
The occurrence of heterosexual pairs has been reported in several

other species of lizards (Ruby 1981; Vitt & Cooper 1985; Bull 1988;
Olsson 1993b). It may be a form ofpostcopulatory mate guarding in
species where multiple paternity is common (Olsson 1993b; Olsson et
al. 1994). Alternatively the proximity of the male may stimulate the
female to ovulate and become receptive (Crews 1975). Either way,

pairing is likely to be associated with mating behaviour, if only
because it occurs coincidentally with sperm production in males and
ovulation in females.
The absence of a significant positive relationship between female

body size and clutch size in C. fionni is unusual among Australian
dragon lizards (Greer 1989). This lack of a size-fecundity
relationship suggests that fecundity selection does not act in this
species. It is interesting to note that several other closely related
species ofCtenophorus (eg. C. decresii Greer 1989; C. rufescens
Johnston, unpublished data) do show a fecundity size relationship. At
85
least one of these (C. rufescens) has lower sexual dimorphism in

body size than most other members 'of this genus. This could result
from fecundity selection favouring large female size, and sexual
selection favouring large male size, so that both sexes have similar
optimal body sizes. However, the idea of fecundity selection is
difficult to reconcile with other Australian dragon lizards in which
females are larger than males, but there is no female size-fecundity
relationship (eg. Moloch horridus, Pianka & Pianka 1970).
Most Australian dragon lizards lay eggs in isolated, specially

constructed burrows (Sporn 1955; Mitchell 1973; Smith 1974;
Hudson 1977; Cogger 1978; Johnston 1979). But Gonocephalus
spinipes lay their eggs in shallow scrapes in leaf litter at communal
nesting sites (Shea, Husband & Weigel 1991; Manning 1991).
Information about egg laying sites has been gained largely through
chance observations. My observation of a female digging a nesting
burrow and the chance finding of an old burrow during this study
were similarly fortuitous. During a twenty year study of Ctenophorus
ornatus inhabiting rook outcrops in Western Australia no nesting
burrows were found, despite considerable effort to look for them
(Bradshaw, personal communication 1993).
In summary, the reproductive cycles of male and female C. fionni

were highly synchronised. Enlarged testes, ovulation and mating all
occurred during the austral spring. At this time males and females
may be found in pairs. This pairing behaviour may offer a field assay
of mating success in the absence of actual observed matings. Most
females produced a single clutch of eggs, but some females in some
86
populations produced multiple clutches. Clutch size varied between

two and six. There was no relationship between female body size and
clutch size. So there was no mechanism by which fecundity selection
might act on female body size in this species.
87
Measurement N mean SD min-max

Eggs
length(mm) 45 20.3 1.52 16.8-24.0
width(mm) 45 11.0 0.81 9.0-12.7
weight(g) 45 1.4 0.28 0.7- 1.8
Hatchlings
SVL(mm) 23 30.9 1.68 28.0-34. 8
weight(g) 23 1.3 0.22 0.9- 1. 7
TABLE 3.1: Measurements of eggs and hatchling Ctenophorus

fionni.
88
4 0 0 0 8 8
~u
3
• •
rl.l
2 • •• • • • •
1 • I I • •• • •
1S
~
lial ••
~ 10
• ••
~
0
•
•
•
rl.l
0
~ s •
• •• • •• •
~ •
" • • • ••
0 • ••
J F M A M J J A s 0
• •
N D
MONTH
FIGURE 3.1: Upper panel: annual reproductive cycle in female

Ctenophorus fionni. A score of 1 = females with no vitellogenic
follicles; 2 = females with vitellogenic follicles < 5mm in diameter; 3
= females with vitellogenic follicles > 5mm in diameter; 4 = females
with oviductal eggs. Filled symbols = specimens from Iron Duke;
each dot represents an individual lizards. Open symbols=
observations made at Midgee Rocks, Mount Young and Siam Station;
each circle represents several lizards. Lower panel: annual
reproductive cycle in male Ctenophorus fionni from Iron Duke. The
curve is a LOWESS function which represents a weighted value of y
based on nearby values (Cleveland 1979, 1981).
90
8 • I I
' - ..... -
s • • -
• --·-
r.ll
=
uE--
4 ...
• -
;;;,
~
u
.
•• •• -
2
• -
-
I I
0 '
50 60 70 80 90
FIGURE 3.2: Relationship between clutch size and female Snout-

vent Length in Ctenophorus fionni.
91
FIGURE 3.3: Grav id female (#46) Ctenophorus fionni digging a

burrow at Midgee Rocks on 30 October 1991 .
92
FIGURE 3.4: Hatched eggs in a nesting burrow of Ctenophorus

fionni found beneath rock at Midgee Rocks in February 1993 .
Camera lens cap provides scale (diameter = 50 mm).
93
FIGURE 3.5: Hatchling Ctenophorus fionni with eggshells.

Australian one dollar coin provides scale (diameter = 25 mm) .
94
FIGURE 3.6: Female Ctenophorus fionni exposing her cloaca 'in a

'rejection' posture.
95
CHAPTER4
GROWTH AND SURVIVORSHIP:

PROXIMATE CAUSES OF SEXUAL SIZE
DIMORPHISM ?
In the majoritr of dioecious animal species, males and females differ

in body size, but the direction and extent of this sexual size
dimorphism (SSD) varies widely among populations and taxa.
Different body sizes in males and females may be favoured for at
least three reasons. First, males and females may be energetically
optimal at different body sizes (Downhower 1976; Robinson 1986).
Secondly, size selective predation may favour one sex (Owen-Smith
1993). Finally, the relationship between body size and reproductive
success may differ for males and females (Arak 1988).
In many species females are larger than males (Ghiselin 1974). This
size relationship is characteristic of mo st invertebrates and many
'lower' vertebrates (Shine 1979, 1994; Berry & Shine 1980). In at
Chapter 4 GROWTH & SUR VIVAL
least some of these taxa large female size may be due to the
advantages of increased fecundity with increases size (Darwin 1871;
Head 1995). Conversely, in many birds (Selander 1972; Wiley 1974),
mammals (Clutton-Brock et al. 1977; Alexander et al. 1979) and
some reptiles (Stamps 1983; Watkins 1996), males are larger than
females.
Large male body size may occur (1) because this is an ancestral
condition which has been derived from the parent taxon (eg.
Cheverud, Dow & Leutenegger 1985), (2) because genes for large
size confer a greater selective advantage in males than females (eg.
Darwin 1871; Trivers 1972), or (3) because males and females would
normally grow to the same size, but local environmental conditions
, favour survival and growth in males, more than in females ( eg.
Howard 1981; Woolbright 1983, 1989).
Males are larger than females in most species of iguanian lizards

(Fitch 1981 ). This pattern of SSD is presumably the ancestral
condition for this lineage and is widely assumed to be a result of
sexual selection for large male size in iguanians (Darwin 187 l;
Olsson & Madsen 1995; Tokarz 1995). But it is just as plausible that
higher mortality among females leads to smaller average female size,
than male size (Dunham 1981; Howard 1981), especially for species
which continue to grow after reaching sexual maturity (Shine 1990;
Stamps 1993). Higher female mortality could result from direct costs
of reproduction being greater in females than males ( eg. Schwarzkopf
& Shine 1992). Size-dependent mortality giving rise to unequal age-
distributions (and thus size distributions) between males and females
97
Chapter 4 GROWTH & SURVIVAL
offers an alternative proximate explanation for smaller female size.

These two hypotheses make different predictions which may be used
to test between them.
The hypothesis that male biased SSD is the result of sexual selection
for large male size makes two predictions. First, males should grow
faster and reach a higher asymptotic size than females because of a
positive relationship between body size and reproductive success
among males. Secondly, larger mature males should suffer equal or
greater mortality than females, due to indirect costs of reproduction
such as displaying for mates or maintaining territories.
The hypothesis that male biased SSD results from higher mortality
among females predicts that males and females should not differ in
growth pattern, unless there were a positive relationship between
size and fecundity in females. In this case females should grow faster
and to a larger asymptotic size than males. If females have a greater
direct cost of reproduction than males, then females should suffer
higher mortality at the onset of reproduction.
Male Peninsula Dragons reach a larger size than females (Houston

1974; Chapter 2). This chapter describes the pattern of growth and
survivorship among male and female Ctenophorus fionni to assess the
alternative hypotheses for SSD. The possibility that differential
mortality between males and females through predation is examined.
98
METHODS
Data were collected from lizards (N = 372) captured on seventeen

visits to Midgee Rocks on Eyre Peninsula (Figure 1.2) between April
1991 and November 1994. The average interval between trips was
2.5 mo. During each visit the study area was traversed in search of
lizards for five hours per day for five days. When lizards were
sighted and approached, they readily entered rock crevices, from
which they could usually be extracted. Each lizard was measured for
snout-vent length (SVL) to the nearest 0.1mm using vernier calipers,
sexed, permanently marked by clipping unique combinations of up to
two toes on each foot (Fenner 1979), and any damage to the tail was
noted. Sex was determined by probing (Schaefer 1934) or by everting
the hemipenes with gentle pressure at the base of the tail. Sex could
not be determined reliably for many juveniles. These were excluded
from all analyses, unless their sex could be determined upon
recapture at a later date. The process of capture, measurement and
release normally took less than 15 minutes per lizard, although some
lizards were retained in calico bags overnight. In the latter case they
were returned to the site of capture within 12 hours.
Growth
Growth rates (GR) in mm/day were estimated for lizards measured at

least twice in the study (104 males and 118 females). They were
99
Chapter 4 GROWTH & SUR VIV AL
calculated as the difference between size (SVL) at first capture

(SVL 1 ) and size at a subsequent capture (SVL2 ), divided by the time
interval between the two captures (in days). If a lizard was
recaptured more than once I used its size difference between the first
and fmal capture. Only captures greater than ten days apart were
considered. These raw growth rates were then transformed into
Specific Growth Rates (SGR) by dividing GR by the average of SVL 1
and SVL2 following Kaufmann ( 1981 ). Linear regression was then
used to fit Logistic, Gompertz and Von Bertalanffy growth models to
the data by using appropriately transformed values of SGR (see Table
4.1) as the dependant variable and the mean of SVL 1 and SVL 2 as the
independent variable. The slope of the linear estimate of the
differential form of each equation (a) describes the rate at which the
growth curve approaches the asymptote (S"') and is called the
characteristic growth rate. Of the three models, the one which

produced the highest value of R2 from a linear regression of the
original data was taken as the model which best fitted the data. The
best regression model was then regarded as a differential equation
and integrated to provide· a growth function (Table 4.1 ). This was
solved to tmd the constant of integration (to) assuming that the SVL
at hatching (31mm; Table 3.1) represents the size at age zero.
Initially, curves for males and females were fitted separately to data
pooled from all years of the study. This was to determine which
growth curve best fitted the data. Then curves were calculated
separately for males and females in each year of the study to test for
significant variation in a and S.,. Analyses of covariance (ANCOVA;
Sokal & Rohlf 1981), with mean SVL as the covariate, were used to
100
test for significant differences between sexes and among years using
the differential forms of the growth curves.
Size distribution of males and females
Size frequency distributions were constructed for all lizards that

could be sexed and that were captured between August and October
(the time at which mating and egg laying occurs; Chapter 3) in each
of the four years of the study. For those lizards that were recaptured
during one of these periods, the size at the first capture for that
period was used. The size frequency distribution for females was
used to construct an expected size frequency distribution for males,
using the appropriate growth curve, and assuming that males and
females have the same age distributions following Stamps et al.
(1994). This approach used a growth curve to find the age of each
female, and then used the male growth curve to find the size that a
male would be at the same age. It assumes that an equal number of
each sex hatch into the population. The use of separate growth
curves for each sex avoids the assumption that females and males are
the same size at a given age (as would be the case if one were to
compare size frequency distributions of male and females directly). A
significant discrepancy between the observed and expected size
distributions may indicate that males and females differ in
survivorship, or in dispersal.
101
Chapter 4 GROWTH & SUR VIV AL
This approach was used to test between the alternative hypotheses

that the SSD in Peninsula dragons is a result ·Of either (1) decreased
survivorship among adult females, or (2) sexual selection for large
body size among males which gives rise to greater mortality among
large males. ~he first hypothesis predicts that there would be an
excess of males in all adult size classes. The second hypothesis
predicts that there would be a deficit of very large males, but not of
small and medium-sized males. The difference between the observed
and expected size frequency distribution of males during the breeding
season in each of the four years was compared using Kolmogorov-
Smirnov tests (dmax, Sokal &Rohlf 1981)
Sul"vivorship
The maximum age attained by each lizard (fx) was estimated to the
nearest year. This was done using the logistic growth equations for
males and females (see above) to estimate age at first capture, and
then from a register of captures (Caughley 1977) for 372 C. fionni at
Midgee Rocks between 1991 and 1994. Lizards were presumed dead
following the last date of capture because they rarely move from the
home ranges they establish during the first few months of life
(Johnston, unpublished data). Thus the absence of a lizard from a
sample is unlikely to result from emigration. The number of lizards
that entered the population during the study was estimated from the
number of clutches produced by females found gravid during the
study, assuming that each gravid female produced a clutch of four
102
eggs (Chapter 3). The primary sex ratio was assumed to be 0.5, so
the estimated number of recruits was divided equally between males
and females in this estimate of survivorship.
Maximum age was expressed as the number of spring breeding

seasons for which an individual lizard was present in the population.
An individual that did not survive to its first breeding season was
assigned a maximum age of zero. Survivorship (/x), age specific
mortality rate (qx), and life expectation (e.,.) were calculated from
these longevity data following Caughley (1977). The Kolmogorov-
Smirnov statistic was used to test the equality of the age distribution
of males and females.
Predation
Two .attempts were made to test the null hypothesis that males and
females suffered equal levels of predation. First, an experiment
measured the attack rate on male and female lizards by a bird of
prey, one of the main predators of dragon lizards (Gibbons &
Lillywhite 1981). Secondly, the frequency of broken tails in lizards
caught in the field was used as an index of predation pressure on
males and females.
103
Predation by birds of prey
Peninsula Dragons were subjected to predation by a single Australian

Kestrel (Falco cenchroides) housed in a 4.0m x 3.5m x 2.5m aviary
which had a cement floor. The two shorter ends and half of the roof
of this enclosure were constructed of corrugated iron. The other
walls of the enclosure were constructed of 35mm wire mesh. Wooden
perches were supplied for the kestrel at each corner of the cage. A
pile of 12 grey concrete blocks of various sizes were placed in the
centre of the enclosure where they were exposed to direct sun for at
least half of the day. To prevent lizards from escaping, a 0.6m high
. sheet of black plastic was taped around the bottom of the cage walls.
Water was available from a bowl placed midway along one of the
short ends of the enclosure.
Ev~ry second day from 12 March to 19 April 199~ two adult

Peninsula Dragons, a male and a female, were released into the pile
of cement blocks in the aviary before 0900hrs. The cage_ was
thoroughly searched after l 700hrs on the same day to determine
whether either, or both, lizards were present. If a lizard was not
found it was assumed to have been eaten by the kestrel. This was
usually confirmed by dissecting pellets produced by the kestrel,
although the kestrel did not produce a pellet after each experiment.
Any lizards remaining at the end of an experiment were removed
from the cage and the kestrel was fed a mouse. The kestrel was left
undisturbed and not fed on the day prior to each predation trial. A
different pair of lizards were used for each trial, but only one kestrel
104
was available. This experiment assumes that this individual kestrel is

representative of kestrels in general, at least in terms of whether it
shows different preferences for male or female Peninsula Dragons.
Trials in which both the male and the female lizard were eaten by the
kestrel on the same day were excluded from analyses because I could
not determine which lizard was consumed first. The remaining data
were analysed using x. 2 (Sokal & Rohlf 1981) to test the null
hypothesis that male and female Peninsula Dragons were consumed at
equal frequencies.
Frequency of broken tails
The frequency of damaged tails among field caught lizards was noted
separately for males and females greater than 50mm SVL when each
individual lizard was first captured. These frequencies were analysed
using x. 2 (Sokal & Rohlf 1981) to test the null hypothesis that adult
male and female Peninsula Dragons exhibited the same proportions· of
tail loss.
RESULTS
Growth
A total of 372 lizards were captured and measured 738 times during
the study. These recaptures provided values of SGR for 104 males
105
and 118 females. The remaining lizards were captured only once, or
were recaptured within ten days of the first capture and were not
included in the analysis.
Linear estimates of the differential forms of all three growth models

were significant (Table 4.2), but the logistic model fitted the data
best for each sex, explaining 23% of variance in SGR in males and
18% in females. Females showed no significant differences among
years for both slopes (F3 •115 = 1.15, P = 0,333) and y-intercepts
(F3 , 115 = 1.04, P = 0.376) of the linear estimates of the logistic
equation. But males showed a significant difference in slopes among
years (F3,100 = 3.26, P = 0.025). Post hoc ANCOVA's demonstrated
that this difference was due to the slope of growth rate against mean
SVL being significantly lower for males during 1992 than in all other
years (F1, 101 = 8. 75, P = 0.004). The slopes were not significantly
different between males and females in any year or for data pooled
across all years (all years pooled FJ,11s = 0.16, P '< 0.0001), but the
y-intercept was larger for males (F3 , 118 = 10.35, P = 0.001).

Peninsula Dragons grow an average of l.3mm/mo (standard deviation
= l.98), and up to 7mrn/mo. The difference in y-intercept between
sexes suggests that males and females differ in asymptotic size. The
logistic equation based on pooled data from all years predicts an
asymptotic SVL of 86. 7mm for males and 78.3mm for females.
106
Size distribution of males and females
The ob served size frequency distribution of male Peninsula Dragons

differed significantly from the size frequency distributions predicted
on the assumption that males and females had the same age
distribution (Figure 4.3). This was true for all years of the study
(1991: dmax = 16, P < 0.001; 1992: dmax = 12, P < 0.001; 1993:
dmax = 20, P < 0.001; 1994: dmax = 6, P < 0.01). The ilifferences
result from a deficit of large males in each year (Figure 4.3). Thus
males and females do not share the same age distribution. Males
appear to live for a shorter time than females. This directly
contradicts the predictions of the hypothesis that the SSD in
Peninsula Dragons results from higher female mortality.
Survivorship
The pattern of longevity of individual Peninsula dragons recorded

over the four year study period shows that males greater than three
years old were rare, relative to females of the same age (Figure 4.4 ).
The oldest known male was five years old at the end of the study,
whereas the oldest female was six years old. Overall there was a
significant difference in the distribution of longevities shown by
males and females (dmax 203, P < 0.001).
107
The survivorship curves for both sexes showed two distinct segments
to the mortality regimes of adult C. fiomii (Table 4.3; Figure 4.5). A
period of low mortality lasted for five years in females but only three
years in males. After this there was a senescent period of high
mortality. Female C. fionni exhibited an age-specific mortality rate
of less than 5 0% per year during the first three years of life.
Thereafter the mortality rate of females increased steadily. The
annual mortality rate of males was 9% higher than females in the first
year of life, but was similar to that of females over the second and
third years of life. After the age of three years the male mortality
rate rose sharply.
The life expectancy for males who survived until their first spring
breeding season was 89% that of females at the same age. By the end
of their third year, males had a life expectancy 68% that of females
of the same age (Table 4.3).
Predation on Males and Females
Eight potential predators were seen incidentally at Midgee Rocks

during the study. Three of these potential predators were observed to
kill and eat Peninsula Dragons. In February 1994 a fox (Vulpes
vulpes) killed three male Peninsula Dragons with radiotransmitters
attached during a study on thermal biology (Chapter 5). In August
and September of 1992 and 1993 a pair of Australian Kestrels (Falco
cenchroides), seen regularly over Midgee Rocks, captured and killed
108
several male Peninsula Dragons that were also being radio-tracked

(Chapter 7). One of these males was found dead over two kilometres
away from its usual home range on top of a large granite boulder,
along with three other juvenile Peninsula Dragons. All four of these
lizards had puncture marks in their skin which were consistent with
having been caught by birds of prey. A chance finding at Midgee
Rocks of the skeleton of an elapid snake (Echiopsis curta) with the
skeleton of a small Peninsula Dragon of unknown sex in its jaws
confirmed that these lizards are also preyed upon by snakes. These
field observations provide anecdotal evidence that males suffer
higher predation than females.
Predation by birds of prey
Of the twenty replicate trials the kestrel consumed both lizards in

five. Ten males and five females were consumed in the remaining
fifteen trials. This tendency for males to be preyed upon more than
females was not significant (x 2 = 1.667, 0.25 > P > 0.10), but the
trend is in the same direction as the field data, ie. for more males to
be taken than females
109
Tail breaks
The tail was recorded as whole or damaged for 25 I lizards first

collected as adults, and which could be assigned to a sex. Nineteen
of 113 males (20%) had damaged tails. Twelve of 138 (9%) had
damaged tails. Although there was a tendency for more males than
females to have damaged tails, this difference was not significant (xZ
= 3.72, 0.10 > P >0.05).
Damaged tails were manifest in a number of ways. Most lizards were

missing various amounts from the end of their tails. Some had
distinct kinks in their tails and irregular scalation, representing scar
tissue, at the point where the tail had been damaged. A few lizards
had partially regrown tails. In these lizards the regrown part of the
tail was covered with very small, granular scales and was grey in
colour, even in the normally brown females.·
DISCUSSION
The results show that male and female Peninsula Dragons have
'
similar rates of growth, but that males achieve a greater asymptotic
size than females. The expected size frequency distribution of males,
constructed on the assumption that females and males have the same
age distribution and only differ in growth pattern, showed significant
differences resulting from a lack of very large males. This suggests
110
that males and females may not have the same age distributions, and
that females tend to live longer than males. An analysis of
survivor.ship confirmed that males suffered consistantly higher annual
mortality than females. This resulted in a significantly younger age
distribution for males. Studies of predation failed to show any
significant differences in the frequency of predation between the
sexes, and there was no significant difference in the proportion of
males and females with damaged tails. This presumably resulted from
unsuccessful predation attempts. However, in both cases there was a
non-significant tendency for males to suffer more than females.
All three equations used to describe growth in Peninsula Dragons

gave a statistically significant fit to the data (Table 4.2). The
equation which explained the greatest amount of variance in growth
rate with respect to body size (ie. had the highest R2 ) was arbitrarily
chosen. Thus the fact that the logistic growth equation, rather than
the Gompertz or Von Bertalanffy equations, is used here should not
be taken as indicating anything abo.ut the intrinsic nature of growth
in Peninsula Dragons, as might normally be suggested by the
acceptance of one formal growth model over another.
Growth in Peninsula Dragons is similar to growth in other reptiles

(Andrews 1982) in that they grow towards an asymptote and mature
well before they achieve the asymptotic size (59% and 68% of
asymptotic size for male and female Peninsula Dragons; Chapter 3).
In common with most other lizards in which males are territorial
(Andrews 1982; .Stamps 1983), Peninsula Dragons exhibit sexual
dimorphism in asymptotic size, with males achieving a larger body
111
size (86. 7mm) than females (78.3mm). Despite this difference in

asymptotic size between the sexes, male and female Peninsula
Dragons do not differ in their characteristic growth rates. Thus males
and females approach their respective asymptotic sizes at the same
rate, but males grow for a longer period than females to achieve a
greater size.
The only other agamid for which a formal growth model has been
published is Japalura swinhonis from Taiwan (Lin & Lu 1982).
Growth in J. swinhonis fitted a Von Bertalanffy growth equation, but
no attempt was made to see if any other growth models gave a better
fit to the data. This species is similar to C. fionni in that the sexes
have similar characteristic growth rates, but males reach a greater
asymptotic size than females.
Peninsula Dragons grow at a similar rate (x = 1. 3mm/mo, SD = 1. 98,

maximum= 7mm/mo) to other long lived species of dragon lizards
such as the closely related C. ornatus (x = 3. lmm/mo, maximum=
7mm/mo; Bradshaw 1986). Japalura swinhonis (Lin & Lu 1982) and
Physignathus /eseurii (Thompson 1993) also have similar average
growth rates. One large, short-lived species of dragon lizard (C.
nucha/is) may grow up to a remarkable 25mm/mo (Bradshaw 1981).
Variation in growth rate between lizards from different populations

raised in common environments suggest that there is a genetic
influence to growth rate (Baverstock 1972; Nierwiarowski 1995).
But it is also clear that environmental characteristics such as food
availability (Schoener & Schoener 1978; Dunham 1980), water
112
availability (Bradshaw 1971; Stamps & Tanaka 1981; Krekorian

1984), the thermal environment (Bradshaw 1971; Sinervo & Adolph
1989) and hormonal factors (Licht 1967) may all affect growth rate.
Perhaps the best example of how the interplay between genetic and
environmental factors may affect the size and age structure in a
population comes from Bradshaw's (1971, 1986) study of C. ornatus,
in Western Australia.
Ctenophorus ornatus (the Ornate Dragon), is very similar to C. fionni

in many respects. Both species have similar body sizes and both
occur in discrete populations on rock outcrops. Ornate Dragons
exhibit considerable variation in growth rate such that some liza:rds
mature and reproduce in their first year, while others do not become
mature until after they are at least one year old and do not breed
until their second or third summer. Bradshaw (1971) refers to these
two groups as fast and slow-growers, respectively, although growth
rate is not bimodally distributed (Bradshaw 1986). Fast and slow
growers represent extremes in a continuum of growth rates which
differ only in the age at which they are able to reproduce. A great
deal of the variation in growth rate seen in C. ornatus can be
explained by climatic variation. Fast growers are frost hardy and
drought sensitive, whereas slow growers are frost sensitive and
drought hardy. Variation in the mean growth rate among populations
of C. ornatus is determined by the frequency of frosts in winter and
of droughts in summer, and corresponds with the predictability of the
frosts a.nd droughts (Haverstock 1972).
113
It is interesting to note that C. ornatus at Bradshaw's ( 1971) study

site show considerably less variation in growth rate (mean =
3. lmm/mo, standard deviation= 0.4; coefficient of variation= 12.9)
than do Peninsula Dragons at Midgee Rocks ( coefficient of variation
= 152.3). This observation suggests that further investigation into
the relationship between climatic variation and growth rate in
Peninsula Dragons may be fruitful. The difference in growth rate
exhibited by male Peninsula Dragons at Midgee Rocks during 1992
compared with the other three years of the study suggests that
growth in this species may be effected by environmental factors.
Neverthless, even though my data may allow a preliminary
investigation of these relations,. such a study is considered beyond
the scope of this thesis.
In addition ,to effecting growth rate, environmental conditions may

also affect asymptotic size (Clutton-Brock, Guiness & Albon 1982;
Ralls & Harvey 1985). For example, female red deer normally
achieve a smaller asymptotic body size than males, but if food is
scarce, asymptotic size is curtailed more strongly in males than in
female (Clutton-Brock, Guiness & Albon 1982). This may also be the
case in Peninsula Dragons because there was a significant difference
for males in the value of a between data collected in 1992 and data
collected in all other years. Despite this variation in growth rate
between years, there were no significant differences in growth rate
between males and females during any single year, or when data from
all years were pooled.
114
Sexual Size Dimorphism
Where differences in body size occur between the sexes, they may
arise due to sexual differences in growth patterns, age-distribution
and/or phenotypic responses to local environmental conditions
(Stamps et al. 1994). The consistency of the relationship between
male and female growth patterns over the four years of this study
suggest that it is unlikely that the SSD in C. fionni is a result of
sexual differences in phenotypic response to environmental variables.
It is more likely that SSD in this species is the result of sexual
differences in growth pattern because males grow to a larger
asymptotic size. The sexual size dimorphism remains apparent
despite higher mortality among large males than among females.
There was a clear difference between the size distribution of male

Peninsula Dragons expected if their age structure is the same as
females(see Stamps 1995), and the observed size distribution 'of
males (Figure 4.3). This difference was consistent over the four years
of the study and, given that sexual differences in growth were used
to construct the predicted size distribution, suggests that the sexes
do differ in their age-distribution (at least in so far as the logistic
growth model accurately describes growth in this species; see
Halliday & Verrell 1988). Assuming that primary sex ratio is 1:1,
such a difference in age structure of a population may result from
males dispersing from the population (and not being replaced) or
from males dying earlier than females.
115
There are several examples of sexual differences in the age

distribution of lizards which result from large males forcing smaller
males to disperse by behaving aggressively toward them (Schoener &
Schoener 1980; Guyer 1988; Andrews & Stamps 1994). In all of
these cases SSD was increased by this behaviour, whether or not the
species involved showed sexual differences in growth pattern. The
pattern of male size distribution at Midgee Rocks was different from
this because there was a deficiency of large males. Dispersal is
unlikely to explain differences in expected and observed size
distribution among male C. fionni because of the limited
opportunities for dispersal. Rock dragons are thought to be closely
tied to suitable patches of rocky habitat (Houston 1974; Gibbons &
Lillywhite 1981). The study population at Midgee Rocks occurs on
an isolated granite outcrop which is separated from other outcrops
by extensive areas of sand dunes. Despite searches of nearby
outcrops throughout this study, only one (male) Peninsula Dragon
that was marked in the study population was located on another rock
outcrop. Furthermore dispersal of young males in response to
a·ggression from older males would result in a deficit of younger
males, which is not a noteworthy feature of Figure 4.3. Dispersal
away from the natal population in Peninsula Dragons appears to be
relatively rare and is unlikely to offer an explanation for the
discrepancy in observed and expected male size distributions.
Increased mortality among larger males, on the other hand, does

appear to offer an explanation which is consistent with the lack of
large male ··Peninsula Dragons at Midgee Rocks. Males may grow to
a larger than optimum size for survival, relative to females, because
116
of the reproductive benefits (Arak 1988), and thereby incur multiple

costs. Larger males may loose condition faster than smaller males
during the breeding season (Wells 1978; Ruby 1984), suggesting that
they suffer higher costs of reproduction. These costs of reproduction
may include the energetic costs of displaying to females and rival
males (Clutton-Brock, Guiness & Alb on 1982; Marler & Moore 1988;
Grahn 1993}, and the risk of injury or death while fighting with rival
males (Sherman & Morton 1984). There may also be more subtle
costs to large size in males. The physiological demands of a longer
period of growth among yearling males may increase male mortality.
Increased energetic costs of further growth among mature males, due
to the longer time males spend engaged in reproductive activities
during the breeding season (Woolbright 1983), may effect future
reproduction.
In addition to the energetic costs of displaying and fighting, males

may also suffer increased mortality due to predation as a result of
their increased conspicuousness. Among frogs, for example, larger
males suffer higher predation than smaller males due to their more
conspicuous calling behaviour (Howard 1981; Ryan 1985). In species
in which males are much larger than females, males may suffer
increased mortality due to predation (Owen-Smith 1993), presumably
as a result of optimal foraging by the predator. There is a counter
suggestion that conspicuous male plumage in some species may
signal that males are 'unprofitable prey' (Baker & Parker 1979;
Gotmark 1992; Olsson 1993a). The idea here is that potential prey
might signal to predators that they are hard to catch, and
117
conspicuousness would make such a vigorous mutant more

memorable and selectively advantageous.
Male Peninsula Dragons have more conspicuous colouration than

females (Chapter 2), use higher perches than females (Chapter 5), are
more active than females (Chapter 5), and are able to run more
swiftly than females (Johnston, unpublished data). One could easily
argue that the increased conspicuousness of males might expose them
to greater predation. Alternatively, it might s_ignal that they are
harder to catch , and therefore reduce their susceptibilty to
predation. All of the trends from field observations, the laboratory
study and the frequency of damaged tails suggest that males are more
susceptible to predation, even though these trends were not
statistically significant.
Broken tails may occur in lizards due to predation (Rand 1954;

Pianka 1967, 1970; Tinkle & Ballinger 1972; Parker & Pianka 1973;
Bellairs & Bryant 1985), but may also occur as a result of damage
during male combat (Vitt, Congdon, Hulse & Platz 1974). However,
agamids lack fracture planes iri their tail vertebrae (Etheridge 1967)
and are not easily damaged during fights (Heideman 1993).
Furthermore, cryptically coloured agamid suffered a lower frequency
of tail breaks than a more colourful species (Heideman 1993),
suggesting that predation, rather than fighting, is the major source of
tail injuries in these lizards.
If females were suffering an increase in mortality due to the costs of

reproduction, one would expect a surfeit of males in all mature size-
118
classes, not of only larger males in Figure 4. 3. The survivorship

schedule constructed from recapture data for Peninsula Dragons at
Midgee Rocks (Figure 4.5), also supports the suggestion that males
(and particularly older males) suffer higher mortality than do
females.
In summary, the pattern of SSD in Peninsula Dragons at Midgee
Rocks, in which males are larger than females, appears to be largely

a result of differences in growth pattern between the sexes. The full
extent of the size dimorphism implied by the larger asymptotic size
of males, appears not to be realised due to high mortality among
large males. This is presumably a result of indirect costs of
reproduction, such as maintaining a territory and gaining mates.
These results suggest that the SSD in Peninsula Dragons at Midgee
Rocks is not a result of proximate factors reducing female body size,

but rather are consistant with the hypothesis that sexual selection
acting on male size may have acted to produce the SSD in this
population.
119
...
Growth model Differential equation Integrated equation Asymptotic si2:e
Logistic SGR=b-aS S = S.,[l +exp-b(t+t0 )J· l S., = bla

Gompertz SGR= b-alnS S = S.,exp[-exp-a(t+to)] S., = expbla
Von Bertalanffy SGR = b-a( 1/S) S = S00 [1-exp-b(t+t9 )] S00 = b/a
....N TABLE 4.1: Equations for the three growth curves used to descn"be growth in male and female
0
Ctenophorus fionni. The first column gives the name of each model. The second column gives the
differential form of the equations showing the relationship between specific growth rate (SGR) and si2:e
(S). The relationship between SGR and S should be linear if these variables are appropriately transformed.
The third column gives the more commonly used integrated form of the equations, which gives si2:e as a
function of time (t). The fourth column gives the equation for calculating the asymptote (S.,) of each
. .
growth equation. The parameters a and b are the slope and y-intercept, respectively, of the linear estimate
of the differential forms of the equations, and to is the constant of integration (Kanffinan 1981).
Males (N =104) Females (N =ll8)
Model Equation · Equation R2
Logistic SOR= 0.00424 • 0.000049SVL 0.226* GR= 0.00428 - 0.000055SVL 0.176*
Gompertz SOR= 0.01398 • 0.003ll8InSVL 0.197* GR= 0.01536 • 0.003515InSVL 0.171*
--
N
Von Bertalanffy SGR = -0.001996 + 0.18629(1/SVL) 0.163* GR= -0.002643 + 0.2136(1/SVL) 0.158*
TABLE 4.2: Comparison of the differential forms of the logistic, Gompertz and Von Bertalanffy growth models
calculated fur male and female Ctenophorus fionni at Midgee Rocks based on data collected from 1991 to I 994. N =
sample size; R 2 = coefficient of determination; probabilities show the significance of the pearson correlation coefficients
for each equation.* P<0.0001.
Age Males Females
(years)
X fx qx ix ex jx qx Ix ex
0 171 0.34 1.00 2.28 203 0.25 1.00 2.67
1 112 0.36 0.66 1.94 153 0.33 0.75 2.23
2 72 0.57 0.42 1.48 l.02 0.48 0.50 1.85
3 31 0.89 0.18 1.11 52 0.54 0.26 1.63
....N 4 4 0.75 0.02 LOO 25 0.66 0.12 1.37
N
5 I 1.00 0.00* 0.00 9 0.91 0.04 1.00
6 0 l 1.00 0.00* 0.00
TABLE 4.3: Static life-table for male and female Ctenophorus

/ionni at Midgee Rocks during the period 1991-1994./x = frequency
of lizards living to age x; Ix= survivorship; qx = mortality rate; ex=
expectation of life in years. * indicates values that are too small to
show due to rounding-up to two decimal places.
00<1
•
• • •••
0
.003 0
-
0
w 0 0.
i
:c
.002
.--- -------,.• 0
0
0
• •
••. •--·--~!)-
Co
!
(!)
.001
.000
o•
• od'
••
~ 0
0
0 --
frl •.001~ • 0 • •
0. •
•.002•I
(I)
0
•
0
•.003
40 50 60 70 80 so
MEANSVL
Figure 4.1: Specific Growth Rate in Snout-Vent Length for

Ctenophorus fionni at Midgee Rocks as functions of size in
males(•, solid lines) and females (0, broken lines) based on
recapture intervals> 10 days.
123
100
90
I
._,
80
~
~ 70
~ 60
~
50
~ ----Male
5
0
40. ---Female
~ 30
0 1 2 3 4 5
AGE(yean)
Figure 4.2: Logistic growth curves for Snout-Vent Length in

Ctenophorus fionni at Midgee Rocks.
124
30 15
1991 1992
20 10
10 s
0 0
-10 -S
-20 -10
ffiiXl .JO -IS

20 40 100 20 40 80 100
::E 30 7.5
::> 1994
z 199.3
9
......:i:.
:IO
.9, l.
..
s
......... ..
: !
:. :.
10
' I
2.5
.
: 1
0
\ 0
\
-10 -:1.S
-20 .5
20 40 60 80 100 20 40 60 80 100
Figure 4.3: Observed size distributions (•)of adult male and female
Ctenophorus fionni at Midgee Rocks over four years along with the
expected size distribution of males (O). In each panel the frequencies
of males are shown above the horizontal line, and the frequency of
females are shown below the horizontal line. The expected
distribution of males was calculated using the size distributions of
females in each sample on the assumption that males and females had
the same age distribution, but differed in growth pattern (see Stamps
et al. 1994).
125
40
30
--
~
._,
0
>
V
20
z'1;l
;;;i
g 10
~
~
0
0 I 2 3 4 s 6 7
LONGEVITY (years)
FIGURE 4.4: Maximum age recorded for 372 of male (D) and
female (•} Ctenophorus fionni at Midgee Rocks based on a mark-
recapture study between 1991 and 1994.
126
0
Female
~ -1
~ Male
I
rlJ
e.c
0
..;i
-2
-3
-4
-5
0 l 2 3 4 5 6 7
AGE (ye~rs)
Figure 4.5: Static survivorship curves for male (0) and female(•)
Ctenophorusfionni at Midgee Rocks during 1991-1994.
''
127
CHAPTERS
ECOLOGICAL DIFFERENCES BETWEEN

MALES AND FEMALES
Where there are ecological differences between males and females of a species it is
often difficult to determine whether they are an indirect consequence of sexual
dimorphism, or whether the ecological differences led to the evolution of the
sexual dimorphism (Hedrick & Temeles 1989). This is a crucial issue in assessing
the relative importance offecun_dity, mortality and sexual selection in the evolution
of sexual dimorphism, because many species show ecological differences between
the sexes (see Shine 1989 for a review).
Theoretical and empirical studies suggest that there are two ways in which
ecological differences between the sexes may be related to sexual dimorphism
(Slatkin 1984). First, each sex may have a separate adaptive optimum to which it
may evolve. This may be the case if males and females have different energetic
requirements ( eg. Sandell 1989). For example, a pre-existing sexual dimorphism
may give rise to behavioural and ecological differences between the sexes. These
Chapter 5 NICHE DIVERGENCE
ecological differences may then amplify the dimorphism, but the ecological
differences are fundamentally a consequence of the sexual dimorphism. This is the
'social roles' hypothesis. Secondly, there may be a single adaptive optimum for
both males and females, and the sexes may force each other away from this
optimum in a process ofintraspecific competitive exclusion. Here the ecological
differences between the sexes cause the evolution of sexual dimorphism. This is the
'niche-divergence' hypothesis.
The niche-divergence hypothesis assumes that (1) the sexes should differ in their
use of resources, and predicts that (2) there should be a correlation between the
dimorphic trait(s) and the resource used. This hypothesis also predicts that (3) the
evolution of the sexual dimorphism should be accompanied phylogenetically by a
shift in resource use. In contrast the social roles hypothesis assumes that ( 1) there
need be no difference in resource use between the sexes, but if there is it predicts
that (2) there should be no correlation between the dimorphic traits and the
resources used, and (3) the evolution of sexual dimorphism should be accompanied
by a shift toward either a relationship between body size and fecundity in females,
intrasexual aggression, or mate choice. In the absence of any published phylogeny
for the genus Ctenophorus, or sufficient data on resource use by most
Ctenophorus species it is not presently possi'ble to test predictions about the order
in which traits have evolved. But it is possible to test the other assumptions and
predictions of the two hypotheses.
Reptiles partition resources along at least three niche axes: food, space and time
(Pianka 1973; Toft 1985). Whether or not reptiles partition a fourth niche axis,
heat, has been controversial. Some authors view the way in which the thermal
environment is used as a simple consequence of the way in which space and time
are used (eg. Roughgarden, Porter & Heckel 1981). Others view heat as a resource
129
in its own right. The latter group argue that animal$ compete for, and partition
thermal resources, and that this process may contribute directly to fitness
(Maguuson, Crowder & Medvick 1979). I treat heat as a resource in its own right
for two reasons. First, some obvious differences in activity patterns and habitat use
are not accompanied by corresponding differences in thermal biology in some
organisms. For example females of the aquatic, tropical snake Acrochordus
arafurae forage for large fish in deep water and males forage for small fish in
shallow water (Shine 1986) without any siguificant differences in field body
temperatures (Shine & Lambeck 1985). Similarly, at an interspecific level, several
sympatric species of Camllean Ano/is have different activity patterns and use
microhabitats with different thermal characteristics, but overlap broadly in the body
temperatures they achieve in the field (Hertz 1992). The second reason is that it is
important to identify ecological differences between the sexes for the purposes of
this chapter. It is, therefore, better to include the thermal niches axis, rather than
risk overlooking an important difference between the sexes.
This chapter looks at whether there are ecological differences between males and
females along each of the four niche axes. The chapter is divided into four sections,
each of which deals with one niche axis. In each section I ask the following
questions, which stem from assumptions and predictions of the social roles and
niche divergence hypotheses: (1) Is there a difference in the way that males and
females utilise resources along this niche axis? Where differences are found an
attempt is made to determine whether the differences may have given rise to
sexually dimoi:phic traits in Ctenophorus ftonni, or whether they may be artefacts
of the dimoi:phism by asking: (2) Does the difference in resource use represent an
obligate difference (ie. a difference in their fundamental niches), or a facultative
difference (ie. a difference in their realised niches)? and (3) Is there a correlation
between any dimoi:phic trait and the way in which resources are utilised?
130
DIET
Males and females in several vertebrate groups may specialise on different prey
types, with corresponding differences in relative size and shape of the head, jaws or
beak (Selander 1966; Schoener 1967; Petrov 1956). This seems to be especially
tme among gape-limited predators, possi"bly because consumption of food items of
different sizes may give rise to an evolutionary response in jaw size or shape (Shine
1989). The correlation between relative head size and prey size in many reptiles
supports the interpretation that they are gape-limited (De Marco, Drenner &
Ferguson 1985; Shine 1991).
Differences in prey sizes or types between the sexes have been documented in
several species of lizards. In most cases this has been explained in terms of the
social roles hypothesis, perhaps because there is 'ample evidence that male lizards
compete for access to females (Trivers 1976; Ruby 1981, 1984; Hews 1990).
However, niche-divergence has been invoked to explain some cases of extreme
dietary differences between males and females in some species (eg. Schoener 1967;
Perez-Mellado & De La Riva 1993; Van Sluys 1993).
In Ctenophorus fionni the heads of males and females differ in shape as well as size
(see Chapter 2), suggesting that the niche-divergence hypothesis may be relevant.
Thus I report the diet of Peninsula Dragons in the field to determine whether males
and females consume different sizes or types ofprey.
131
Methods
. This study is based on 120 Ctenophorus ftonni from Iron Duke in the Middleback
Ranges (Figure 1.2). This collection was made by the author just prior to Iron
Duke being cleared for an extensive open cut mine and is lodged in the South
Australian Museum. Iron Duke was visited once in most months between April
198S and April 1986. During each visit, liurds (N =4-14) were collected by hand.
and killed by freezing within 12 hours of capture. They were subsequently fixed
with a 10% formalin solution and then stored in 70% alcohol These specimens
also form the basis for a study of reproductive cycles in C. fionni (Chapter 3).
Snout-vent length (SVL) was measured and the stomach removed from each Hurd
in 1991. Only the contents of the stomach were counted; those in the posterior gut
were omitted to prevent a bias towards less easily digested items. All food items
were identified to the level of order. The length of the largest food item in each
stomach was measured to the nearest 0.1mm using a vernier calliper when the
items were intact. If the largest food item was not intact, its length was estimated
to the nearest 1mm with reference to voucher specimens collected in the
Middleback Ranges. A Hurd found in one stomach was excluded from
I
measurements of maximum prey siz.e because it consisted only of skin fragments

and its siz.e could not be estimated with any accuracy.
Raw counts were used to calculate the relative numerical importance of each prey
type, expressed as a proportion of the total number of food items, and the
proportion of stomachs containing each prey type. A discriminant function analysis
was used to test for seasonal differences based on raw counts of stomach contents.
For this analysis the liurds whose stomachs contained food were divided into four
groups: males (N =27) and females (N = 19) collected in summer (August-
132
February) and males (N = 26) and females (N = 17) collected in winter (March-
July). Kolmogorov-Smimov tests were used to compare the distributions of the
average number of items in each food type per stomach between sexes. Linear
regressions were used to descnbe the relationship between maximum prey size and
SVL for each sex. An analysis of covariance was used to test whether these two
regressions were different
Results
Eighty nine of the 120 lizards dissected contained 1133 prey items in their
stomachs (Table 5.1). The stomachs of the remaining 31 lizards were empty,
although food was usually present in their intestine.
Hymenoptera (mostly ants) were by fur the most important prey type for both male
and female Ctenophorus flonni, both in terms of the number of stomachs
containing them and the proportion they represent of the total number of food
items. Beetles were the second most important food type consumed. Orthopteraus ·
accounted for less than five percent of the total number of food items consumed by
both sexes, but were important because they were present in over 40% of stomachs
and were among the larger food items consumed. Arachnids, blattodeaus, diptera,
hemiptera, lepidoptera, mautodeans, neuroptera and vertebrates combined made up
less than 15 % of the total food items consumed (see Table 5.1 ). The single record
of a vertebrate was based on a piece of Peninsula Dragon skin in the stomach of an
adult male collected on 7 November 1985. Cannibalism was also obseIVed

incidentally at Secret Rocks (33°12'S, 135°51 'E) in February 1992 when an adult
male was caught with a recently killed hatchling (SVL = 32 mm) in his gullet.
133
The discriminant function analysis failed to distinguish males collected in winter,

females collected in winter, males collected in summer and females collected in
summer. None of the three discriminant functions derived from the analysis were
significant (P's= 0.43-0.99) and only 52% of cases were classified correctly. When
the data were pooled with respect to season the frequency distributions of the
average numbers of each prey type did not differ significantly between the sexes
(Kolmogorov-Smirnov Z = 0.6396, P = 0.808; Figure 5.1).
Males and females consumed prey of similar maximum sizes (males: x = 11.0 mm,
SD=6.6 mm; females: x= 11.2mm, SD= 5.9 mm;Mann-Whitney U= 0.411,P=
0.681). The slopes of maximum prey size on body size did not differ between the
sexes (F1,86 = 0.04, P = 0.85) and were not significantly different from zero for
either sex (t-tests: males:= 0.596, P = 0.55; females: 0.830, P > 0.41; Figure 5.2).
Discussion
Dietary studies have been conducted on a number of Australian dragon lizards

(Pianka 1986), but none have addressed the question of whether males and females
differ in diet. This study reports the diet of Cterwphorus fionni for the first time
and demonstrates that males and females share very similar diets. It also shows that
there is no relationship between maximum prey size and lizard body size.
Consequently there is no evidence for dietary differences between the sexes or that
niche divergence has played any part in the evolution of the sexual differences in
relative head size or head shape in Peninsula Dragons.
134
The lack of data on types and abundances of prey types available to the lizards
used in this study prevents an assessment of how selective Peninsula Dragons are
in their diet, but this is unimportant to the aim of this chapter. A demonstration that
males and females feed on similar types of prey requires only a comparison
between individuals ofboth sexes that have had the same prey available to them
This is likely to be the case here because the liz.ards used in this study were
collected at the same place and at the same times.
Peninsula Dragons are predominantly insectivorous and feed primarily on ants and
beetles. However a wide variety of other arthropods are also consumed. Ants are
an important component of the diet of dragon lizards, frequently making up over
20% of the girt contents by volume ( Harris 1964; Pianka 1986; Robson &
Lambert 1980; Al-Johany 1995; Znari & El Monden in press). The abuodance of
ants in the diet of these liz.ards may be explained by the abuodance of ants in xeric
environments (Pianka 1986) or by their high energetic value (Schoener 1971;
Robson & Lambert 1980). The absence ofisopterans from the stomach contents of
C. ftonni is noteworthy because they are often very abuodant in the stomachs of
other desert dragon lizards (Pianka 1986; Znari & El Monden in press). Similarly
the complete absence of plants in C. jionni is unusual because several other dragon
liz.ards may consume large amounts of plant material (Harris 1964; Al-Johany
1995).
The diet of C. fi0!1Tli is consistent with a sit-and-wait mode of foraging in that it

consists mostly ofmobile, terrestrial or flying prey such as ants, beetles, and
orthopterans (Regal 1983). The diet of lizards which are active foragers is usually
composed of more sedentary terrestrial and cryptic prey (Regal 1983). However,
the presence of'lerps' (scales ofCoccoidean Hemiptera) in the stomachs of two
Peninsula Dragons suggests that some individuals do a certain amount of active
135
foraging. Flexibility in foraging tactics has been reported in several other lizards
(Pietmszki 1986).
SPACE
Many iguanian lizards are sit-and-wait insectivores which use perches to survey
their home ranges for food, and for potential mates or competitors. Arboreal
species may use the branches of trees or shrubs as perches and terrestrial species
may use rocks or mounds of soil as perches. Larger individuals often use higher
perches than smaller ones (Rand 1967; Schoener 1967; Lin & Lu 1982) and in
species which show sexual size dimorphism the 1argennales use higher perches
than the smaller females (Rand 1967; Schoener 1967). One proximate explanation
for thi$ pattern of perch use comes from optimal foraging theory (Schoener 1967).
Large lizards may use high perches to survey a greater area for less common, large
food items (Schoener 1967; Rummel & Roughgarden 1985).
In some lizards seasonal changes in perch heights chosen by males and females
correlate with seasonal differences in reproductive and social behaviour. This has
been used to support the importance of sex differences in social roles as a
determinant of perch height (Pounds & Jackson 1983). However, seasonal patterns
in the use of perches may also reflect seasonal differences in the abundance of
other resources, such as food (Pounds & Jackson 1983; Zucker 1986). Thus,
seasonal differences in perch use may also be explained by niche-divergence. An
experimental approach is required to test between these alternative explanations. In
the iguanid Urosaurus omatus, males used higher perches than females even when
availability of other resources (food) was kept constant (Zucker 1986), lending
136
some support to the social roles hypothesis. But a more explicit test of the
importance of social factors in the choice of perch height requires manipulation of
the social environment to see if the sexes respond differently in different social
milieux.
In species which show sexual differences in perch height the social roles hypothesis
predicts that (1) there would be no difference in perch height chosen by each sex
when they are housed alone, and (2) at least one sex would respond differently
when alone than in the presence of member of the opposite sex. Alternatively, if
the sexual difference in perch height represents a fundamental niche difference
between the sexes there should be (1) a consistent difference in perch height
between the sexes even when housed alone, and (2) no difference due to the
presence of a member of the opposite sex. Being able to distingnish between these
hypotheses relies on the assumption that different social roles have not given rise to
obligate differences in perching behaviour between the sexes in Peninsula Dragons.
This is unlikely because other lizards are known to alter their choice of perch
height with season (Pounds & Jackson 1983; Zucker 1986) and their level of
hunger (Stamps 1977).
Peninsula Dragons are similar to other iguanians in that they utilise perches to
survey their home ranges. Given the size difference between the sexes in this
species (Chapter 2), the literature would suggest that males and females might
utilise rocks of different heights as perches. I use field data and an experimental
manipulation of the social environment to answer the following questions: (1) Do
the sexes differ in their use of perches of different heights in the field? (2) Is there
a relationship between perch height and body size in C. fionni?, and (3) Do any
differences in the use of perches change with the presence or absence ofa member
of the opposite sex?
137
Methods
Field Observations
Field data were collected to determine whether the choice ofperch height was
influenced by body size or sex, and whether any influence of sex might vary with
season. Lizards might alter their choice 'of perch height in the breeding and non-
breeding seasons if social interactions are important determinants of perch choice.
While I cannot discount the possx'bllity that food availability changes seasonally as
well, a seasonal perch height shift would imply that social roles may be important.
Data were collected from a population of C. ftonni at Midgee Rocks (Figure 1.2)
between August 1991 and April 1993. Data on perch utilisation were obtained by
walking across the study area throughout the day and noting the position of 337
perching lizards before they were disturbed. Each lizard was then captured and its
SVL, sex and the perch height were recorded. Perch height was measured as the
vertical distance between the position of the lizard on a rock and the closest level
area of soil. Determining perch height was usually a simple matter because the
study site consists of numerous isolated granite boulders separated by flat areas of
soil. Care was taken so that search time was evenly spread throughout the day.
Breeding activity at Midgee Rocks occurs between August and November. Data
taken in these months (N = 183) were considered to have been within the breeding
season and data collected from January to August (N = 154) were considered to be
in the non-breeding season.
Mann-Whitney U tests were used to test for the effects of sex and season (breeding
vs non-breeding season) on perch height, because the data were not normally
I38
distributed. Speannan's rank correlation (r,) was used to test for any relationship
between body size and perch height.
Experiment
Nme male and nine female Peninsula Dragons were collected from either Midgee
Rocks or the Middleback Ranges and were held in outside pens for between six
months and two years prior to the experiment. All lizards were aduh and more than
two years old.
The experiment was conducted between 19 March and 22 April 1993 and 22
March and 6 June 1994. Both of these periods were outside of the breeding
season. This should not influence any conclusions reached because the field data
suggested that season had no influence on perching behaviour (see results). Two
outdoor enclosures were constructed of galvanised sheet metal at the Flinders
University campus in Adelaide (34°56'8, 138°36'E). This site has a similar climate
to the Midgee Rocks area. Each enclosure measured 3.0mx 1.5m, with 0.7mhigh
walls and was covered with wire mesh to keep predators out. Concrete blocks
were used to construct five perches between 0.05m and 0. 7m high (Figure 5.3).
The experiment consisted of nine replicates of four treatments: (1) male alone, (2)
female alone, (3) the same male as in treatment one with a female placed in his
enclosure, and (4) the same female as in treatment two with a male placed in her
enclosure. Each lizard was used in one experimental sequence of two treatments
only. Liz.ards were placed in an enclosure under the experimental conditions at
least 24-72 hours prior to recording behaviour for each treatment. This period
139
depended on weather conditions because lizards were only active on fine days
when the air temperatnres exceeded 25°C. Time lapse video was used to record
the height of each lizard on the concrete blocks every ten minutes between 0800
hours and 1900 hours local time. Perch heights were only recorded from the
experimental subjects, ie. no data were collected from the lizards that were
introduced into the enclosures for treatments three and four. Treatments were
rotated between the two enclosures so that an animal under one experimental
treatment could be acclimating while data were being collected from an animal in
the other cage. Experimental treatments were randomised between the pens. Three
replicates of the experiment were run in 1993 and another six replicates were run in
1994. There were no significant differences between the replicates run in 1993 and
1994, so the data from both years were pooled for all subsequent analyses.
Lizards were fed daily on crickets, mealworms and chopped fruit placed on the
ground at least one metre from the base of the concrete blocks. No food was
provided while experiments were in progress, although naturally occurring food
(ants and dipterans) and escaped crickets were available in the enclosures at those
times. Not all food was consumed before or after trials, so all of the lizards used
were assumed to be satiated during the experiment. Successful mating, egg laying
and incubation occurred in holding enclosures of the same construction as the
experimental enclosures during both years of the study, indicating that the lizards
were healthy and behaving normally.
Since the number of perch heights recorded for different individual lizards varied
between 6 and 66, mean perch heights were calculated for each experimental
treatment to ensure an equal contnlrution of each replicate to the results. The mean
perch heights were analysed using a two-way ANOVA to test the effect of sex and
140
Chapter S NICHE DIVERGENCE
presence or absence of a member of the opposite sex. Scheffe's range test was used
to test the significance ofpairwise comparisons.
Results
Field observations
At Midgee Rocks Peninsula Dragons typically perched on granite boulders during

daylight hours when air temperatures exceeded approximately 20 C. Perch height
was not correlated with snout-vent length for all lizards combined (r, = 0.104, P =
0.22) or when the sexes were analysed separately (males: r, =0.019, P = 0.87,
females: r, = -0.185, P = 0.20).
During the nonhreeding season average perch height for males was 0.55m (SD=
0.52) and for females was 0.43m (SD = 0.36). During the breeding season average
perch height for males was 0.65m (SD= 0.60) and for females was 0.49m (SD=
0.50). Neither sex showed a significant difference in perch height between the
breeding and nonbreeding seasons (Males: U= 4584,P = 0.143; Females: U=
1975, P = 0.714). When samples for each season were pooled within each sex,
there was a significant differences in perch height between the sexes; males perched
higher than females (Mann-Whitney U = 15548.5, P = 0.02) (Figure 5.4).
141
Chapter 5 NICHB DIVERGENCE
Experiment
Male and female C. fionni chose similar perch heights when housed alone (females:
.
x = 33.8cm, SD= 12.43; male: x =32.4cm, SD= 14,27) (Figure 5.5). Males chose
higher perches when a female was present (x = 38.8, SD= 10.36) and females
chose lower perches when a male was present (x =26.4, SD= 9.18) (Figure 5.5).
The ANOVA showed a significant interaction between sex and treatment (F1,32 =
4.238, P < 0.048) and a significant effect due to sex (F1.l2 = S. 718, P = 0.023), but
no significant effect of treatment on perch height (F1,32 = 0.028, P = 0.868). An a
posteriori range test confirmed that the significant interaction was entirely due to
the opposite reactions of males and females to the presence of a member ofthe
opposite sex (Scheffe, P = 0.033).
Discussion
The field data show that male Peninsula Dragons chose higher perches than.
females and that neither sex showed any seasonal changes in their choice of
perches. These observations,together with the finding that male and female C.
fionni chose similar perch heights when housed in isolation but different perch
heights when they were housed togethe'lsupports the hwothesis that different
social roles of the sexes explains the sexual difference in perch height observed in
this species. While previous studies have shown that perch heights chosen by male
and female lizards vary with season and that these seasonal differences persist
when the availability of other resources, such as food, are held constant (Pounds·&
142
Jackson 1983; Zucker 1986), this is the first report of a change in perch height
resulting from direct manipulation of the social environment of lizards.
Peninsula Dragons appeared to use perches as vantage points to survey their home
ranges for potential social interactions (ie. territorial disputes and courtship
opportunities) and to scan for potential predators and prey. But vigilance alone
does not explain the pattern of perch use in C.fionni because (1) felllllles choose
lower perches than males in the field even though both sexes appear to be
territorial (Chapter 6), and (2) in the experimental enclosure males chose lower
perches when alone than in the presence of a female. These observations llllly be
explained by the disadvantages of perching for each sex.
The choice of perch height by lllllles is more probably a compromise between (1)
the advantages ofvigilance for potential llllltes, food items, or competing males,
and (2) the disadvantages of increased exposure, such as increased mortality due to
predation. This may explain why permanently territorial male Urosaurus ornatus
choose high perches at all times of the year, whereas females choose higher
perches after the breeding season (Zucker 1986) when they establish territories
(Dunham 1980). Male Japalura swinhonis use progressively higher perches as they
grow and obtain territories (Lin & Lu 1982) suggesting that perch height plays a
role in territory acquisition and maintenance in this species too.
One disadvantage ofhigh perches for female lizards comes from incessant
courtship by males during the breeding season. To avoid harassment from males,
females may choose lower perches where they are less visible to males (Andrews
1971; Fleming & Hooker 1975; Zucker 1986). During this study male C. fionni
were seen chasing females from high perches onto lower ones at all times of the
year, but the frequency and intensity of this behaviour was higher during the
143
breeding season (Johnston, unpublished data). Females use perches less often than
males (see sample sizes in Figure 5.4), and they usually use lower rocks.
Harassment by males may also explain why females used lower perches in the
presence of males during the experiment.
Predation is another disadvantage of perching which may be particularly relevant

for males. Where aerial predation is important lizards should choose low perches to
avoid exposure to predators. Male C. ftonni are preyed upon by raptors (Gi'bbons
& Lillywhite 1981; personal observation) and this may explain the lower survival
rate among males (Chapter 4). Suscepti'bility to predation may explain why males
chose lower perches when alone than when a female was present. Vigilance over
potential mates and against potential competitors may counteract the increased risk
ofpredation due to choosing a high perch.
The responses of males and females in the experiment may be the result of size
related dominance. Since females are smaller than males they would be expected to
perch lower in the presence ofa larger (male) lizard, whereas males might be
expected to perch higher by way of asserting their dominance. This hypothesis
could be tested with an experiment of similar design to the one reported here. One
could measure perch height preference of a liz,ard before introducing another liz,ard
of the same sex. The results would have to be analysed with due regard to the size
of the lizards used, even though perch height was not related to body size in the
field. This is because relative body size is an important determinant of dominance
in many lizards (Carothers 1984; Tokarz 1985; Olsson 1992).
144
Chapters NICHE DIVERGENCE
TEMPERATURE
Body size, shape and colour are important determinants ofheat flux in reptiles
(Porter, Mitchell, Beckman & DeWitt 1973; Porter & James 1979; Tracy 1982).
Colour is an.important determinant ofheat absorption from solar radiation because
about half of the energy in sunlight is in the visil>le spectrum (Norris 1967). Many
liz.ards can alter body colour so that they are darker during cooler periods of the
day than during hotter periods (Bradshaw & Main 1968; Heatwole 1970; Cogger
1974). These colour changes have been shown to influence heat absorption (Norris
1967; Porter & Norris 1969; Bradshaw & Rice 1981). The effect of colour in
lizards is mediated by the dermal vasculature which transports heat through the
body after it has been absorbed by the skin (Bradshaw & Rice 1981). Body size
and shape affect heat flux in that they determine the ratio of surface area to
volume. No studies appear to have explicitly examined the causal relationships
between sexually dimorphic body size and colour and sex differences in thermal
biology in liz.ards.
Many sexually dimorphic lizard species show differences in thermal biology

between the sexes (eg. Heatwole 1970; Patterson & Davies 1978; Perez-Mellado
& De La Riva 1993). The extent of these differences may vary greatly. They may
be quite subtle. For example male and female Ctenophorus nucha/is overlap
broadly in body temperature in the field, but males have greater variance in
temperatures than females (Heatwole 1970). The differences may also be quite
marked. Female Tropidurus melanopleurus are thennoconformers, and appear not
to thennoregulate, whereas males thennoregulate actively (Perez-Mellado & De
La Riva 1993). Differences in body temperature in animals may reflect their
145
particular thermal requirements or may be due to constraints imposed by

differences in the thermal environment of the animals.
Adult male C. fionni are larger and considerably darker than females. This species
does not exhibit the temporal variation in colour that many other dragon lizards do
(Sharrad 1969). Males absorb 70% more energy than females in the visible
spectrum (Chapter 2; Figure 2.12). Males are territorial and defend large territories
which overlap with a number of smaller female home ranges (Chapter 6). The
home ranges of both males and females usually encompass several granite boulders
and a number of crevices suitable for use as refuges from extreme climatic
conditions or predators.
In this section I ask the following questions: (I) Do male and female Peninsula
Dragons differ in their thermal ecology in the field? (2) Do any differences field
body temperatures reflect differences in preferred body temperatures between the
sexes? and (3) Do any differences between the sexes in body size and colour affect
the rate at which they gain heat?
Methods
Field Body Temperature
Field work was conducted at Midgee Rocks. A description of this study site is
given in Chapter I (see Figure 1.3). Body temperatures (Tb) were measured by
telemetry in February and November 1994. These months were chosen to allow a
comparison between the nonbreeding and the breeding seasons, respectively.
Temperature sensitive transmitters (Tidey, New South Wales) were taped to the
146
side of the base of the tail and a thermocouple inserted l 5-20mm into _the intestine ·
via the cloaca. Transmitters were placed on five males and five females at the
beginning of each study period, but data could not be collected from all of these
due to predation and transmitter failure (see Table 5.2 for final sample sizes). Body
temperatures were measured hourly overnight between 2000h and 0600h and half-
hourly during the day between 0600h and 2000h. Only one gravid female was
found at Midgee Rocks in 1994 (Chapter 3), so no valid comparison could be
made between gravid and nongravid females in the field. The transmitters were
calibrated two days before the study was initiated and calibrations were rechecked
after the transmitters were recovered.
Environmental temperatures (Te) were recorded with a further two temperature
sensitive transmitters in February and with copper-constantan thermocouples in
November. Te was measured on the exposed surface of a 2.0m high granite
boulder (Tr), and in an open crevice (Tc) in the same boulder. These two sites
were 1.2m apart. Both transmitters and thermocouples were held to the rock with
white masking tape, to avoid dislodgment and to reduce errors due to radiation
(Tracy & Christian 1986). Exposed rock surfaces and deep crevices such as these
represent the most extreme thermal environments at the study site. All other Te' s
would have fallen between these extreme measurements and only radiative heat
exchange would allow Tbs outside of the temperature range they define.
Lizards were only active during the day, so only data collected between 0900h and
2000h were used in the analyses. Descriptive statistics and linear regression were
used to explore the relationship between Tb and Te's for males and females, during
and after the breeding season following Huey & Slatkin (1976). A regression slope
of zero indicates perfect thermoregulation; a slope of one indicates perfect thermal
conformance. Nonparametric analyses were used to test for differences in means
147
and Fmax tests were used to compare variances. Analyses of covariance were used
to test for differences between the slopes of the regression lines of Tb on Te (Sokal
&Rohlfl981).
' Measuring the thermal niche
Ecological constraints can influence thermoregulation in the field. Body
temperatures unbiased by these constraints were measured by allowing post-
absorptive lizards to move about freely in a photothermal gradient. Twenty seven
lizards were run once each in the gradient during July and October 1992. Each
lizard was run in the gradient once, so the measurements for each season are
independent. Experiments were run at these times of the year to investigate
seasonal variation in the fundamental thermal niche. Several lizards of each sex ( see
Table 5.3 for sample sizes) were run in the gradient in each of the two months.
Gravid females were only available in October ( chapter 3) and were analysed
separately from non-gravid females. Body temperatures were recorded every ten
minutes using copper-constantan thermocouples placed 15-20 mm into the
intestine of each lizard via the cloaca. Measurements were taken over 24 hours for
each individual lizard.
The photothermal gradient consisted of a 2. Omx2. Omx0.5m enclosure placed into a
controlled temperature room. The room was held at 5"c and 60% relative
humidity. Flourescent lights on the ceiling of the.room provided a lighting regime
which reflected the natural photoperiod at the time of each experiment (lights on
0800-1700 in July and 0630-2030 in October). Heat was supplied by two 150W
heat lamps at one end of the enclosure. One lamp was placed 0.4m above the floor
148
of the cage and directed straight down. The second lamp was placed 0.8m above
the floor and directed at 45 toward the centre of the enclosure. This arrangement
oflights provided a temperature gradient that ranged from 50 C to 5 C, as
-indicated by the floor temperatures. The relationship between light and heat in
photo-thennal gradients has been discussed by Sievert & Hutchinson ( 1989).
Although the separate effects of heat and light were not determined here, I
considered that a photothermal gradient was appropriate for determining the set
point range, because Peninsula Dragons are heliothennic.
The set-point range (Tset range) (Berk & Heath 1975) was used as a measure of
thethennalniche, following Magnuson, Crowder &Medvick (1979). The Tset
range is the range of temperatures between the temperature at which lizards begin
thermoregulating to increase Tb (lower Tset) and the temperature at which lizards
thermoregulate to decrease Tb. This measure of the thennal niche was chosen
because it has a sound physiological basis (Cabanac, Hammel & Hardy 1967; Firth
& Turner 1982). The mean Tb chosen is of uncertain biological importance and
tends to vary enormously with different conditions of acclimation (Firth & Turner
1982).
Lower and upper Tsets are normally measured using a shuttlebox (Berk & Heath
1975), but this apparatus was not available for this study. Instead the Tsets were
estimated from Tbs measured in the temperature gradient, following Hertz, Huey
& Stevenson ( 1993). Lower Tset was estimated by the 25th percentile and upper
Tset by the 75th percentile from the frequency distribution of Tbs measured for
each lizar.d. A mean upper and lower Tset was calculated for males and for females
in each season from these individual measurements. Comparisons of Tse ts were
made using Kruskall-Wallis nonparametric oneway analyses of variance (Sokal &
Rohlf 1981) because the Tsets showed significantly differentvariances between the
149
sex and season categories (lower Tset; Cocltrane's C"' 0.651, P < 0.05; upper
Tset; Cocltrane's C"' 0.610, P < 0.05),
The effect ofgender on heat balance
Heating rates were measured for ten males and ten females from Midgee Rocks
and the Middleback Range. These lizards had been held in captivity for between six
months and two and a half years prior to the experiment and were all more than
two years old. The conditions of housing have been described in the previous
section of this chapter.
Measurements were made between 7 and 9 March 1994. Each lizard was weighed
to the nearest O. lg before each experiment. They were precooled to about 10°C in
a refrigerator and then placed directly beneath a SYLVANIA 150W white globe on
a brown cardboard substrate in a room with minimal air movement. The globe was
held on a retort stand at a height sufficient to heat the air around the lizard to a
stable temperature of S0-5s•c before the lizard was put there. Body temperatures
were measured with a copper constantan thermocouple attached to a FLUKE 52
digital thermometer. The thermocouples were placed 15-20 mm into the intestine
of each lizard via the cloaca. A second thermocouple attached to the same digital
thermometer was held on the cardboard next to the lizard with white masking tape
to measure the environmental temperature. The masking tape completely covered
the thermocouple and was used to avoid dislodgment and to reduce errors due to
radiation (Tracy & Christian 1986). Body temperatures were measured at 30
second intervals (see below) from the time that each lizard was placed beneath the
150
globe until it either: (I) voluntarily moved away from the heat source, or (2) began
panting, at which time it was removed from the globe.
At each reading the difference between lizard Tb and ambient temperature (liT)
was determined. This was plotted against the time of the reading, and the slope of
the regression was determined for each lizard (Figure 5.6). The modulus of the
slope (because slopes were negative) was used as a measure of the rate ofheat
gain and is equivalent to the thermal time constant of Spotila, Lommen, Bakken &
Gates (1973). The larger the number, the faster the lizard was gaining heat. The
slope for each lizard was then used as raw data in an analysis of covariance
because the data appeared to be normally distributed (Cocbrane's C = 0.530, P >
0. 05). Sex was treated as the faetor and weight as the covariate.
Heat loss was not measured because radiative cooling occurs mostly in the infrared
and emission of infrared radiation has no relation to visible colour (Bradshaw
1986).
Results
Field Body Temperatures
Over 450 field Tbs were measured from 14 lizards on 21 lizard days (Table 5.2).
There was significant heterogeneity in mean body temperatures due to sex and
season (Friedman two-way ANOVA = 55.684, P < 0.01). Males had significantly
higher Tbs than females in both November (Mann-Whitney U = 4375.5, P < 0.01)
and February (U = 2743, P < 0.01). Females showed no difference in mean Tb
151
i
between seasons (U = 3280, P > 0.05), but males had significantly higher Tbs in
November than in February (U = 7243, P < 0.05). Males showed significantly
higher variances in Tb than did females duriug both seasons, although the
differences was less outside of the breeding season (November: Fmax = I. 75, P <
0.01; February: Fmax = 1.045, P < 0.05).
The slopes of the regressions of Tb on Te' s were significantly different from zero
duriug November, but not duriug February (Table 5.2). This suggests that both
sexes were thennoregulating duriug February, but following Te more closely
during November. For each sex and each season the coefficients of determination
(R2) were higher for the regression of Tb on Tr than for the regression of Tb on Tc,
indicating that lizards are more closely coupled with the temperature of exposed
rock surfi!ces than with the temperature in crevices.
The thermal niche
Table 5.3 shows body temperatures selected by lizards in the thermal gradient.
Neither high or low Tset showed any significant differences between males, gravid
females and nongravid females (low Tset: Kruskall-Wallis H = 3.012, P > 0.05;
high Tset: H = 0.765, P > 0.05) or between seasons (low Tset: H = 0.002, P >
0.05;high Tset H = 0.352, P> 0.05). Therefore single values ofhigh and low Tset
calculated from pooled measurements of all lizards appear to summarise the
thermal preferences of C. fionni. The overall Low Tset (mean± SD) was 31.6±
3.23 and high Tset was 36.7±1.51 (Table 5.3).
152
The effect of colour on heat exchange
The slopes during heating were not significantly different between male and female
C. flonni (F1,18 = 0.06, P > O.OS)(Table 5.4), indicating that the differences in
colour between the sexes does not influence the rate at which these li7.ards gain
heat. Weight did not have a significant influence on the rate ofheat gain (F1•18 =
0.124, P > 0.05).
Discussion
The traditional approach to analysing field temperature data (see Huey & Slatkin
1976) used here has several disadvantages. These have been outlined by Hertz et
al. (1993), who also proposed a different approach to studying field thermal
relations in ectothenns. A comparison ofthe results of a traditional regression
analysis of my field data and an analysis using the approach advocated by Hertz et
al. (1993) did not result in any'difrerent conclusions. Therefore, I have only
presented the results of the simpler traditional analyses for brevity.
Although male Peninsula Dragons had higher mean Tbs than females, there was a
broad overlap in the actual values of Tb in the field. Both sexes had similar thermal
requirements, as measured by Tset range. Males showed a greater variance in field
Tbs than females during the breeding season. This may be explained by the costs of
territorial vigilance leading to less precise thermoregu]ation among males. But

much of the variation in male Tbs falls within the Tset range.
153
The difference in colour and body size between the sexes had no discernible impact
on the rate ofheat exchange. Sev~al other studies have also failed to show any
effect of colour on the rate ofheat exchange (Bakken & Gates 1975; Crisp, Cook
& Hereward 1979). The heat flux in the infrared part of the spectrum may mask
any effect of radiation at visible wavelengths (Norris 1967). Ahematively, factors
other than dermal reflectance may have a significant modifying influence on the
rate ofheating and cooling. For example integumentary circulation varies
independently of skin reflectance, but has a large effect on the rate ofheat uptake
and loss (Bradshaw & Rice 1981). While in general larger body size lowers the rate
ofheat exchange (Stevenson 1985), the differences in body size between male and
female Ctenophorus ftonni is only around 20%. This magnitude of difference in
body size is probably too small to influence the detectable rate ofheat exchange
(Stevenson 1985).
In SUllllll3l)', males and female Peninsula Dragons show no difference in the rate of
heating, had·similar thermal requirements, and overlapped broadly in field Tbs. The
differences in Tb that do occur between males and females in the field are
consistent with expectations if these sexes respond differently to constraints
imposed by territorial vigilance and the need to avoid exposure to predators. The
most parsimonious inteqiretation ofthe observed differences in field Tbs is that
they represent independent adaptations by each sex to preexisting sexual
differences in moiphology and social behaviour.
154
TIME
Activity patterns result in a gradual and sequential replacement of species dwing

the day in several lizard conmnmities on at least three continents (Pianka 1986).
These daily patterns of emergence may change seasonally, so that emergence
occurs later in the cooler winter months than in summer. Species with a unimodal
activity patterns dwing cooler months often have bimodal patterns of activity in
summer (eg. Schwaner 1989).
Sexual differences in activity patterns have been reported for both diurnal ( eg.
Christian, Tracy & Porter 1983; Robinson 1990; Bull, McNally & Dubas 1991;
Jenssen, Greenberg & Hovde 1995) and nocturnal (eg. Cooper, Caffi:ey & Vitt
1985) species of lizards. The higher activity in males appears to be, at least partly,
under the influence of androgens (Moore 1987, 1988). In other species, males and
nongravid females do not differ in activity patterns, but gravid females tend to
spend more time basking (Shine 1980; Schwarzkopf & Shine 1991 ).
Activity may be measured in two ways. First, lizards may be regarded as active if
they are exposed on the surrace and not secreted in refuges, whether or not they
are actually moving. Secondly, activity can be expressed as the amount of time
spent actually moving. In most iguanian lizards the proportion of time moving is a
subset of the time exposed. More secretive species may seldom be seen away from
refuges, but may move about within them (eg. Bustard 1967).
This section asks whether there are differences in activity pattern between male and
female Peninsula Dragons. I measured the proportion of time spent away from
155
refuges in captivity, and the proportion of time spent in motion by lizards which
were not in refuges.
Methods
Observations in captivity
Activity was measured from videos of the experiment on the perching behaviour
( see above). The videos were of nine adult male and nine adult female Peninsula
Dragons, and were taken between 19 March and 22 April 1993 and 22 March and
C
6 June 1994. Aytlvity was only measured for lone males (treatment one of the
perch height experiment) and lone females (treatment three), so each individual
lizard was housed alone in a 3.0m x 1.5m outside pen. Liz.ards were placed in the
enclosure at least 24-72 hours prior to being videoed. Each lizard was scored as
being active or inactive during 60 s periods every ten minutes between 0800 hours
and 1900 hours local time. Liz.ards were regarded as active if they were visr'ble at
anytime during the 60 s sample period. Liz.ards were scored as inactive if they were
not visible and secreted in crevices between concrete blocks for the whole 60 s.
Data were only collected on fine days when the air temperatures exceeded 25°C.
Activity was measured for three males and three females in 1993 and another six of
each sex in 1994. There were no significant differences in activity between 1993
and 1994, so the data from both years were pooled for analyses. Activity was
expressed as the number of lizards of each sex scored as being active over each
hour for males and females separately. The maximum score possible was 54 (ie. 9
lizards of one sex by six 60 s samples). The distn'butions of hourly activity scores
l56
for males and females were compared using Kolmogorov Smimov statistics (dmax,
Sokal & Rohlf 1981 ).
Obsen,ations in the field
Field observations of active lizards were conducted to determine the proportion of

time that active lizards spent in motion. A lizard was regarded as being in motion if
it was engaging in vigorous social display or locomotion. All observations were
made opportunistically at Midgee Rocks in January and February 1993
. (nonbreeding) and between August and November 1993 and 1994 (breeding).
Observations were made from a distance of 5-10 m from each lizard with the aid of
Nikkon 9x25 binoculars. At least ten different males and ten females were
observed for between 10 and 90 minutes each during the breeding and nonbreeding
seasons.
The data were expressed as the percent time spent active during observations made
during each hour of the day. These distributions were compared between the sexes
and seasons using Kolmogorov-Smimov statistics (dmax, Sokal & Rohlf 1981).
Results
Observations in captivity
Male and female Peninsula Dragons showed no significant difference in their

activity patterns in captivity (P = 0.848; Figure 5. 7). The proportion of lizards
157
active rose steadily from 0900 to a maximum at 1400 and decreased gradually until
1900.
Observations in the field
Peninsula Dragons were observed for 23.9 hr (males 10.9 hr; females 13 hr) during
the spring breeding season, and for 35. 7 hr (males 20.8 hr; females 14.9 hr) after
the breeding season, in late summer. Active males were moving for an average of
1.5% and females for 0.5% ofthe time during the breeding season (Figure 5.8).
Active males were moving for 0.8% and females for 0.6% ofthe time after the
breeding season (Figure 5.8), None of these differences were significant (dmax, P's
> 0.5).
Discussion
There were no differences in activity patterns between male and female Peninsula
Dragons. The amount of time spent on the surface was only measured outside of
the breeding season. But the lackof difference in the amount of time spent in
motion between seasons suggests that the sexes may not differ in activity at oilier
times of the year.
158
GENERAL DISCUSSION
Male and female Peninsula Dragons did not differ in their diet or activity patterns.
They did differ in their use of perches and in thermal biology. Males perched on
higher rocks than females in the field. Males also had higher mean and variance (at
least in the breeding season) in Tb than females. An experiment showed that males
and females changed their choice of perches depending on their social environment.
Males chose higher perches when a female was present, and females chose lower
perches when a male was present. Both sexes chose similar perches when housed
alone. Males may thermoregulate less accurately than females because they perch
on high boulders and thus may be exposing themselves to a different thermal
environment to females. Thus the differences between the sexes in perching
behaviour and thermal biology are readily explained in terms of their different
social roles.
The idea that ecological differences between the sexes may give rise to sexual
differences in body size and head size is largely based on optimal foraging theory,
and the way in which males and females might partition food resources (Shine
1989). In liz.ards this idea was first developed to explain patterns of covariation
between body size, head size and food particle size in insectivorous, sit-and-wait
foragers (Schoener 1967). Larger liz.ards were thought to use higher perches as
vantage points from which they search for larger food items. They were also
thought to have larger heads(= jaws) to handle these large insects. However,
optimal foraging theory does not provide a universal explanation for sexual
differences in perch height because a population of Urosauros ornatus which
showed no sexual size dimorphism still showed a sexual difference in perch height
(Zucker 1986). Furthermore male and female C. fto1mi do not differ in diet, and
159
exhibit no correlation between either body size and perch height, or head size and
maximum food particle size. There were no seasonal changes in the use of perches
which might correlate with changes in insect abundance or with changing energy
requirements due to reproduction. Thus there is little evidence for dietary or
foraging differences between the sexes which may explain the differences in perch
height between male and female C. ftonni.
Although there is no evidence that ecological differences may have given rise to
sexually dimorphic traits in Peninsula Dragons, three other species of reptile are
consistent with the predictions of the niche-divergence hypothesis. First, the
iguanid lizard Tropidurus melanopleurus exhtllits such extraordinary sexual
differences in morphology and ecology that males and females were initially
descnlled as separate species. Males prey on a wide variety of arthropods, whereas
females specialise on ants; males thermoregulate actively and females are
thermoconformers (Perez-Mellado & De La Riva 1993). Unfortunately, there are
no published data which allow an assessment of whether the head and jaws of this
species exhtllit sexual differences which might be consistent with the dietary
differences.
The second species is the Arafura Filesnake, Acrochordus arqjurae. This species
has been studied extensively by Shine and his coworkers. Females are larger than
males, feed on larger fish and forage in deeper water than males (Shine & Lambeck
1985; Shine 1986; Houston & Shine 1993). In this species large female body size is
related to increased fecundity (Shine 1986). The difference in body size between
the sexes is related to differences in foraging behaviour and diet, which are, in tum,
related to differences in jaw morphology and head shape (Shine 1986; Houston &
Shine 1993). Thus fecundity selection appears to have given rise to the dimorphism
160
in body size and then the dimorphism in head shape seems to have arisen through
niche-divergence.
The third species is Ano/is ferreus from Marie Galante in the Carrlbean
(Roughgarden 1995). This species is particularly interesting. in terms of the niche-
divergence hypothesis because it is an exception to a pattern ofbody size variation
among Anoles on Cam'bean islands (Roughgarden 1995). The pattern ofbody size
on these islands can be Jargely explained by ecological resource partitioning,
resulting from interspecific competition (Roughgarden 1995). On islands which are
inhabited by a single species ofAnolis, the lizards are usually of medium size and
have a catholic diet. On islands which are inhabited by two species, one is large and
the other is smail, and they each feed on appropriately sized insects. Since A.
fe"eus is the only species on Marie Galante it should be of medium size if
competition for food is an important determinant of body size. Instead, males are
very large, the same size as the larger species in two species communities. Females
are the same size as the smaller species in two species communities. Male and
female A. fe"eus appear to be acting like two species, at least in terms of body size
(Roughgarden 1995). This species warrants further investigations for two reasons.
First, its pattern ofbody size dimorphism makes it an exception to a hypothesis ·
' explains a large amount ofvariation in body size among Cam"bean Anoles,
which
which have themselves been a very important source of empirical studies clarifying
the role ofinterspecific competition in community ecology (Roughgarden 1995).
Secondly, Anolisfe"eus appears to be the most size dimorphic lizard on the planet
(Johnston, unpublished review of the literature).
A causative relationship between ecological differences and sexual dimorphism has

been invoked for several taxa (reviewed in Shine 1989) and lizards represent an
historically important example of this (Schoener 1967; Rand 1967). The evidence
161
for this idea. has principally been the relationship between perch height, food
particle size and sexual differences in head size ( eg. Schoener 1967). These
relationships may equally well be used to support the argument that ecological
dillerences may be the result, rather than a cause, of sexual dimorphism. The latter
view is supported by the finding that social environment is a primary determinant
of perch choice, and correlated dillerences in field Tbs, in Peninsula Dragons.
162
Chapler s NICHE DIVERGENCE
Females Males
N p N p
{n=516} {N,,,.36} {n =617} {N= 53}
Arachnida 01.4 16 01.1 12
Blattodea 02.5 24 01.9 17
Coleoptera 09.7 68 14.6 74
Diptera 01.2 08 03.7 22
Hemiptera 01.0 08 05.5 19
Hymenoptera 80.0 86 66.5 88
Lepidoptera 00.4 05 00.8 05
Mantodea 00.2 03 00.2 02
Neuroptera 00.6 08 01.0 12
Orthoptera 03.0 43 04.5 41
Vertebrata 00.0 00 00.2 02
TABLE 5.1: Proportion of total food items (N) and proportion of

stomachs containing each food type (P) for male and female
Ctenophorns fionni in the Middleback Ranges. n = number of food
items in each category; N = number of lizards in each category.
163
(')
i...
"'
Season& Tb Tb vs Tc Tb vs Tr
Sex n N {°C) b R2 b If
Breeding
Males 4 183 31.8 43.07 0.043* 0.018 0.303"' 0.106
Females 4 75 28.1 29.40 0.142* 0.116 0.555* 0.279
Nonbreeding
.... Males 3 100 34.5 24.61 -0.074"' 0.016 0.180"" 0.033

°'... Females 3 101 30.3 26.91 0.015"' 0.001 0.068"" 0.005
TABLE 5.2: Snmmacy statistics describing temperature regulation in male and female
Ctenopharus fionni in the field during the breeding season and after the breeding season. n =
number ofliz.ards; N = total number of temperature measurements; Tb= mean body temperature;
s' = variance in Tb; Tc= crevice temperature; Tr= rock temperature set-point range; b = slope of
regression of Tb on environmental temperature; R! =coefficient of determination associated with
each regression of Tb on environmental temperature. The difference(*, P < 0.05) or lack of
difference (NS, P > 0.05) of the slope from zero for each regression is indicated based on the
analysis of covariance.
f.
V,
Set Points
Season& 25th 75th
Status n N X SD Min Max percentile percentile
Breeding
Males 7 340 34.5 4.014 I6.ll3 9.5 41.5 33.0 36.8
Nongravid Females 5 275 34.7 4.015 16.116 19.0 40.S 33.7 37.1
-"'
V,
Gravid FelllJlles 4 213 30.9 6.971 48.589 10.2 46.0 27.0 37.7
Nonbreeding
Males 4 209 36.2 2.111 4.455 29.6 40.1 34.8 37.8
Fema1es 7 577 32.8 5.395 29.106 IO.I 41.2 29.6 34.l
TABLE 5.3: Body temperatures selected by lllJlle and female Ctenophorus ftonni in a therlllJII gradient during the breeding season and
after the breeding season. n = number of lizards; N =total number of temperature measurements; x == mean temperature selected; SD
standard deviation; s2 = variance; Min= minimum temperature selected; Max= maximum temperature selected. The 25th and 75th
percentiles of each distribution are used to estimate the set point temperatures following Hertz, Huey & Stevenson (1993).
Heating
Sex n X SD min. max.
Male IO -0.0301 0.0068 -0.0450 -0.021
Female ·10 -0.0311 0.0074 -0.041 -0.209
TABLE 5.4: Thermal time constants for male and female Ctenophorus
fionni during heating. n = number oflizards; x = mean; SD = standard
deviation; min. = minimum; max. = maximum. the method of
calculating the thermal time constant is shown in Figure 5.6.
166
Ar Bl Co Di He Hy Le Ma Ne Or Ve
PREY CATEGORY
FIGURE 5.1: Mean(± one standard error) number of each fuod type found in the
stomachs of male (solid bars) and female (hollow bars) Ctenophorus fionni from
the Middleback Ranges. Ar =Arachnida, Bl = Blattodea, Co = Coleoptera,
Di= Diptera, He = Hemiptera, Hy Hymenoptera, Le = Lepidoptera,
Ma= Mantodea, Ne= Neuroptera, Or= Orthoptera, Ve= Vertebrata.
167
0
• 0
00
• 0
• 0 0
0 • 0 • 0 •
• 0
0
.o• •
0
a.._____~---~----~~--~--~-~-,._----.
•
30 50 60 70 80 90
SNOUT-VENTLENGlll(mm)
FIGURE 5.2: The relationship between maximum prey size and body size for
male ( 0, ---) and female (•, -) Ctenophorus fionni from the Middleback Ranges.
168
FIGURE S.3: The arrangement of concrete blocks used to provide lizards with
:five perches of different heights (0.05m to 0. 7m) in experimental enclosures used
to test the effect of social environment on perch height in Ctenophorus fionni.
169
0.3 MALES
mean = 0.604 m
SD=0.560
N=205
0.2
~ 0.1
......
~
~ 0.0
Cl
re 0.4
FEMALES
mean= 0.463 m
0.3 SD=0,425
N= 132
0.2
0.1
I
0.0
0 1 2 3 4
PERCH HEIGHT (m)
FIGURE 5.4: Histograms of perch heights chosen by male and female

Ctenophorus fionni at Midgee Rocks, South Australia.
170
Chapter 5 NICHE DIVERGBNCE
45
40
30
25
TREATMENT
FIGURE 5.5: Mean ( ± one standard deviation) of mean perch heights chosen by
nine Ctenophomsfionni in each of four experimental treatments, showing that
males chose higher perches in the presence of a female and females chose lower
perches in the presence of a male.
171
••
20
•T
10
0.----.----..---~----,-----l
-100 100 300 500 700 900
TIME (seconds)
FIGURE 5.6: Plot of the difference between ambient temperature and body
temperature (LI. T) against time while a 17.8 g female Ctenophoros jionni was
heating. The thermal time constant was calculated as the slope ofthe linear
regression (AT"" 26.285 - 0.029.Time). Thus the thermal time constant for this
animal was -0.029. See Table 5.4.
172
60
40
20
0
8 10 12 14 16 18
TIME
FIGURE 5.7: Number out ofa possiole 54 male(•) and female (D) Ctenophorus
jionni outside of rock crevices at different times of the day.
173
• • • • • • •
2 ,..
-
-
1 ~
~
- -
-
- - -
0
J__ -- -- -~ 18-- 20
8 10 12 14 16
TIME
FIGURE 5.8: Percent of time spent moving by male<•) and female (D)
Ctenophorus jionni in the field during the breeding season (upper) and
nonbreeding season (lower).
174
CHAPTER6
MATING SYSTEM AND OPPORTUNITY FOR

SEXUAL SELECTION
The term mating system refers to the number ofmates an individual has, how
mstes are acquired, characteristics of pair bonds, and patterns ofparental care
(Davies 1991). Mating systems are the outcome of competition among individuals
to maximise their reproductive success. They can be classified with respect to their
structure ( eg. monogamy, polygamy), or in terms of the factors which determine
their structure. The key determinants of mating systems are (1) male and female
dispersion, and (2) the extent to which each sex cares for offspring (Emlen &
Oring 1977; Davies 1991).
Most males have the potential to father offspring at a faster rate than females can
produce them. Females, on the other hand, often need to mate only once to fertilise
all of their gametes. Therefore, a males' reproductive success is often limited by
the availability of females (Bateman 1948; Trivers 1972), and male dispersion is
often influenced by the distribution of females (Bradbury & Vehrencamp 1977;
Chapter6 MATING SYSTEM
Emlen & Oring 1977. Where males provide no nuptial gifts or parental care (as in
most reptiles; Shine 1988a), a females' reproductive success may be limited by
access to resources, rather thau by access to males. Consequently patterns of male
settlement would be expected to mirror the distnl>ution of females.
This framework has been influential, and has lead to considerable advancement in
our understanding of mating systems. However, it does not make clear the
relationship between mating systems and the potential for seXW1l selection in non-
resource based mating systems. More recent approaches, which make the link
between mating systems and sexual selection, rely on haviug reliable data on the
mating success and the resulting reproductive success for all males and females in a
sample (Arnold & Duvall 1994). These data are becoming available as molecular
genetic techniques are applied to behavioural studies (Burke 1989, Queller,
Strassmann & Hughes 1993), but they are not currently available for Peninsula
Dragons. Consequently this chapter relies on quantifying various proximate
determinants of mating systems in the way proposed by Emlen & Oring (1977).
But the approach is modified to take account of formal selection theory (following
Duvall & Arnold 1994), where I have suitable data.
An important step in understanding how mating systems relate to the potential for
sexual selection is the determination of the variance in male reproductive success
(Bateman 1948; Wade & Arnold 1980). The potential, or opportunity, for sexual
selection may be estimated in several ways (reviewed by Andersson 1994). These
range from relatively simple estimates of the operational sex ratio (Emlen & Oring
1977), through estimates of the relative time spent by each sex seeking mates
(Sutherland 1985), to estimates based on the variance in actual mating success
(Payne 1979; Wade & Arnold 1980; Arnold & Duvall 1994).
176
Mating systems of reptiles are poorly known (Dugan & Wiewandt 1982; Stamps
1977, 1983; Olsson & Madsen 1995). There is a tendency in the literature to
regard all liz.ards as being polygynous. However, this generalisation belies the true
variation in mating systems observed among liz.ards, which ranges from obligate
monogamy, through polygyny, to promiscuity (see Chapter 1).
Home range defence (territoriality) is a key feature of the social organisation of

many liz.ards because reproductive success of males is often limited by access to
females (Hixon 1987). In some families ofliz.ards territory defence is the norm (eg.
Agamidae and Iguanidae), whereas in others (eg. Lacertidae, Scincidae and
Teiidae) it is less common (Stamps 1983). Territoriality occurs when the benefits
of defending the territory outweigh the costs (Brown I 964). The benefits of
territoriality for male liz.ards include access to mates (eg. Deslippe & Mccloskey
1991), and access to other limited and defensi'ble resources (eg. Simon 1975; Hews
1993). In species with small female home ranges, males have the potential to
defend a larger territory including several female home ranges, and are more likely
to be polygynous (Stamps 1983). If female home ranges are large, males may only
be able to defend a single female, leading to a monogamous mating system.
Conspecific males compete against each other to defend exclusive territories, and
some males may be able to defend larger territories and mate with more females
(eg. Manzur & Fuentes 1979; Trillmich 1983).
In addition to descn"bing the structure (monogamy, polygamy) of mating systems it

is also important to determine the factors control their structure (Emlen & Oring
1977). In choosing their home ranges, females may respond either to males
themselves or to the quality of their territories. It is important to determine which
ofthese two filctors determines female dispersion. Hthe mating system is
determined by female choice among males, certain male traits might be subject to
177
Chapter 6 MATING SYSTEM
selection (ie. traits indicating vigour, or 'good genes'). Whereas if males must
compete with other males for territories in order to gain access to mates, quite
different traits might be selected for (ie. traits associated with male combat). Of
course both factors and patterns of selection may be important simultaneously
(Gtoson & Bradbury 1985; Gibson 1996), or sequentially (Burley 1981; Moore
1990).
This chapter asks the following questions: (1) What is the structure of the mating
system in Ctenophorus fionni ? (2) Is variance in male mating success based on
defense of females, or on defence of some resource (sucb as rock crevices) which
may determine the distn"bution of females? and (3) to what extent may sexual
selection be acting on males ?
METHODS
Study Site and Variables
Data were collected from lizards captured at Midgee Rocks (Figures 1.2 & 1.3)
during the four spring breeding seasons (between August and October) between
1991 and 1994. During each spring the study area was traversed in search of
lizards for five hours a day for 15 days in 1991, 20 days in 1992, 42 days in 1993
and 28 days in 1994. Field work was curtailed for personal reasons during the
spring of 1992, and no data were collected on home ranges and territory ownership
in that year. However data collected during August and early September 1992
provided some information on sex ratios and pairing behaviour. Male home ranges
were also mapped during January and February 1993 with the same methods used
178
in the spring of other years. These data were used to provide some information on
the dynamics of territory ownership during the summer of 1992-1993, but they
were not used to analyse the mating system When lizards were sighted and
approached, they readily entered rock crevices, from which they could usually be
extracted. An attempt was made to capture all resident males in all four years.
Residents were lizards seen on.more than ten days during any one spring.
When each lizard was first captured during the spring breeding season of each year
it was measured, permanently marked by toe-clipping (Fenner 1979), and its toe-
clip number was painted on its back with white, non-toxic, pliable acrylic paints.
Snout-vent length (SVL) was measured with vernier callipers to the nearest
0.1mm Body weight (Mass) was measured to the nearest 0.5g on Salter SOg
scales. Dorsal colouration was scored from one to five with increasing scores
indicating increasing degree of melanisation (Figure 2. 4). All lizards were returned
to the site of capture within 12 hours. Sex of all adults was determined by probing
(Schaeffer 1934) or by everting the hemipenes with gentle pressure at the base of
the tail. Not all traits were measured on all individuals, so sample sizes differ
among traits.
Transmitters were attached to all.adult males, on the side of the base of the tail
using quick curing araldite. Each transmitter weighed 2.0g and represented on
average 7% of body weight. The painted numbers on the back of each lizard
allowed identification of individuals at a distance without the need for recapture.
Home range area was measured as the minimum polygon (Jennrich & Turner
1969) which encompassed all koown locations for a male from systematic searches
of the study grids, radio-tracking and focal observations (see below).
179
I mapped all suitable rock crevices at .a scale of 1: 1000 on a 1.6 hectare study grid
within the larger study area. This study grid was covered by an array of stakes in a
lOxlOmgrid (see Figure 6.1). All crevices that had been used by Peninsula
Dragons within the grid during this study were included on the map. I also mapped
any other crevices which appeared to be suitable for use by dragons. Rock crevices
are important thermal refuges during periods of extreme temperatures (Chapter 5).
They serve as shelters from predators, and may also be humid refuges which
prevent desiooation during hot dry periods (Warburg 1966). Crevice density was
therefore used as a measure ofterritory quality. Maximum and minimum shaded air
temperatures over the previous 24hrs were measured at 0900 on every day that
field work was undertaken to provide some idea of the variation of climate at .the
site.
Since no actualmatings were observed during the study, fitness was estimated as
the number of females with which a male was found paired (see Chapter 3 for
justification). Only large males could be radio-tracked, so there was a potential bias
toward finding larger males in pairs. I attempted to counteract this bias by doing
daily systematic sear~s of the study site, paying particular attention to known
home ranges of yearling males. These searches were made in between forays aimed
at locating radio-tagged males.
Territory and mating system
Although dorsal colour was scored on a one to five scale there were very few
lizards with low colour scores, so all males present during each breeding season
were divided into two groups. One group were males with colour scores of four or
180
five. These males were referred to as black males. The other group were males
with colour scores of three or less and were referred to as brown males. I tested
the null hypothesis that the proportion of paired males should be equal in each of
these two classes of males using G tests with William's correction for small sample
sizes (Sokal & Rohlf 1981) for each year.
I captured and tethered males from off the study area. Resident males were
classified as being territory holders if they responded with hind-limb pushup
displays (Gt"bbons 1979) or wrect attacks to a male tethered in their home range, or
to natural territorial incursions by neighbouring males. Focal observation (Altmann
1974) were made on all resident males for between 20 and 150 minutes between
August and October in 1991, 1993 and 1994, and in January 1993 (see above), to
determine territorial status.
An attempt was made to locate each territory holding liz.ard on at least fifteen
different days during the spring of each year. Fifteen is the minimum number of
locations required to provide an estimate ofat least 90% of the home range for
male Peninsula Dragons at this site following Rose (1982). Five locations were
sufficient to provide an estimate of90% of the home range for females. Territory -
sizes reported are based on territory areas determined from> 15 locations for
males and > five locations for females. Patterns of overlap between the sexes were
based on all home ranges, no matter how few locations they were based upon.
Therefore the estimated numbers of opposite sex lizards in each home range are
conservative.
In 1991 only a few lizards were located a sufficient number of times to provide a
reliable estimate ofhome range area. In 1993 and 1994 reliable estimates ofhome
range area were obtained for all black males (N = 24) and some of the adult
181
females (N = 15) whose home ranges were entirely on the study area. Only data for
1993 and 1994 were used in statistical analyses involving home range sizes. The
home range data were not normally distnlmted (Cochrane's C = 0. 71591, P =
0.001 ), so nonparametric statistics were used for all comparisons. All territories,
no matter what nnmber oflocations they were based on, were mapped (Fignre 6.1)
to provide information on the length of each territorial male's tenure.
Determinants of home range area
To determine whether larger lizards had larger home ranges, body mass was
regressed against home range area for males and females in two separate analyses.
Ifhome range area was detennined by the availability of food resources, these
regressions should yield similar slopes for males and females . If males had a
steeper slope than females it would suggest that male home range area is related to
male body size. Sµch a relationship might arise if male were competing with each
other for access to females, or some other resource.
The hypothesis that rock crevices are an important detenninant ofhome range area
was tested by regressing home range area against crevice density in separate
analyses for males and females. A negative slope would indicate that crevice
density may be a detenninant of home range area. A larger slope in one sex: might
indicate that crevice density is more important to that sex:. The regression lines
were compared using analyses of covariance (Sokal & Rohlf 1981).
The alternative hypotheses that the number of females in a territory or the number
of crevices in a territory were detenninants of territory area among male Peninsula
182
Dragons was tested using partial correlation coefficients (Sokal & Rohlf 1981).
Territozy area was square root-transformed for all regression analyses.
Opportunity for sexual selection
Three approaches were used to estimate the potential for sexual selection. The first
involved calculating the breeding sex ratio, following Arnold & Duvall (1994). An
unbalanced sex ratio would suggest greater competition in one sex for mates. The
second approach is based on the idea that mating success. can be restricted by the
time necessacy for mating and caring for offspring (Sutherland 1985), in the same
way that food intake can be restricted by handling time (Holling 1959). Using this
approach the strength of sexual selection can be assessed in terms of the
proportion of time spent seeking mates by each sex (Sutherland 1985). The greater
the difference between the sexes in time spent seeking mates, the stronger the
competition for mates. This will be referred to here as the time budget approach.
The third approach to measuring the potential for sexual selection is based on
measurements of the variance in male reproductive success (Wade & Amold
1980). This variance originates from two sources. The first is the variance in clutch
size of females with whom a male mates. The second is the variance in the number
of females with whom a male mates. Both factors combined define the total
intensity of selection on male reproduction. The variance in the number of mates
obtained defines the intensity of selection due to sexual selection only. I used the
method proposed by Wade &Amold (1980), as modified by Kluge (1981), to
estimate the intensity of sexual selection on male Peninsula Dragons. I will refer to
this method as the variance approach.
183
sex ratio
The total number of mature lizards of each sex, or breeding sex ratio (Arnold &
Duvall 1994), was estimated from the lizards found between August and October
of each year. Auy lizard with an SVL ~ 55mm was assumed to be mature (Chapter
3). It is more usual to use the operational sex ratio as an index of competition for
mates, following Emlen & Oring (1977). But simulation studies have shown that
the operational sex ratio is a poor predictor of variance in male mating success in
mating systems with mate searching and guarding (Imms 1988). Furthermore, the
breeding sex ratio constitutes a constraint on the mean mating success for each sex,
and can be related directly to the selection acting on mating success (Arnold &
Duvall 1994). Sex ratios calculated for each year separately, and the sex ratio
pooled across years were compared to an expected sex ratio of unity using G-
statistics (Sokal & Rohlf 1981).
Time Budget Approach
Liz.ards are able to spend a certain amount of time ( 1) on searching for mates and
on activities related to obtaining mates. Each mating leaves a lizard unable to seek
other mates for a time (H) available associated with courtship, mate guarding,
copulation, time spent gestating eggs or replacing gametes. So the average number
of matings (F) obtained by each sex affects the time left to search for additional
mates. We assume that mates are encountered at a random rate (:I.) during the time
spent seeking mates. The value of:I. can be estimated (fullowing Sutherland 1985)
as:
184
;\. = 1 / [(T/F)-H]
These parameters may be used to estimate various parameters which are of interest
in examining the mating system of Peninsula Dragons (following Sutherland 1985;
see Table 6.5). The strength of sexual selection (S) on each sex may be estimated
as:
S= (F/1) I;\..
The investment in offspring (I) by each sex may be estimated as:
I= FI [(T!H)-F].
The relative strength of sexual selection and of parental investment may be
expressed as the ratio (males/females) of these parameters for the sexes. The ratio
of maximal breeding by males to females may be estimated as:
RB=Hr!Hm.
These measures were originally formulated for lifetime estimates of the parameters
T, F and H (Sutherland 1985). However,. I have based my estimates on values for a

single breeding season and taken from Chapter 3 (see Table 6.5).
185
Variance approach
The variables needed are the mean (x,) and variance (s2,) of male mating success,
and the mean (xr) and variance (s2r) of female clutch size. ·These data are given in
Table 6.6. The male mating success variables are given Table 6.3. The clutch size
data are from Chapter 3. The same clutch size data were used in estimating the
intensity of selection for all four years between 1991 and 1994. Following Kluge
( 1981) the variance in male reproductive success is:
The total intensity of selection on male reproductive success is:
The intensity of sexual selection on male reproduction is:
Is= s2. I x.2.
Im measures the intensity of selection due to variance in male mating success and
clutch size, while Is only considers the parameters affected by sexual selection (ie.
mating success only). Is/Im gives the proportion of selection on male reproduction
that is due to sexual selection alone. These indices give a measure of the
opportunity for sexual selection. Evolutionary change can result only if there is a
heritable basis for variance in fitness. Determinations of these indices are useful
186
because they pennit quantitative comparisons of variation in the intensity of

selection within and between species.
RESULTS
Territory & Mating System
All black males (ie. with.colour scores of four or five) held well defined territories.
Younger, brown males (colour scores ::;; 3) were never observed to successfully
defend a territory, but lived within older males' territories. Of the 41 males who
established territories on the study site during the four years of the study, 36 (66%)
were present during a single year, 12 (29%) maintained territories for two years,
and two (#s 76 and 259) maintained territories for three years (Figure 6.1). Female
home ranges were widely spaced and seldomly overlapped with each other (Figure
6.2).
Black males had larger home ranges (x = 0.113 ha, SD= 0.0720, n = 24) than
females (x =0.026, SD= 0.0207, n = 15) (U= 12.5, P= O.OOl)wbendata from
1993 and 1994 were pooled. The home range area of black males was significantly
larger in 1994 than 1993 (U = 16, P = 0.043), but females showed no such
difference (U = 4.5, P = 0.2975). Male home ranges were significantly larger than
those offemales in both years (1993: U = 3.5, P = 0.0101; 1994: U = l, P =
0.019). The larger male home ranges present males with an opportunity to be
polygynous by including more than one female home range within their own. An
average ofl.6 (SD= 1.46, minimum= 0, m.axiinum= 8) mature females overlap
each male home range (Table 6.1 ). There is also considerable opportunity for
187
females to mate with more than one male (Figure 6.2). An average of 1.5 (SD=
1.25}, minimum= 0, maximum= 6) mature males overlap each female home range
ti,es '5'f a,,p,iuv_ +,, hue... 1>eev,.
(Table 6.1 ). However, this potential for polyandry~ realised.
In most cases only one of the males with whom a female's home range overlapped
held a territory. The remainder ofthe males were smaller, brown, yearling males
who lived within the older, black male's territory by adopting secretive or
subordinate behaviour. Older, darker, territory holding males had almost exclusive
access to any females occurring on their territories. In 1991 and 1992 younger,
brown males paired significantly less frequently than older black males (Table 6.2).
This trend continued in 1993 and 1994, but was not significant. ·Of 47 pairs
recorded during the study, 45 (96%) involved territorial males with colour scores
of 4 or 5. Only two younger, brown males, that lacked territories, were found
paired with females. The situation was similar in both cases. A female had been
found paired with an older territory holding, black male, and a yearling, brown
male was known to live nearby. The older male was killed by a kestrel and, within
hours of this happening, the yearling was found paired with the female.
Th~ patterns ofhome range overlap translate into a pattern ofpairing in wµich
most males (73%) and females (68%) were not found paired at all (Table 6.3).The
males that <lid pair, were found with either one (24%) or two (3%) females.
Females were also found paired with either one (31%) or two (0.7%) males. The
patterns of pairing did not differ significantly between the sexes (Kolmogorov-
Smirnov test, P = 0.41).
188
Determinants of home range area
Tue climate at Midgee Rocks is harsh. Air temperatures varied from -2°C to 49°C
during this study. Peninsula Dragons characteristically use rock crevices to sheher
from environmental extremes, as well as from predators. They used crevices that
were between 4-10 mm in width, and at least 60 mm in depth. These included
crevices beneath thin sheets of exfoliated granite, through to crevices between
large boulders. Among both males and females there was a significant negative
relationship between home range area and crevice density (males: F 1;M = 6. 79, P =
0.016; females: F1•14 = 7.54, P = 0.017) (Figure 6.3). Home ranges were smaller
where crevices were abundant. Tue slopes of the male regression was significantly
steeper than the slope for females (F1;38 = 10.66, P = 0.002). This indicates that
crevices may be a limiting resource for both sexes, but that the number of crevices
may be more important fur males than females.
Tue home ranges of black males included an average of 42.7 crevices (SD= 30.73,
minimum= 5, maximum= 139), whereas female home ranges included
considerably fewer crevices (x = 16.1, SD= 8.34, roinirnuro-4, maximuni= 27).
This difference was significant (U = 80, P = 0.003). Tue raw relationship between
both number of females and number of crevices in a males home range, and male
home range size were both significant (Figures 6.4 & 6.5). When the effect of the
number of crevices was removed, there was no correlation between the number of
females and territory area (rp =-0.33, P = 0.123). After partialling out the effect of
females, correlation between the number of crevices and territory area remained
significant (rp =0.77, P < 0.001).There was, however, a significant partial
correlation between the number of crevices and the number of females on a male's
territory (rp = 0.72, P < 0.001). These data suggest that male Peninsula Dragons
189
may defend large territories which contain many crevices, rather than defending
females. A large number of crevices in a male's territory provides him with access
to more females, even though there is no direct link between the number of females
and the ~ of a males' home range. There was no correlation between female
home range size and the number of males, or the number of crevices in female
home ranges (P's> 0.13).
Opportunity for sexual selection
Sex ratio
The sex ratio of adult lizards during the breeding season at Midgee Rocks varied
from 0.64 to 1.32 over the four years of the study, but none of these values were
significantly different from unity (P's > 0.1) (Table 6.4).
Time budget approach
Both male and female Peninsula Dragons may spend the whole breeding season,
from August to October (Chapter 3) searching for mates. So the value ofTmaybe
estimated as 90 days. The average number of mates (F), calculated from the .
pairing success (Table 6.3), was 0.292 for males and 0.321 for females. The time
that males were unable to seek other mates (H) would be the average time spent
paired with each female during the breeding season(2.2 (1-10) days)(Chapter 3)
multiplied by the number of females with whom he mates. For females, H would be
190
the time spent gravid (minimum 39 days; Chapter 3). Thus the ,strength of sexual
selection is considerably higher on males, and females invest considerably more
time on their offspring (primarily cail)'ing eggs) than males. These calculations
suggest that males can breed at a rate nearly eighteen times that of females (Table
6.5).
Variance approach
Mean male mating success varied between 0, 16 and 0.41 over the four years of the
study and was.highest in 1992 (Table 6.6). The total opportunity for selection on
male reproduction (lm) varied from 2.5 to 6, 1, and was lowest in 1992. The
opportunity for sexual selection (ls) varied from 2.32 to 5.63, and was also lowest
in 1992. The Is value ofS.63 in 1994 is the highest value of this parameter
recorded for any 1izard (see Table 6. 7). The proportional opportunity for sexual
selection (Is/Im) was consistently 93% of the total opportunity for selection in all
four years (Table 6.6).
DISCUSSION
Mating System
Male Peninsula Dragons bave larger home ranges than females, and may overlap
with more than one female. Large males defend their home ranges against other
territorial males, but smaller, yearling males live within older males' territories.
191
Cbapter6 MATING SYSTEM
Females defend their home ranges against other females, but may overlap with one
or more males. This pattern ofhome range size and overlap permits both sexes to
be polygamous. However, the observed pattern was that most males and females
found in pairs, were fuund with only one partner in any one year (see Table 6.3).
Thus most liz.ards that pair appear to be monogamous, at least within a year,
despite the potential fur polygamy.
Polygyny is the most commonly reported mating system among liz.ards (Stamps
1983). Many liz.ards that are characterised as being polygynous have a substantial
proportion of males that mate with only one female per year (eg. Tmlde 1967;
Manzur & Fuentes 1979; Ruby 1981; McCloskey et al. 1990). On the other hand,
females are known to mate with multiple males in at least one lizard (Olsson et al.
1994), and a snake (Madsen et al. 1992). Males in both of these species are non
territorial, and mate with multiple females. The only obligately monogamous
reptile, the skink Tiliqua rugosa, has similar sized male and female home ranges
(Bull 1994). In Tiliqua both males and females overlap home ranges with several
mature opposite sexed liz.ards, and males exh.toit prolonged mate guarding
behaviour (Bull 1988, 1994).
Like mate guarding, territoriality is a means by which males attempt to gain

exclusive access to females (Stamps 1983). Most lizard territories appear to
provide access to females through control of crucial resources, rather than
providing direct access to the females themselves. Food (Simon 1975; Hews
1993), sites for thennoregulation (Trillmich 1983), shelter from predators (Vitt &
Zani 1995), and trees (Manzur & Fuentes 1979; McCloskey et al. 1990) have been
shown to be important resources which are related to the distn"bution of females.
192
Crevices may be an important detenninant of home range size in Peoinsula

Dragons because there was a negative relationship between home range area and
crevice density for both sexes. Male Peninsula Dragons may defend
'superterritories' (sensu Stamps 1983), which include a large number of crevices
because the number of crevices had a greater relationship to territory area than did
the number of females overlapping the territory. Control of a large number of
crevices may increase the likelihood that a male will have access to females.
Crevices may be important for thermoregulation (Chapter 5), water conservation

(Warburg 1966), and as shelter from predators. Crevices are probably an important
resource for females because there is a negative relationship between home range
area and crevice density. However, this relationship disappears when the effect of
the number of overlapping males was controlled for using partial correlation.
analysis. There was also no relationship between home range size and the number
of overlapping males when the effect of crevice number was excluded using a
partial correlation. This pattern would be explained if females settle in home ranges
which include an adequate but small number of crevices and males. Larger female
home ranges may only be necessary in so far as they allow females to have access
to both males and crevices. The likelihood of including males and crevices in an
area increases with the size of that area, and the occurrence of males is correlated
with the occurrence of crevices (see above ). Independently neither males nor
crevices may be important to females above a low threshold value.
Smaller, yearling male Peninsula Dragons were unable to defend a territory. They
remained within the territories oflarger older males by adoptiog subordinate
behaviour and maintaining the brown dorsal colouration of females (see Chapter
2). These yearling males were probably sexually mature because they were larger
than the size at which maturity is reached (Chapter 3). They typically had some
193
yellow colouration on the throat and a grey wash where the black chest patch
develops in older males. The black chest patch probably indicates a higher
androgen titre associated with enlargement of the testes (Cooper & Greenberg
1992). Young, mature males resemble adult females in many other sexually
dichromatic species (Darwin 1871; Rohwer, Fretwell & Niles 1980). This appears
to represent an age-related alternative male reproductive tactic (Cooper & Vitt
1987; Hews 1995) because it allows young males to survive and grow until they
are large enough to defend a territory of their own, without being attacked by
territorial males. Some of these young males may be able to pair with females if a
territory holding male dies and leaves a receptive female unattended.
Unfortunately, few yearling males were found at enough different localities to
obtain an accurate estimate of their home range size. Most yearling males were
repeatedly found at one or two localities, indicating that their home ranges may be
small.
The pairing behaviour in Peninsula Dragons during spring may be a form of mate
guarding. Mate guarding could be a very important determinant of a male's
reproductive success in a species such as this, where many females overlap with the
home ranges of several males. Yearling males may represent a threat to a territorial
male's reproductive potential if their subordinate behaviour and female mimicry
allows then to cuckold older males. Mate guarding has been reported in at least
three other lizards (Vitt & Cooper 1985; Bull 1988; Olsson 1993b).
194
Cbapter6 MATING SYSTEM
Opportunity for Sexual Selection
The approximately equal number of mature males and females found at Midgee
Rocks suggests that neither sex were in short supply. However most of the mature
males did not hold territories (Table 6.2), and the few territorial males obtained the
majority of pairings (and presumably matings) with females (Table 6.3).
Territoriality severely skewed the pattern of reproductive success among males.
Other factors further enhanced thls skew. For example, most mature females did
uot breed every year (Chapter 3). The actual number of females that were receptive
was lower than raw counts of females would suggest. Thus there is considerable
variation in reproductive success among males associated with whether they have a
territory, and whether their territory overlaps the home range of a receptive female.
M.ale and female Peninsula Dragons differed in the amount of time they spent
searching for mates. males spent an average oftwo days paired with any female
with whom they presumably mated, and were able to mate multiple times. They
were able to spend the majority of their time searching for mates. Females had a
refractory period after mating of about forty days. During thls period they gestated
eggs, and were unavailable for further reproduction. These differences in time
spent searching for mates and in parental investment meant that individual males
were potentially able to reproduce at a considerably higher rate than females.
Therefore males were likely to be under greater selection associated with
reproduction than females.
Both the mean and variance of male pairing success varied among the four years of
the study. Both the mean and the variance were largest in 1992, the wettest year.
There was considerable opportunity for selection on traits associated with
195
reproduction (Im) in all years. The majority of this opportunity for selection was
due to the opportunity for sexual selection (ls) in all four years. The magnitude of
the Is varied from 2.32-5.63. This is similar to the range of values calculated from
studies of other territorial iguanian lizards (0.12-5.00; Table 6.7). All of the species
included in Table 6. 7 have sexually dimoi:phic traits which were assumed by the
authors to have evolved through sexual selection. The value ofIm could only be
calculated from data available in three of the studies reported in Table 6.7 (Trivers
1976, Ruby 1981, 1984). In all three cases the value ofIm and Is were identical, so
all of the opportunity for selection was due to variation in mating success (ie.
sexual selection).
Sexual selection may take the form of mate choice (intersexual selection) or male
combat (intrasexual selection) in male Peninsula Dragons. Males compete with
each other for territories which provide almost exclusive mating access to females
occurring within them Therefore intrasexual selection is likely to act upon traits
which allow males to establish and maintain territories. A female may exln'bit
choice among males before or after they establish their home range. Females may
base their decision about where they should establish a home range on the
distn'bution of resources or on male characteristics. Females generally settle in -
home ranges which contain a small number of suitable crevices and at least one
male, but neither of these factors independently explains any variation in female
home range size. Nevertheless females in established home ranges often overlap
with several mature males. Therefore there is a potential for females to exht'bit
choice among males.
In summary, Peninsula Dragons appear to have a mating system best descn'bed as
resource defence polygyny. In any one breeding season most males do not pair at
all, others appear to pair with a single female, and only a few pair with more than
196
one female. Males spend more time searching for mates than females and are able
to reproduce at a higher rate than females. Males show no parental care and exhibit
greater variance in reproductive success than females. These factors provide an
opportunity for sexual selection to act on males. The circumstances of the mating
system permit the action of both intrasexual and intersexual selection. But
intrasexual selection., through male combat over territories, is likely to be a major
determinant of fitness among males.
197
# lizards N 0 1 2 3 4 5 6 7 8
overlappin~
Male home ranges
, 1991 II 4 5 2
1993 17 4 2 4 5 2
1994 21 8 1 6 2 1 2 1
Total 49 16 8 12 7 3 2 l
....
'C Female home ranges
00
1991 8 0 7 1
1993 23 4 10 6 2 I
1994 19 5 4 5 '2 2 1
Total 50 9 21 12 4 3 l
TABLE 6.1: Number oflizards of the opposite sex overlapping a lizard's home
range in Ctenophorus fionni at Midgee Rocks. Numbers in brackets represent
gravid females
~
~
~
I
brown black
Year N % found paired N % found paired G - statistic
-
1
1991 09 0 43 26 032.14*
\0
\0
1992 17. 6 32 47 107.30*
1993 05 20 38 37 000.530
1994 07 0 11 46 002.870
TABLE 6.2: Proportion of brown and black male Peninsula Dragons found
paired with a females over four years at Midgee Rocks. N = total number of males
in each colour class.*= P < 0.01.
Number ofPairs
Year 0 1 2
Males
1991 43 9
1992 26 14 3
1993 27 13 l
1994 22 3
Total 118 39 4
Females
1991 52 9
1992 13 20
1993 12 l3 1
1994 16 3
Total 93 43 1
TABLE 6.3: Frequency of pairing success for male and female

Peninsula Dragons at Midgee Rocks between 1991 and 1994.
200
year females males sex ratio

1991 61 52 0.85
1992 33 21 0.64
1993 27 35 1.30
1994 19 25 1.32
total 140 133 0.95
TABLE 6.4: Number of mature male and female

Cte,wphorus ftonni, and the sex ratio at Midgee
Rocks from 1991 to 1994. The sex ratios are not
significantly different from unity.
201
n
Parameter Symbol Males Females

Time spent searching for and handling mates (days) T 90 90
Time spent handling mates (days) H 2.2 39
Mean number of mates F 0.292 0.321
Strength of sexual selection s -0.001476 -0.000092
t..:> Parental investment Pl 0.0072 0.1616
s Relative strength of sexual selection RS 16.0
Relative parental investment RPI .045
Ratio of maximal breeding RB 17.8
TABLE 6.5: Indices of sexual selection based on time spent searching for mares
(following Sutherland 1985). The upper panel gives raw values used to calculate
the indices of sexual selection, which are given in the middle panel The bottom
panel gives the ratios (males/females) of the indices for each sex.
1991 1992 1993 1994

Male mating success
Xs 0.1698 0.4146 0.2973 0.1579
s2. 0.1437 0.3988 0.2703 0.1404
Female clutch siz.e
Xf 4.240 4.240 4.240 4.240
s2r l.277 1.277 1.277 1.277
Im 5.40 2.49 3.30 6.08

Is 4.98 2.32 3.06 5.63
Isllm 0.93 0.93 0.93 0.93
TABLE 6.6: Parameters estimating the intensity of sexual selectinn using the
variance approach for Ctenophorus fionnl at Midgee Rocks over four years. x. =
mean male mating success; s2s = variance of male mating success; xr = mean clutch
siz.e; s2r = variance of clutch siz.e; Im = total opportunity for selection of male
reproduction; Is = opportunity for sexual selection on males; Is/Im = opportunity
for sexual selection as a proportion of total opportunity for selection on male
r7Production.
203
Species Is Reference
A,w/is carolinensis 0.94 Ruby(l984)
Ano/is garmani 5.00 Trivers (1976)
Amblyrhynchus cristatus 4.15 Trillmich (1983)
Sceloporus ja"uvi 1.42 Ruby(I981)
Agamaagama 0.12-0,51 Harris ( 1964)
Cte,wphorus fionni 2.32-5.63 this study
TABLE 6.7: Calculated values of the opportunity for sexual selection (Is)
for six species of territorial iguanian lizards for which appropriate data are
available.
204
(')
[.~: ...
·~. ' ' '
1993
1991 "'
1992 1994
"'
('£):
FIGURE 6.1: Map of the home ranges of territorial males on a 1.6 ha study area over summer of four consecutive
years at Midgee Rocks. Bold lines= home range boundaries; numbers show identity of individual lizards. Shading
indicates home,ranges of males present in the previous year. Dots show position of jarrah stakes arranged in a !Om x
lOm grid. The maps labelled 1991, 1993 and 1994 were based on data collected between August and October in those
years. The map labelled 1992 was based on data collected in January and Pebmary 1993 (see methods for explana1ion ).
FIGURE 6.2: Example ofthe pattern ofhome range size and overlap between
female (thin lines) and male (thick lines) Ctenophorusfionni. Scale bar= 10
metres.
206
.3----------------------~
• •
•
t5 ••
...
.2
~
w
C)
!
"c9• •••
w
::?i .1
0
I
-0____ •
-c/.•---0..a~---- eg,. ~ I
O.O,l..o--~--@~_;·::,__o
_ _~·-.:::O:_-_-~-_-_.c~-~-~---..::--=-;-.,,.-';;;;,::i_~1600\'
200 400 eoo soo 1000 1290 1400
CREVICES DENSITY (crevices/ha)
FIGURE 6.3: Relationship between crevice density (crevices/m2) and home range
size for male ( •, solid line) and female (0, broken line) Ctenophorus ftonni.
207
10
r:F.1
~
8
•
•
~ ••••
6
•• • ••
• • •
- -
i:,:t.;
4
• .-
z 0
•
•• • •
2 ·-
•
QI
-2
••
- •
0.0 .1 .2 .3
TERRITORY AREA (ha)
FIGURE 6.4: Relationship between th:number of mature females in a male's

home range and home range size in Ctenophorus ftonni. Only bJack, territorial
males are included.
208
Chapter 6 MATING SYSTBM
•
• •
0
z
0
0.0 • .I .2 .3
TERRITORY AREA (ha)
FIGURE 6.5: Relationship between the number crevices in a male's home range
and home range size in Ctenophorus fionni. Only black, tenitorial males are
included.
209
CHAPTER 7
TARGETS OF SEXUAL SELECTION IN MALES
Differences in the variance in mating success between the sexes are necessary for
sexual selection to occur. However, measures ofthe potential for sexual selection
based on variances in mating success alone must be viewed with caution (Ralls
1977; Payne 1979, 1984; Wade & Amold 1980; Amold & Wade 1984a,b).
Showing that there is higher variance in mating success in one sex, is not sufficient
to demonstrate that sexual selection is occurring. Rather, it indicates the potential
for sexual selection to produce an evolutionary response.
Environmental mctors, such as temperature and water availability, have a large

influence on activity patterns, reproductive biology, and survivorship in desert
reptiles (eg. Bradshaw & Main 1968; Bradshaw 1971; Bradshaw, Saint (iirons &
Bradshaw 1991). Different individuals may be exposed to small, local differences in
environmental conditions which may translate into differences in mating success.
For example, a lizard living in a deep valley may be exposed to the sun for only a
fuw hours each day and have a greatly reduce activity period in which to search for
Chapter? TARGETS OF SEXUAL SELECTION
mates. On the other hand, a lizard living on top of a hill may have a much longer
activity period and be able to find many more mates. The high variation in mating
success between the two lizards would be a result of were they live, and would not
reflect selection on any heritable trait.
Variance in mating success may be particularly influenced by environmental

variation in species which are polygynous and in which females do not mate as
frequently as males (Sutherland 1985; Hubbell & Johnson 1987). Ifvariance in
mating success is environmentally induced, it cannot result in sexual selection
because a response to selection requires that a trait be heritable (Lande & Arnold
1983). The differences in variance in mating success between male and female
Peninsula Dragons (Chapter 6) may be an artefact of environmentally induced
variance in mating success, or it may result from heritable variation and give rise to
sexual selection.
To demonstrate sexual selection one must show that selection is acting on

particular traits. That is, one must identify targets of selection. This means that a
trait must vaxy with mating success among individuals of the same sex. The
identification of direct targets of selection may be problematical because traits that
explain variance in mating success are often interrelated. The independent effects
of correlated traits must be teased out using appropriate statistical techniques
(Lande & Arnold 1983; Brodie, Moore & Janzen 1995).
Traits which are subject to sexual selection have been identified in eight species of
lizards. This low number of studies on sexual selection in lizards is surprising
because they show strong sexual dimorphism in body size, body shape, colouration
and behaviour (eg. Stamps 1983), and are important model organisms in ecology
(eg. Huey, Pianka & Schoener 1983). The traits identified as being subject to
211
Chapter7 T ARGBTS OF SEXUAL SELECTION
sexual selection in lizards are male size (Ruby 1981, 1984; Hews 1990; Andren
1986; Madsen 1988; Madsen & Shine 1992, 1993; Madsen, Shine, Loman &
Hakansson 1992; Olsson 1992; Cooper & Vitt 1993; Baird, Acree & Sloan 1996),
male shape (Hews 1990), male visual ornaments {Greenberg & Noble 1944; Crews
1975; Sigmund 1983; Olsson 1994a,b; Diaz 1993), male display (Greenberg &
Noble 1944; Crews 1975) and male territory characteristics (Trivers 1976; Hews
1990). Most of these traits are associated with male-male competition for
territories, but body size has been shown to be the subject of female choice
independent of territory quality in one lizard species (Cooper & Vrtt 1993).
The relative importance of intra- and intersexual selection remains a topic of

debate, both generally (Thomhill 1980; Bradbury & Davies 1987), and in studies of
reptiles in particular (Olsson & Madsen 1995; Tokarz 1995). In part, this may be
due to the difficulties in determining whether a female is choosing a male or his
territory (eg. Schmale 1981; Alatalo, Lundberg & Glynn 1986; Hastings 1988; von
Schantz et al. 1989). Territorial lizards offer a conservative test of whether mate
choice gives rise to sexual selection in this group.
Male Peninsula Dragons compete for territories, within which mating occurs
(Chapter 6). Female home ranges may overlap with one or more territories held by
older, black males, and with the home ranges of a number of non-territorial,
yearling, brown males. Yearling males remain on adult male territories by adopting
subordinate behaviour patterns (Chapter 6). Females may mate with any of the
males occurring in their home range. So there is the potential for both intra- and
intersexual selection to occur in this species. Any trait that allows a male to gain a
Ix S:tJbjc:-t~ '/l,,e. ,;.,ftve..<E. of
territory (and thus gain access to females) may be assumed to\l'eSlilt .&em.I
intrasexual selection. Any trait which is identified as a target of sexual selection,
212
Chapter7 TARGETS OF SEXUAL SELECTION
and which cannot be explained by male combat, may have arisen through the action
of intersexual selection.
In this chapter I ask two questions; (l) Can any of several moxphological traits
which reflect body size and shape in male Peninsula Dragon lizards be identified as
targets of sexual selection, and (2) what mode(s) of sexual selection are acting on
these traits. My approach to the second question is to determine which of the traits
under selection play a role in male combat over territories, and therefore result (at
least partly) from intrasexual selection. If a target of sexual selection cannot be
explained by male combat then intersexual selection (mate choice) may be
occurring.
GENERAL METHODS
Data were collected during the spring breeding seasons (August to October)
between 1991 and 1994 at Midgee Rocks (Figure 1.2 & 1.3). Two study areas
were located on two separate, but adjacent, rock outcrops and were defined by a
10mxl0m grid of Jarrah stakes ( 1.6m) to allow accurate location of lizards. The
northern study site was the same one used to study the mating system in Chapter 6,
and had an area of 1.6 ha. The southern site had an area was the same site used for
an experimental manipulation of male colouration (see Chapter 8), and had an area
of2.4 ha. An attempt was made to capture all resident males in the more northerly
study area in all four years. The southerly area was only studied during 1992 and
1993, and the data reported here were collected prior to the experimental
manipulation. Residents were lizards seen regularly on the study areas between
August and October.
213
Chapter7 T ARGEI'S OF SEXUAL SELECTION
When each lizard was first captured in spring of each year it was measured,
permanently marked by toe-clipping (Fenner 1979), its toe-clip number was
painted on its back with white, non-toxic, pliable acrylic paint, and it was returned
to the site of capture within 12 hours. Transmitters were attached. to all aduh males
over 15g in body mass, on the side of the base·of the tail, using quick curing
araldite. Each transmitter weighed 2.0g and represented less than 7% of each
lizards body weight. The painted numbers on the back of each lizard allowed
identification of individuals at a distance without the need for recapture.
Measurements taken (Figure 2.3) were snout-vent length (SVL}, depth of the head
innnediately behind the eye (HD), maximum head width (HW), distance from the
tip of the snout to the anterior margin of the tympani (HL), and body weight
(Mass). All linear measurements were made with vernier callipers to the nearest
0.1mm. Lizards wei:e weighed to the nearest 0.5g on Salter 50g scales. Home
range area was measured as the mininmm convex polygon (Jennrich &. Turner
1969) which encompassed all known locations for a· male from systematic searches
ofthe study grids, radio-tracking and focal observations (Chapter 6) over the
spring. Each lizard was located at least fifteen (and up to 81) times during the
spring of each year of the study. Fifteen is the minimum number oflocations
required to provide an estimate of at least 90% of the home range for male
Peninsula Dragons at this site, following Rose (1982). Not all traits were measured
on all individuals, so sample sizes differ among traits. Motphological measurements
were available for many more lizards than were data on home range area .and
pairing success.
Fitness was estimated indirectly as the number of females with, whom a male was
found paired because no actual matings were observed during the study ( see
214
Chapter 3 for discussion). Since only large males could be radio-tracked there was
.a potential bias because those males. could be located more often than smaller
males, without transmitters. Thus large males might be found in pairs more often,
even if large and small males paired equally frequently. I attempted to counteract
this bias by doing daily systematic searches of each study site, paying particular
attention to known home ranges of yearling males. These searches were done
between forays aimed at locating radio-tagged males.
The data were used to: ( 1) calculate coefficients of sexual selection on continuous
morphological traits and home range area, and (2) test whether male competition
for territories might explain the pattern of sexual selection.
SELECTION ANALYSES
Methods
Estimates of selection coefficients cannot normally be made using cross-sectional

data (Lande & Arnold 1983; Brodie, Moore & Janzen 1995). This is because
cross-sectional da~a are collected at a single point in time and may include animals
from different generations. Cross sectional estimates of selection reflect both
current selection, and any response to selection between generations. Longitudinal
data, on the other hand, include measures of fitness and trait values for individuals
in a single cohort, and provide the best estimates of selection (Lande & Arnold
198~).
2.15
Chapter7 TARGETSOFSEXUALSELEcrION
Each male lizard in this study was assigned to a cohort based on estimates ofits
age at first capture. Age estimates were derived from a logistic growth model for
SVL (Chapter 4). Each cohort was defined by the year in which lizards of that
cohort were thought to have hatched. Selection coefficients were then estimated, in
two separate analyses, for 36 male lizards assigned to the 1990 cohort, and for 27
male lizards assigned to the 1991 cohort. The estimates of selection coefficients
were based on data collected during the spring breeding seasons of 1992 and 1993.
Measures of selection were calculated by comparing male Peninsula Dragons

which did pair to those which did not. Since fitness was measured as presomed
mating success, the measures of selection calculated here are coefficients of sexual
selection (Arnold 1983; see Chapter 1). Selection may act directly on a trait, or
indirectly via the effects of selection on correlated traits (Lande & Arnold 1983).
Selection differentials measure the total. direct and indirect selection, acting on a
trait. Selection gradients measure selection acting directly on a trait, independent
of all other measured correlated traits (Lande & Arnold 1983; Mitchell-Olds &
Shaw 1987).
The total selection acting on each trait was estimated using standardised selection
differentials (Endler 1986):
j =va-Vb /Vb
where i is a measure of directional selection,} is a measure ofvariance selection, Xb

is the mean of a trait before selection, Vb is the variance of a trait before selection,
x. is the mean of the trait after selection, and Va is the variance of a trait after
216
selection. In this study, all males were used to calculate Xb and Vb, and only those
males which paired were used to calculate x. and v•. If i is positive, directional
selection is acting to increase the si7.e ofthe trait. If i is negative then there is
directional selection to decrease the size of the trait. Negative values ofj indicate
stabilising selection, and positive values ofj indicate dismptive selection. The null
hypotheses that values of i andj were not significantly different from zero were
tested for each trait in each cohort using modified Student's I-tests (for i) and
Fmax tests (forj) (Endler 1986).
Targets of direct selection were identified using standardised partial regression

coefficients from multiple regression analyses as estimates of selection gradients
(Lande & Arnold 1983; Brodie, Moore & Janzen 1995). Standardised coefficients
allow easy comparisons among difierent studies, because they express selection in
standard deviation units. Linear multiple regression models were used to calculate
linear selection gradients (13) and quadratic multiple regression models were used
to calculate quadratic selection gradients (y), Linear selection gradients were used
to estimate directional selection and quadratic selection gradients were used to
estimate variance selection. These selection gradients only provide crude estimates
of the form that selection takes (Mitchell-Olds & Shaw 1987) but small sample
sizes prevented the use of more sophisticated techniques (eg. spline analysis;
Schluter 1988). Linear selection gradients were estimated in separate analyses from
those used to estimate quadratic selection gradients because linear and quadratic
variables were correlated (Lande & Arnold 1983; Wade & Kalisz 1990).
Significance testing of multiple regression analyses assumes that the data are
parametric and that the variables are uncorrelated. These assumptions· did not
always hold for my data, so the use of F-statistics to test the significance of the
selection gradients is not strictly valid (Lande & Arnold 1983; Endler 1986;
,..INDERS UNIVm&IJY
LIBRARY
217
Chapter 7 TARGETS OF SEXUAL SELECTlON
MitcheJl..Olds & Shaw 1987). Instead, Speannan's partial correlation coefficients

were used to test whether selection gradients were significantly different from zero
wherever possible. For most analyses, only parametric tests of significance were
available on the software I had available. I addressed the problem of correlations
between the variables by reducing the original set of traits to a smaller set of
orthogonal variables derived from a Principal Components Analysis (Lande &
Arnold 1983; Endler 1986). Principal components (PC) scores are uncorrelated
summaries ofvariation in the original traits. I then estimated selection gradients for
the PCs.
The small sample sizes, coupled with the large number of variance and covariance
terms required for a quadratic model, precluded estimation ofvariance selection o...
each individual trait. Therefore quadratic selection gradients were not estimated for
raw traits, but only for the PCs derived from them.
In the analyses ofboth cohorts SVL, Mass, HL, HW and HD were included as
independent variables. HR was included in the analysis of the 1991 cohort, but not
the 1990 cohort, for which insufficient measurements ofthis variable were
available. Separate sets of PCs were extracted from the phenotypic covariance
matrix for all males with complete data for each cohort (N = 32 for 1990 cohort; N
= 19 for 1991 cohort).
Results
All five morphological traits were highly correlated with each other (Table 7.1).
Home range area showed a less strong, but nevertheless significant, positive
correlation with all of the morphological traits (Pearson's r = 0.32-0.45, all P's<
218
Chapter 7 TARGETS OF SEXUAL SELECTION
0.02). The correlations among these variables confirmed the need to calculate
selection gradients to identify traits which are direct targets of selection.
1990cohort
The 1990 cohort showed a significant sexual selection for HD.' Paired males had
larger headsJ.t1tan males from the whole population. Paired.males were also
significantly less variable in SVL and HD, and had more variable HLs, than all
males. However, no single trait was identified as a significant target of direct
..,
selection, since none showed a significant selection differential.p (Table 7.2).
V
The first two PC's from an analysis of males in the 1990 cohort were interpretable
in terms ofilieir relationship to the original traits (Table 7.3). The first PC was
interpreted as a measure of overall size because SVL, Mass, HL and HD had their
highest loadings on this component. The second PC reflected variation in head
length because it was defined by a high loading for HL. Together these two PC's
accounted for 95% of variance among inilividualmales. A significant negative
directional selection gradient was detected on PC2 Table 7.3). This may reflect the
smaller HL values fur paired males (Table 7.2) in 1990. No directional selection
was detected on PC 1, and neither PC was subject to variance selection.
Selection differentials suggest that both directional selection and disruptive

selection were acting on HL (Table 7.2). The signific~t linear selection gradient
s
on PC2 also sugge~at sexual selection may have been acting to reduce HL in
this cohort. This selection must have been weak because no single raw head
measurement had a significant selection gradient (Table 7.2).
1991 cohort
None of the traits showed any significant directional selection differential among
males of the 1991 cohort (Table 7.2). But all morphological traits measured
showed significant reduction in variance among paired males. Home range area did
not show any significant relationship to fitness. The lack of any significant
directional selection differentials may be due to the interrelation among these traits
(Table 7.1). Significant linear selection gradients for SVL and HW suggested that
directional selection is acting independently on these traits (Table 7.2).
The first PC from an analysis of males in the 1991 cohort was interpreted as a
measure of overall size because all five morphological traits had high positive
loadings on this component (Table 7.4). The second PC reflected variation in
home range area because it was defined by a high loading for HR. Together these
two PC's accounted for 97% of variance among individual males. A linear
selection gradient that was very nearly significant was detected on PC 1. But
neither of the quadratic gradients or the directional gradient on PC2 approached
significance (Table 7.4).
The combination of the significant positive directional selection, significant

negative variance selection on all morphological traits, combined with the ahnost
significant positive directional selection gradient on PC l (overall size) demonstrate
that sexual selection was acting on body size in this cohort. The significant positive
selection gradient on HW (Table 7.2) also suggests that selection was acting
independently to increase head size.
220
Chapter7 TAR.GETS OF SEXUAL SELECTION
The selection analyses gave quite different results for each cohort. Only weak
selection acting to reduce BL could be detected in males ofthe 1990 cohort,
whereas there was clear evidence for sexual selection acting to increase SVL and
HW independently in males of the 1991 cohort.
MECHANISMS OF SELECT10N
Methods
To examine the potential of male competition to exert the observed phenotypic

selection I (1) compared the morphological measurements for the tenitorial and
for nonterritorial males, and (2) staged encounters between pairs of males in
neutral arenas to determine if any morphological traits were associated with
winning aggressive interactions. Territorial males defended an area of the study site
against other mature males. Nonterritorial males usually lived within territories .of
other males by adopting subordinate behaviours (Chapter 6).
Morphology and territ.ory status
I used 71 males for which all morphological traits had been measured. I extracted
four PCs from these measurements to obtain orthogonal summaries of the factors
associated with tenitorial status. Univariate oneway ANOVAs were used to test
whether territorial and nonterritorial males differed either in raw measurements or
in scores from the four PCs.
221
These are not tests of the significance of selection coefficients for territorial status
because they are based on cross-sectional data (Lande & Arnold 1983). They
compare different aged males with and without territories at one point in time.
They may, however, indicate the general importance of the traits to territory status.
Staged male encounters.
Fighting between males during territory establishment and maintenance involves

ritualised displays (Gtobons 1979) and physical combat. This includes biting and
reciprocal jaw locking. I staged 22 male-male encounters in neutral arenas, using
males captured at the two study areas. Individual liz.ards were tested once only
during November 1995. The same morphological measurements used in the field
study were taken for the 44 liz.ards used in the staged encounters.
The arenas were two glass terraria measuring 300x740x350mm which were
divided in halfby a removable cardboard partition. Both terraria had 20mm of sand
on the bottom. A 150W light globe was suspended over each half of the arena, so
that each liz.ard could thermoregulate to its preferred temperature. Each trial was
video-taped so that liz.ard behaviour was not affected by the presence of an
observer. Video tapes were viewed at a later date. Two liz.ards were placed in the
arena, on either side of the cardboard partition, for one hour before the trial began.
Removal of the cardboard marked the start of the trial, and each trial lasted for 30
minutes. Losers in the encounter were lizards which retreated or lay prostrate on
the sand in response to displays or attacks by their contestant. There was a clear
winner and loser in sixteen of the 22 trials.
222
Four PCs were extracted from measurements of the 44 males used in the trials, to
obtain independent measures of the factors associated with winning a fight.
Univariate oneway ANOVAs were used to test whether the means of the raw
measurements and the four PC scores differed among winners and losers in the
sixteen trials in which there was a clear winner.
Results
Morphology and territory status
Data pooled across all years showed that territorial males were significantly larger
than non-territorial males for all five morphological traits(Table 7.5). This
suggests that overall size may be an important factor in gaining and maintaining a
territory. This impression was confirmed when the effects ofoverall size (PCl in
Table 7.6) was found to differ significantly between territorial and nonterritorial
males (Table 7.5), whereas three PCs representing various aspects ofhead size and
shape (Table 7.6) did not differ between the two groups of males (Table 7.5).
Staged male encounters.
Sixteen (73%) of the staged encounters involved some display or combat

behaviour. Displays conformed to previous descriptions-ofthe hindleg pushup
display that is characteristic of rock dragons (Gibbons 1979). Males lowered their
throats while still at some distance from each other. They then moved toward each
other and alligned themselves laterally. Once alligned they performed a variable
number ofhindleg pushups. Some liz.ards circumducted the forelimbs or performed
head bobs in between each hindleg pushup. A few contests progressed to physical
223
Chapter 7 TARGETSOFSEXUALSELECfION
combat, which involved biting, and throwing each other bodily around the
enclosure.
Males which won in staged contests were significantly larger than losers for four of
the five morphological traits (Table 7.7). There was no difference in IID between
winners and losers. These results suggest, once again, that overall siz.e may be an
important factor in gaining and maintaining a territory, at least insofar as male
combat determines territory ownership. A comparison of PC scores between
winners and losers showed significant differences in two separate PCs. The first of
these (PC 1) represented overall siz.e, and the other (PC3) represented variation in
HW (Table 7. 8). Winners had larger values in both cases. This result suggests that
overall body siz.e and HW have separate roles in determining whether a male is
likely to win or lose a contest.
DISCUSSION
This study is based on two assumptions. The first is that pairing in Peninsula
Dragons is correlated with mating. Pairing, rather than direct observation of
matings, has been used to estimate mating success in several other studies of
lizards (Ruby 1981; Bull 1988; Bull & Pamula 1996) and is usually regarded as
mate guarding behaviour (Cooper & Vitt 1993; Olsson 1993b). When data have
been available on both mating and pairing success, the results using each as a
measure of fitness have not differed appreciably (Ruby 1981). The efficacy of using
pairing as a measure of mating success in Peninsula Dragons has been discussed in
Chapter 3.
224
Cbapter7 TARGETS OF SEXUAL SELECTION
The second assumption is that the chance of finding any male in a pair is equal
among all males. This may not be strictly true for my data. Some males could be
relocated at will because they were large enough to be radio-tracked, whereas
those lizards which were too small to have transmitters attached could not be
located as readily. Thus, even iflarger and smaller males paired equally frequently,
there may have been a greater chance of finding larger lizards in a pair. An attempt
was made to redress this potential bias by daily systematic searches of the study
areas, paying particular attention to known home ranges of smaller males. Most of
these yearling males were located on each search and all were located every 2-3
days. Given that Peninsula Dragons pair for an average of two days, and up to ten
days (Chapter 3), the likelihood of missing a pairing was minimised.
All males in this study had numbers painted on ,their backs and were toe-clipped.
These methods of marking might be expected to affect survivorship, and chances
of gaining or defending a territory. However, several studies have found no
reduction of survivorship due to paint-marking lizards (Jones & Ferguson 1980;
Simon & Bissinger 1983 ). Similarly,. toe-clipping does not appear to influence
survival in lizards (Huey, Dunham, Overall & Newman 1990; Dodd 1993). The
effect of the marking teclin.iques used here should not be important for this study
because all lizards were marked in the same way, so any effect should be constant
across all individuals.
Targets of Selection
The two cohorts studied showed very different patterns of selection. The 1990
cohort exhibited evidence for weak selection for shorter heads. This directional
225
Chapter .7 TARGETS OF SEXUAL SELECTION
selection may have caused the reduction in variance for HD. The reduced variance
'in SVL may be the result ofindirect selection, arising from the correlation between
SVLandHD.
The significant positive variance selection differential for HL in the 1990 cohort
suggests that this trait is subject to disruptive selection. When variation in HL was
dissociated from its correlation with overall body size using principal components
analysis, HL v;as found to be subject to significant negative directional selection
(Table 7.3). This might account for the disruptive selection acting on this trait.
Through positive correlations between HL and all other traits, selection may favour
liz.ards with long heads. This indirect selection for longer heads may be opposed by
direct selection for shorter heads. These opposing selective forces coll;ld result in a
net disruptive selection acting on HL. The lack of any significant directional
selection gradients on the raw traits may be due to the low sample size and
consequent lack of statistical power, or alternatively the significant selection
differentials may be the result of selection acting on unmeasured, but correlated
traits.
The 1991 cohort showed clear evidence that sexwtl selection was acting to
increase body size and HW. Although none of the directional selection differentials
were significant, all morphological traits showed significant stabilising selection
(Table 7.2). The lack of significance among the directional selection differentials
was probably due to the correlations among the traits. The stabilising selection
presumably resulted from directional selection for a large body size and a wide
head because both SVL and HW were identified as direct targets of selection
(Table 7.2 & 7.4). There was no evidence that home range area affected a male's
reproductive success.
226
Cbapter7 TARGETS0FSEXUALSELECI10N
Since the two cohorts were studied at the same time and at the same place, these
differences in selective regime do not represent temporal variation in the action of
selection. If the apparent differences in selection between the cohorts is not a
statistical artefact, they may be a result of age-related changes in selection. Males
of many species of lizard exln'bit changes in reproductive tactics as they grow
(Hews & Moore 1995; Baird, Acree & Sloan 1996). Yearling male Peninsula
Dragons do not hold territories, but live in small home ranges within the territories
of other, older males (Chapter 6). As they grow these younger males then
challenge older males for possession of a territocy. Episodes of selection that occur
prior to or upon achieving maturity, and which are associated with the initial
gaining of a territory, could not be included in the analyses. It is posst"ble,
particularly for traits such as body size, that natural selection acting on juveniles
might counter or enhance the sexual selection that was measured in this study. For
instance gaining a territory may be important for future reproductive success
(Stamps 1983) and have an influence on adult behaviour, even though the size of
the territocy was not identified as a target of selection.
It is noteworthy that the youngest cohort of males was the one in which the action
of sexual selection was most clearly apparent. Males ofthe 1990 and 1991 cohorts
were in their third and second breeding seasons, respectively, in 1992. All males in
this study had already obtained territories by 1992, so the most obvious episode of
selection (gaining a territocy) was excluded from this study. However, the 1991
cohort had more recently gained territories. So size may have been important,
particularly since they were competing among their own cohort and the older
males from the 1990 cohort. The selection coefficients recorded here may be
viewed as being conservative estimates of the importance ofiotrasexual selection.
227
Cbapter7 TAROETSOPSEXUALSELECrION
Toe three PC analyses derived wholly from morphological traits produced similar
results. In each analysis PC I had consistently high positive factor loadings for all
variables, but particularly for SVL and Mass (Table 7.3, 7.6 & 7.8). These PCs
were interpreted as representing overall body size. Subsequent PCs in all of the
analyses had their highest loadings associated with various head dimensions. This
pattern of covariation among morphological traits is consistent with the
observation that males differ from females in body size and relative head size
(Chapter 2).
Most (95%) of male Peninsula Dragons hold territories, and are thus able to pair,
for one or two years, when they ai:e two and three years of age (Figure 6.1 ). Thus,
in so far as pairing reflects mating success, the estimates of fitness used in this
study approximate lifetime reproductive success for these lizards.
Modes of Sexual Selection
Territory ownership is a prime determinant of reproductive success in lizards

(Stamps 1983). In Peninsula Dragons 96% of pairs were between a territorial male
and a female that overlapped his territory (Chapter 6). Female Uta palmeri, a
species in which males exlnl>it resource defence (Hews 1993), also seldom mate
with non territory holding males (Hews 1990). Conversely, male Lacerta agilis do
not defend exclusive home ranges, and females commonly mate with multiple
males (Olsson, Gullberg, Tegelstrom, Madsen & Shine 1994). In lizards possession
of a territory appears to give a male priority access to females.
228
Chapter7 TARGETSOFSEXUALSELECI10N
Fighting ability could determine a male's ability to gain and defend a territory, and
could, therefore, be a proximate mechanism of sexual selection. There is some
evidence in other lizards that females choose to mate with large males (Cooper &
Vitt 1993; Ellington 1993), and reject the advances of smaller males (Baird et al.
1996). Both male competition and female mate choice are potential mechanisms of
sexual selection in Peninsula Dragons.
Both body size and head size were foimd to be larger among territorial males. Both
were also important determinants of which lizard won staged male fights. Male
competition appears to be adequate as an explanation for selection on these traits,
without the need to invoke female mate choice. Since overall body size and head
shape were identified as independent targets of sexual selection they must be
subject to different sources of selection.
Size has been shown to be an important determinant of social dominance in lizards

(Tokarz 1985; Hews 1990; Olsson 1992), and many other organisms (eg. Reichert
1978, 1984; Clutton-Brock, Guiness & Albion 1982; Enquist, Leimar, Ljungberg,
Mallner & Segerdahl 1990; Leimar, Austad & Enquist 1991). Selection for body
size may arise because it confers a direct advantage during agonistic encounters
(Tokarz, 1985; Olsson 1992), or simply because larger males are more obvious and
less likely to be challenged.
Selection for larger HW is independent of selection for body size. Therefore the
mechanism of selection for greater HW must be different from the mechanism
leading to selection for SVL. Comparative studies have among lizards have shown
that male head size is greater in lizards which are highly polygynous, than in those
in which male competition over females is less severe (Carothers 1984). A deeper
head is associated with greater mating success, and confers an advantage in male-
229
male contests in Uta pafmeri (Hews 1990). But these explanations are not
independent of the role that body size plays in male competition.
There are several explanations for the identification ofhead climensions as

independent targets of selection: ( 1) BW may still be an indirect target ofselection
if statistical assumptions are violated; (2)males with greater BW may be better
able to maintain their grip on females while mating (Hews 1990); (3) head depth
may confer a mechanical advantage in male-male contests that progress to bites;
(4) females may choose males with large heads.
Carothers (1984) documented an association ofpolygyny and accompanying high

levels of male-male aggression with sexual climorphism in head size in iguanid
li7.ards. He proposed that the selective advantage oflarge male head size was to
increase gape, and thus bite strength. Male Eumeces laticeps undergo hypertrophy
ofthe jaw muscles during the breeding season. This led Vitt &. Cooper (1985) to
suggested that increased head size in male skinks is favoured by intrasexual
selection. While it is not p0SS1'ble to manipulate head width experimentally in
li7.ards. it should be posSI"ble to determine whether greater head size is associated
with the increased or more effective biting in contests between males. Bites
certainly occurred in the trials reported here, but the low .sample sizes precluded
separate analysis of contests that did and did not progress to bites. The
demonstration that greater BW provides some mechanical advantage in severe
fights would not exclude the posstlile role for HW in holding females during
mating.
If females choose mates·based on morphology and if selection is strong enough to
be detected by this analysis, the preferred trait should be identified as a target of
230
Cbapter7 TARGETS OF SEXUAL SELECTION
selection. For the traits that were not identified as direct targets of selection., the
null hypothesis ofno female choice could not be rejected.
The selection analysis provided no evidence that female choice was acting on any
male trait in the field. Selection on both SVL and head dimensions could be
accounted for by intrasexual selection. 11tis may be because there is limited
opportunity for females to express mate choice in the field. Several males may
overlap a female's home range (Chapter 6), but her ability to choose among them
may be limited because a large territory holding male often accompanies her when
she ovulates (Chapter 3). This may provide the male with exclusive mating access
to the female.
In summary, this chapter confirms that sexual selection is acting on body size and
shape in male Peninsula Dragons. Body size and head width were found to be
independent targets of sexual selection in the younger of two cohorts studied at the
same time. Head length was under weak selection in the older cohort. Body size
and head width were important correlates of territorial status and whether a male
won or lost in staged fights. These findings suggest that sexual selection acting on
male morphology in this species can be adequately explained by male competition.
I found no direct evidence that female mate choice was acting on the traits
measured in this study.
231
Chapter? TARGETS OF SEXUAL SELECTION
Trait
Trait SVL HW HL HD
SVL 1.00
Mass 0.98 l.00
HW 0.94 0.94 1.00
HL 0.76 0.76 0.78 1.00
HD 0.90 0.93 0.87 0.70 1.00
Table 7,1: Correlations among measured body traits for 109 male Cte,wphorus
fionni. All correlations are significant (P < 0.001).
232
(")
All males Paired Males

Trait n x(SD) n x(SD) j
1990 col,ort
Snout-veot length (mrn) 36 80.8(3.39) 9 81.8(1.91) 0.087 -0.683*
Mass (gm) 36 23.0(2.61) 9 23.6(2.13) 0.088 . -0.334
Head width (mrn) 34 2!.0(I.17) 8 21.3(1.04) 0.219 -0.21
Head length (mrn) 34 19.5(3.56) 8 17.7(7.08) -0.142 2.955**
Head depth (mm) 34 9,7(0.63) 8 9.9(0.35) 0.504** -0.691 *
.,,..., 1991 col,ort
Snout-vent length (mm) 27 73.7 (13.04) 9 80.6 (2.75) 0.041 -0.956** 3.560•
"' Mass(gm) 27 18.2 (7.04) 9 22.1 (2.01) 0.079 -0.919••
Head width (mm) 24 19.3 (3.41) 9 21.l (0.78) 0.155 -0.948** 4_339•
Head length (mm) 24 18.2 (3.15) 9 19. 7 (1.00) 0.151 -0.899**
Head depth (mm) 24 8.7 (l.34) 9 9.2 (0.67) 0.279 -0.750*
Home range area (ha) 21 0.04 (0.07) 7 009(0.09) 10.204 0.653
• P< 0.05; ** P<0.01
Table 7.2: Mean (and standard deviation) of trait values, standardised directional differentials (i), standardised quadratic differentials (j) and
directional selection gradients (~) due to pairing success in two cohorts of male Ctenophorus ftonni at Midgee Rocks during 1992 and 1993.
Only significant directional selection gradients are shown.
Chapter 7 TARGETSOFSEX:UALSELEcrION
1990COHORT
Factor loadings
Trait PCl PC2
SVL 0.977 0.116
Mass 0.974 0.104
HW 0.968 0.101
HL 0.714 0.698
HD 0.910 0.204
J3 0.116 -0.381*
'Y -0.104 0.015
Table 7.3: Factor loadings of each trait on the first

two principal components (PC), with partial linear (13)
and partial quadratic (r) selection gradients for these
composite variables for males from the 1990 cohort.
For the linear modelK = 0.14, F2.31 = 2.41, P= 0.106
and for the quadratic modelK = 0.10, F2,31 =1.795, P
= 0.183. For each component, the traits with the
highest factor loadings are underlined.* P < 0.05.
234
1991 COHORT
Factor loadings
Trait PCI PC2
SVL 0.990 -0.063
Mass 0.988 -0.063
HW 0.988 -0.033
ID., 0.983 -0.110
HD 0,930 -0.166
HR 0.476 0.878
J3 0.4581 0.167
y -0.170 0.087
1 P=0.052
Table 7.4: Factor loadings of each trait on the first

two principal components (PC), with partial linear (13)
and partial quadratic (y) selection gradients for these
composite variables for males from the 1991 cohort.
For the linear model If= 0.24, F2,16 = 2.50, P= 0.114
and for the quadratic model R2 = 0.21, F2.16 = 2.094, P
= 0.156. For each component, the traits with he highest
factor loadings are underlined. None of the selection
coefficients were significant.
235
Chapler7 TARGETS OF SEXUAL SELECTION
x(SD)
Territorial N onterritorial
Traits (n= 47) (n=24) F
SVL 81.0(3.163) 59.6(6.147) 379.08*
Mass 22.8(2.288) 9.3(2.495) 520.76*0
HW 21.0(0.889) 15.5(1.841) 297.77*
HL 19.5(3.000) 14. 7(1.628) 52.39*
HD 9.4(0.586) 6.8(0.637) 286.39*
PCI .51(0.316) -1.47(0.468) 449.73*
PC2 -0.10(1.434) 0.24(0.491) 1.25
PC3 -0.26(0.929) 0.05(1.184) 1.37
PC4 -0.02(0.829) 0.23(1.164) 1.09
Table 7.5: Mean·(and standard deviation) of trait values for territorial and
nontenitorial male Ctenophorus jionni at Midgee Rocks during 1991-
1994. F= univariateF-statistic; * P < 0.001.
236
Chapter7 TARGEI'SOFSEXUALSELECI10N
Factor loadings
Trait PCI PC2 PC3 PC4
SVL 0.976 -0.116 -0.094 -0.127
Mass 0.980 -0.124 -0.037 -0.110
HW 0.964 -0.040 -0.189 0.184
Ill, 0.841 0,538 0.057 -0.014
HD 0.938 -0.191 0.280 0.070
% 88.6 7.1 2.6 1.3
cumulative% 88.6 95.7 98.6 99.6
Table 7.6: Factor loadings of each trait on the first four principal components (PC)
calculated for 71 male Ctenophorus ftonni measured at Midgee Rocks during the springs of
1991-1994. For each component the traits with the highest factor loadings are underlined;
% is the p ~ t ofvariance explained by each.factor.
237
Chapter 7 TARGETS0FSEXUALSELECI10N
x(SD}
, Winner Loser
Traits (n= 16} (n=l6) F
SVL 73.7(3.141) 69. 7(5.687) 6.238**
Mass 15.2(1.078) 13.3(3. 785) 3.631*
HW 18. 7(0.855) 17.8(1.625) 4.004*
HL 17.8(0.871) 16.9(1.619) 3.730*
HD 7.9(0.411) 8.0(2.646) 0.068
PCl 0.28(0.547) -0.43(1.284) 4.053*
PC2 -0.10(0.198) 0.31(1.580) 1.055
PC3 -0.32(0.320) 0.08(0.710) 4.260**
PC4 -0.04(0.819) -0.00(0.962) 0.009
Table 7.7: Mean (and standard deviation) of trait values male Ctenophorus
ftonni which won and lost in staged contests in neutral arenas. F =
univariate F-statistic; *P < 0.07; ** P < 0.05.
238
Factor loadings
Trait PCl PC2 PC3 PC4
SVL 0.956 -0.103 -0.096 0.029
Mass 0.910 0.026 -0.302 0.251
HW 0.794 -0.285 0.530 0.072
HL 0.925 -0.097 -0.131 -0.330
HD 0.442 0.883 0.154 -0.017
% 68.5 17.6 8.4 3.6
cumulative % 68.5 86.l 94.6 98;1
Table 7.8: Factor loadings of each trait on the first four principal components (PC)
calculated for 44 male Ctenophorus fiormi used in staged male contests in neutral arenas.
For each component the traits with the highest factor loadings are underlined. % is the
percent ofvariaµce explained by each factor.
239
CHAPTERS
MATE CHOICE OR MALE COMPETITION ?
It is over one hundred years since mate choice by females was first proposed as
a mechanism for the evolution of elaborate male traits (Darwin 1871). The
relative importance of the intrasexual (male competition) versus intersexual
(mate choice) modes of sexual selection has remained controversial for most of
this time. This is true for biologists generally (Thornhill & Alcock 1983;
Bradbury & Davies 1987; Andersson 1994), and for those who work on lizards
in particular (Olsson & Madsen 1995; Tokarz 1995).
The controversy has been based on different issues through its long history.
Initially, the debate centred on whether nonhuman organisms had the aesthetic
senses that seemed to be implied by an ability to choose among mates
(reviewed by Cronin 1991). More recently the controversy has arisen from,
among other things, the difficulty in separating mate choice by females based
on phenotypic characteristics of males from choice based on resources
controlled by males ( eg. Alotalo, Lundberg & Glynn 1986). This difficulty
Chapter g MATE CHOICE?
comes about when a trait is correlated with success in gaining both mates and
resources concurrently. In these circumstances intrasexual and intersexual
selection are not mutually exclusive and may occur simultaneously. When this
occurs, the interactions may be complex ( eg. Petrie, Halliday & Sanders 1991;
Sorenson & Derrickson 1994). In species with mating systems which are based
of resource defence, as in IllllllY lizards (Stamps 1983), it is important to
separate mate choice based on a phenotypic trait from a preference for
resources over which potential mates may have competed.
Active mate preference by females has been invoked to explain male body size
and colour in lizards fur many years (eg. Noble & Bradley 1933; Tokarz
1995). Many studies which pmport to demonstrate mate choice in lizards have
actually only demonstrated the importance of elaborate traits for passive mate
recognition (see Chapter 1). For example, male lizards attack females that are
painted with male colours (Noble 1934; Kramer 1937; Kitzler 1941; Werner
1978; Cooper & Bums 1987) and court males painted with female colours
(Noble 1934; Kramer 1937; Kitzler 1941; Harris 1964; Werner 1978; Cooper
& Bums 1987; Cooper & Vitt 1988).
Few studies have explicitly tested the idea that females distinguish among
conspecific males from within the normal level ofvariation in a trait. Females
of at least two species of lizard are able to choose among males. Female
Broad-headed Skinks (Eumeces laticeps) preferred larger males over smaller
I
males (Cooper & Vitt 1993). Female geckoes (Gonatodes albogularis)
preferred males with more brightly coloured heads over those with duller
coloured heads (Ellington 1993). The general lack of evidence for active mate
preference in other lizards may result from a lack of cues which give female
lizards a reliable indication of male quality (Olsson & Madsen 1995).
241
Chapter 8 MATE CHOICE?
Alternatively, the apparent rarity of mate choice in lizards may be a
consequence of the paucity of appropriately designed studies, or the difficulty
of observing normal behaviour.
Sexual selection may explain the sexually dimorphic traits in Peninsula Dragons
because males have higher variance in mating success than females (Chapter
6). The home ranges of female Peninsula Dragons may overlap with those of
more than one adult male, providing an opportunity for females to express
mate choice (Chapter 6). An analysis ofbody measurements in males indicates
that body size and head size are independent targets of sexual selection, but
this appears to result from intrasexual selection due to male combat over
territories (Chapter 7). Female mate choice may be acting on male colouration
because adult males are more brightly coloured than females or immature
males (Chapter 2).
Sexual dichromatismis widespread among diurnal lizards (Stamps 1983;
Cooper & Greenberg 1992). However, no previous studies appear to have
tested whether colour is associated with competitive ability or mating success
in reptiles by manipulating male colouration in the field. Such manipulations of
colour are essential to control for other factors that may be correlated with
colour ( eg. age, size). They are crucial for demonstrating that colour itself
functions as a signal and serves a direct role in communication ( eg. Butcher &
Rohwer 1989; Rohwer & Roskraft 1989).
Under ideal conditions one could subject colour traits to a selection analysis,
similar to the one carried out in Chapter 7 for morphological measurements.
This would allow one to identify which particular aspects of colour pattern
were targets of sexual selection. These could then be investigated
242
experimentally to identify the mechanism(s) by which selection was acting on
them. 1bis was not possil>le in the present study for two reasons. First, colour
is notoriously difficult to quantify objectively, and the complexity of the colour

pattern in Peninsula Dragons would require quantification of many traits to
descnl>e it adequately. Secondly, the low number of males for which
reproductive success could be estimated within each cohort precluded a
selection analysis of such a large number of traits. Instead, I experimentally
manipulated colour patterns of males in the field to investigate whether sexual
selection acts on this trait.
1bis chapter asks the following questions: (1) Do female Peninsula Dragons
show preferences among males? (2) Is sexual selection acting on male
colouration in Peninsula Dragons? and (3) What mode(s) of sexual selection
are acting on male colouration ? It descn"bes two experiments. The first
experiment was aimed to test whether female choice occurs in C. fionni under
laboratory conditions, given that inclement weather, the distnl>ution of males,
and other factors may limit the potential for female choice to occur in the field
(Chapter 6). The second experiment was designed to test whether female
choice and/or male-male competition were acting on the male colouration in
the field.
METHODS
Laboratory Experiment
Fourteen females were each given the choice between two males. The fourteen
trials were run concurrently over five days between 14th and 18th August
243
1991. All females had palpable, enlarged ovarian follicles. They had been kept
isolated from males in outdoor enclosures for one month, so they could not
mate prior to the experiment. In each trial one male was from the same
population as the female and the other was collected at least 200km away. This
was done to increase the difference in colour pattern between the males
because males from geographically separated populations differ in colour
pattern (Houston 1974). It therefore maximised the likelihood that the females
would make an active choice between the males if colour were important. The
collection locality and SVL of all males used in these experiments are recorded
in Table 8.1. This experiment does not preclude the possibility that females are
choosing males based on a trait other than colour (such as size or odour),
which may also differ between populations. Rather, the aim was to test
whether females discriminate among males.
The experimental enclosures consisted of glass terraria measuring 300mm x
740mm x 350mm which were split into three compartments with transparent
acrylic dividers (Figure 8.1). One compartment ran the length of the enclosure
and along one side of each of the other two compartments. A female was
placed into the long compartment. She was able to see both males in the other
two compartments through clear acrylic at each end, but could choose to stay
in the central part of the compartment, where she was shielded from both
males by an opaque divider. The divider between the male's compartments was
opaque to avoid the males reacting to each other in any way that might
influence the female's behaviour. All outside walls of the experimental
enclosure were also opaque so that lizards in each experiment could not see
each other. All experimental enclosures were inside a room at temperatures of
20-25°C. A 100W light globe was suspended 10cm over the intersection of the
dividers to provide adequate heat so that all three lizards in each cage could
244
Chapter 8 MATE OIOICE?
thermoregulate and achieve their preferred body temperature. light globes

were turned on at 0600 and turned off at 1830. Neither water nor food were
provided during the five days of the experiment.
The location of each female was checked hourly between 0700 and 1800 local
time for five days ( except for one trial in which four observations were
excluded because a female escaped temporarily) to provide a total of836
observations. Each female was scored as being adjacent to one male, adjacent
to the other male, or in the central section ofher compartment and unable to
see either male (see Figure 8.1). To be in visi'ble contact with either of the
males in her cage a female had to move away from the heat source. Males were
swapped between compartments mid-way through experiment (ie. at 1200 un
the third day of the experiment) to allow a distinction to be made between
females that preferred one end of the experimental enclosure, from females that
preferred to associate with a particular male.
The number of times each female lizard was in visual contact with each male
was used to calculate G statistics (Sokal & Rohlf 1981). The data were tested
against the expectation that females would associate with each male equally,
under .the null hypothesis that females showed no preference for either male.
Field Experiment
This study was conducted at Midgee Rocks (Figure 1.2 & 1.3) during August
and September of 1992 and 1993. An experimental group of fifteen male C.
ftonni (6 in 1992 and 9 in 1993) were painted with brown, non-toxic, pliable,
245
acrylic paint to obliterate the distinctive dorsal, head and chest colouration.
After painting they resembled immature males and females to the human eye. A
control group of fourteen males (6 in 1992 and 8 in 1993) were painted with
non-toxic, pliable, clear lacquer, so that the male colouration remained visible.
Cate was taken to avoid painting over the femoral pores, so that olfactory
signals were not occluded. The home range area and the number of females
associated with each male were recorded during a ten day period before the
manipulation and then for another ten days immediately aiier the manipulation.
Each male was weighed at the beginning and end of the experiment.
Transmitters were attached to the side of each male's tail at the base, using
quick-curing araldite. Each transmitter weighed 2.0g and represented on
average 7% of each li:zard's body weight at the beginning of the experiment.
Lizards were located one or two times per day depending on weather
conditions. Each liz.ard was located at least fifteen times in each ten day period.
This is the number oflocations required to provide an estimate of at least 90%
of the home range for male Peninsula Dragons at this ·site following Rose
( 1982). Home range area was measured as the mininmm convex polygon
(Jennrich & Tomer 1969) based on all locations at which each liz.ard was
found by radiotelemetry.
I used pairing between adult female and male li:zards as an index of mating
success because I made no direct observations of actual matings (see Chapter 3
for discussion). Paired lizards were found within I.Om of each other while
active, or were found in bodily contact inside a rock crevice when inactive. All
C. fionni encountered during the study were individually marked by clipping
up to two toes from each foot in unique combinations (Fenner 1979) to allow
identification of females that paired with the males under study.
246
Chapters MATE CHOICE'!
Iffemale choice alone were acting to select for male colour one would expect
a reduction the number of females that paired with experimental males, but no
differences in home range size and weight between experimental and control
males. Ifmale-male competition alone were acting to maintain male colour one
would expect no difference in the number offemales associating with control
and experimental males, but home range size and weight of experimental males
would decrease in experimental males. Ifboth female choice and male-male
competition were acting in concert to maintain male colour, one would observe
a reduction in the number of females associating with experimental males, and
a r~ion in home range size and weight in experimental males compared
with control males.
In two separate analyses the number of females paired with each male and the
home range area of each male were used as dependent variables in repeated
measures two-way Analyses of Variance with the Treatment ( control vs
experimental) and Time (premanipulation vs postmanipulation) as fuctors. In a
third analysis the percentage of original mass lost by males in each treatment
group was analysed using a oneway analysis of variance. These percentages
were arcsine transfonned before the analysis.comparing control and
experimental groups (Sokal & Rohlf 1981). ANOVAs were deemed
appropriate because the between groups variances were homogeneous
(Cochrane's C, P's> 0.05) in each case.
247
RESULTS
Laboratory Experiment
Although the pooled data showed no evidence of mate choice overall (pooled
G = l.066, df=l, P > 0.05), individual females varied in their behaviour
(heterogeneity G = 36.583, df= 13, P < 0.01). This heterogeneity was mainly
due three females that exlumted a significant preference for one of two males
(Table 8.1; individual G statistics, P's< 0.05). Two of the three females that
showed significant choice, associated more frequently with the male that was
not from their own population, and that differed in colour from those the
female had previously experienced. This experiment suggests that there is little
evidence that female Peninsula Dragons consistently choose among males,
even when colour differences among the males are greater than normally occur
in a population. Only 20% of females showed any evidence for a choice.
Field Experiment
The mean number of females paired with each male (Figure 8.2) declined after
the manipulation for both experimental and control males , although this
difference was not quite significant (F1,2s = 3.23, P = O. 07). The experimental
males were consistenj;}y less attractive than control males, whether they were
painted or not, but the difference only just failed to be significant (Fus= 3. 74,
P = 0.059). There was no significant interaction between the :lilctors (F1,28 =
248
Chapters MATE CHOICE?
0.48, P= 0.89), so the decline in attractiveness between treatments was
similar.
For hollle range area (Figure 8.3) there was no overall difference between
treatments (F1:.zs = 0.096, P =0. 76), but there was a significant interaction
between tillle and treatment effects (Fus = 8.655, P = 0.007). A posteriori
tests showed that the significant difference due to time (F1:.zs = 9.897, P =
0.004) arose .from different responses by the two treatment groups. The
experimental males suffered a .significant· decrease in home range area due to

the manipulation (F1,13 = 20.96, P = 0.001), whereas the control group
retained similar home range areas before and after the manipulation (Fi.12 =
0.02, P = 0.89).
There was a significant difference between the two groups of males in the
proportion ofthe original body weight that was lost (F1,21 = 8.52, P = 0.008).
Over the period of the experilllent experimental ll13les lost twice as IDUCh
weight as the control males (Figure 8.4).
DISCUSSION
The laboratory study showed that only three out of fourteen female Peninsula
Dragons exluroted a preference,between males. Two of these fell13les chose to
associate with ll131es that did not coIDe their natal population. These males had
quite different colour patterns to males from the female's own population. The
variation in traits between these males was greater than would be available
within natural populations. This suggests that the Jevel ofvariation in colour
249
that males exhibit within populations may not be important for female mate
recognition.
The variation in the apparent degree of female mate preference may be an

artefact if some of the females used in my experiment were not receptive. I
attempted to ensure that all ofthe females I used were receptive by conducting
the experiment at the time ofyear when mating normally occurs (see Chapter
3), and by using females with enlarged, palpable ovarian follicles. Similar
variation in apparent female choice among males was found in Ano/is
carolinensis even though mate choice trials in this species were run over
several ovarian cycles (Andrews 1985). Andrew's resuhs were similar to my

own in that, even though a few female AMlis showed some evidence of mate
choice, her resuhs overall were not statistically significant.
The apparent lack of mate choice exhibited by some female lizards might also
be due to a lack of adequate variation in male traits, so that females are unable
to distinguish among males from their natal population (Andrews 1985). This
is unlikely in my study because the anatomy of Peninsula Dragon eyes suggest
that they are capable ofperceiving very small differences in body size, shape
and colour pattern (Johnston & Locket, unpublished data).
The field experiment showed that the male colour pattern played an important
role in territory maintenance among males, and that male colouration was
unimportant for attracting females in the field. Both control and experimental
males suffered a decline in the number of females with whom they were found
paired after the manipulation (Figure 8.2). This may have occurred because
females were avoiding males treated with lacquer and paint. This effect could
have been tested by including an unpainted group of male lizards in the
250
experiment. The difficulties of carcying out such a field experiment precluded

obtaining enough lizards to have included this extra control group.
Alternatively, the number ofpairs found may have dropped-off as a seasonal
effect because most females in the study area may have mated during the
earlier part of the experiment.
Ig1rnnian lizards have traditionally been regarded as using visual signals more
than olfactory ones (Cooper 1995). However, olfactory communi()l!tion based
on femoral gland secretions may be important in the social behaviour of
Iguanian lizards (Alberts 1992). There is certainly enough variation in the
chemical components of femoral pore secretions to allow population, sex and
individual recognition in this species (La Nafie 1995; La Nafie, Prager &.
Johnston 1995). Care was taken during the field experiment to avoid painting
over the femoral pores. Thus, in the field experiment olfactory cues were still
available for communication between the lizards. During the laboratory
experiment the sealed acrylic wall in the mate choice chamber precluded any
olfactory cues: Thus those females that showed a significant mate preference,
did so on visnal cues alone.
The male colour pattern appears to be relatively unimportant for sex and mate
recognition in female Ctenophorus fionni, but very important for recognition
among males. The field experiment shows that the colour pattern of male
Peninsula Dragons is important for maintaining a territory because the males
whose colour pattern was obliterated suffered a significant reduction in the
area of their home range, and lost twice as much weight as control males. This
weight loss may reflect extra costs for exprimental males of maintaining a
territory against increased incursions by neighbouring males, or lower feeding
opportunities in a smaller territory. The twofold difference in weight loss
251
between the control and experimental males is a measure of the advantage that
the male colour pattern confers. One interpretation is that male colour acts as a
signal which is effective in reducing challenges to male territories.
The colour pattern in Ctenophorus fionni emphasises the head, as does the
pattern of many other lizards (Cooper & Greenberg 1992). This is not
surprising given the importance of the jaws as a weapon in ma1e-male combat
(Hews 1990). Male colouration has been shown to be important in sex
recognition among male skinks ( Cooper & Burns 1987), but has been shown
not to alter female behaviour in other lizards (Vrtt & Cooper 1985; Cooper &
Vitt 1987). Male head colouration is known to be subject to female choice in
one species of lizard (Ellington 1993). Characteristic colours on other parts of
the body are probably important for sex recognition among females in other
species oflizards (Werner 1978; Cooper 1984).
In summary, the bright colouration of male Peninsula Dragons appears to

function in sex recognition among males. Colour does not appear to be a cue
for intersexual selection, but is an important determinant of whether a male can
maintain a territory, and therefore obtain priority mating access to females.
252
Choice
Trial Male 1 Neither Male2 G statistic
1 10 39 11 0,046
2 20 24 16 ·o.446
3 12 14 34 52.598*
4 8 37 15 2.166
5 5 53 2 l.330
6 19 27 14 0.760
7 25 27 8 9.194*
8 17 22 21 0.422
9 23 18 19 0.382
10 15 28 17 0.124
11 26 9 21 0.532
12 4 46 10 2.656
13 21 18 21 0.000
14 11 20 29 8.398*
pooled 216 382 238 1.066
TABLE 8.1: Choices shown by female Ctenophorus

fionni among two males. In all trials Male l was a male
from the same population as the female, and male 2 came
from a different population. The number of observations
in which females did not leave the central compartment is
shown under "neither". * P:,; 0.05.
253
MALEl MALE2
.
LOCATION OF FEMALE
I •
MALE I
r GLOBE II MALE 2
FIGURE 8,1: Enclosure used to assess mate choice in Ctenophorus jionni;

The thick lines represent opaque walls, and the thin lines represent transparent
walls. The enclosure was 740mm long and 300mm wide. A light globe was
placed over the intercept of the dividers for heat.
254
2.0
1.5 N=15
1.0 N=14
0.5
0
CONTROL EXPERIMENT
FIGURE 8.2: The mean number of female Ctenophorus ftonni that paired
with control and experimental males before (solid) and after (hatched) colour
manipulation. Vertical bars show one standard error either side of the mean.
255
1200
N= 14
--
N
e
.._,
fl;l
800
...
~
Ill
~
N= 15
c.,
z
<
~
~
0
400
0
CONTROL EXPERIMENT
FIGURE 8.3: Home range area among control and experimental male
Ctenophorus ftonni before (solid) and after (hatched) colour manipulation.
Vertical bars show one standard error either side of the mean.
256
20
N 14
00
00
0
~ 15
~
Cl
1-4
~
E,. N=15
~ 10
~
J;,;l
~
5
CONTROL EXPERIMENT
FIGURE 8.4: Weight change among control and experimental male ·

Ctenophorus ftonni during colour manipulation. Vertical bars show one
standard error above the mean.
257
'
CHAPTER9
SYNTHESIS
This thesis aimed to explain the .evolution of sexually dimorphic

traits in the Peninsula Dragon lizard ( Ctenophorus fionni). The
hypotheses that attempt to explain these traits include the
suggestion that they are sampling or demographic artefacts, and
that they may be the result of fecundity, natural or sexual
selection. Sexual selection is the most widely invoked
explanation for sexually dimorphic traits in other species. Indeed,
there is a tendency in the literature to assume that sexual
selection is the preeminant (if not the only) explanation for such
traits (eg. Bradbury & Andersson 1987). However, there are
sufficient cases in which sexual selection can not explain the
sexual dimorphism to suggest that other, alternative explanations
merit consideration ( eg. Shine 1989; Perez-Mellado & De La
Riva 1993).
Chapter 9 SYNTHBSIS
Adult male and female Peninsula Dragons differ from each other
in body size, shape and colouration. The sexes do not differ in
any trait before they become mature, but upon reaching maturity
males and females diverge from a common juvenile pattern. Males
exhibit hypermorphosis, growing to a larger size, at a faster rate
than females. Adult males also develop unique, sex-limited traits,
such as bright yellow throat colour. This constellation of sexually
dimorphic traits is common to many lizard species (Stamps 1983;
Hews 1996) and other taxa, and so conclusions drawn from
Peninsula Dragons may have general application.
This thesis shows that the sexually dimorphic traits in Peninsula

Dragons could not be explained as statistical or demographic
artefacts (Chapters 2 & 4). Nor could any of them be explained
by fecundity selection (Chapter 3) or natural selection (Chapter
5). The sexes utilised the space and thermal niche axes
differently, as predicted by the niche-divergence hypothesis.
However, these differences were interpreted as being a
consequence, rather than a cause of morphological differences
between the sexes. Males perched on higher rocks than females,
and had higher, more variable body temperatures than females
'(Chapter S). These differences probably reflect greater vigilance
for potential mates, and against rivals by territory holding males
than by females. Sexual selection acting on males appears to be
the sole explanation for the dimorphic traits in this species.
259
Chapter 9 SYNTHESIS
Peninsula Dragons have a mating system best described as resource

defence polygyny (Chapter 6; Emlen & Oring 1977). Rock crevices
were an important resource for males and females, because they
offered a refuge from both predators and extreme temperatures.
Males defended large home ranges (territories), which conferred
s
priority access to females. The number of creviceion a male's
territory was a measure of territory quality, and was related to the
number of females who lived there. Males defend larger home ranges
il.e ho..e. "'1-r 1
than are necessary for their own maintenance, and thusiappeart..to be
'superterritories' '(sensu Stamps 1983) in that they function primarily
to allow access to females.
In many lizards species opportunities for females to exhibit an active

preference among males is probably limited (Stamps 1983). However,
the pattern of home range overlap among Peninsula Dragons does
provide an opportunity for female choice. One female can overlap
with the home ranges. of several mature males, of_different ages,
colour patterns, sizes and territory holding capabilities (Chapter 6).
Thus the both intrasexual and intersexual s.election are theoretically ··
possible in Peninsula Dragons. However, obliteration of male colour
w~,·a.
pattern did not significantly reduce the number of females wit'X a
male paired. Furthermore, mate choice trials provided no evidence
that females actively chose between males (Chapter 8). Over four
breeding seasons the majority of pairings (96 %)were achieved by
dominant, territory holding males, as has been observed in most
other territorial lizards (Stamps 1983). This probably results from
three factors: (1) older males defend exclusive home ranges
260
Chapter 9 SYNTHESIS
(territories) from each other, (2) older males pair with (guard)
receptive females, and (3) young males are subordinated by older,
larger males (Chapter 6). I found no direct evidence that female mate
choice (active or passive) was acting on the traits measured in this
study. On the other hand, there was strong evidence for intrasexual
selection favouring male traits that allowed individuals to obtain and
defend territories.
Male Peninsula Dragons exhibited an age-related change in

reproductive tactics, similar to those observed in several other lizard
species (Hews & Moore 1995). Young, mature males appeared
unable defend their home ranges. They lived within the territories of
larger, black males by adopting subordinate behaviour and
maintaining brown dorsal colouration. These young males were
occasionally observed to accompany adult pairs during August and
September, when mating was occurring, but were rarely found paired
( 4 % of all pairs) with females themselves (Chapter 6).
The major difficulty in this project was that I did not observe any
matings. Future work should test the assumption that pairing
between males and females during spring indicates that they had
mated. The estimate of mating success by young males may be an
underestimate because they are harder to observe than older, more
conspicuous males. However, it may equally be that the low rate of
pairing observed in these males overestimates their mating success.
Young males were only observed to pair with females who had
already paired with an older, territory holding male who died while
261
Chapter 9 SYNTHESIS
guarding the female. In this situation, even if the young male had
mated with the female, he may not father any of her offspring
because the older male may have a first mate advantage (Smith 1984;
Eberhard 1985). A DNA analysis of paternity would allow one to
measure the true differences in reproductive success between older,
territory-holding males and younger males.
Young males reached a size which allowed them to challenge older

males for ownership of a territory, at the end of their first breeding
season. This appears to be a pivotal time in the life of a male
Peninsula Dragon. Males that failed to gain a territory by the
beginning of their second breeding season were never seen again, and
had presumably died or emigrated. Males that attained a territory
defended it for the rest of their lives, usually for one or two years
(95 percent of males). Territory defence across years has been
reported for other long-lived lizards (McCloskey et al. 1987;
Deslippe & McCloskey 1991).
The primary determinants of' whether a male was able to defend a

territory were his colour pattern, body size and relative head size.
Colour appeared to be important for recognition as a male by
other male Peninsula Dragons. Young, mature males may not
develop full colouration because it would be incur severe costs
associated with being harassed by territory holding males ( eg.
Cooper & Vitt 1987). Among older males, colour is an important
determinant of whether a male can maintain a territory ( Chapter
8), and therefore obtain priority mating access to females.
262
Chapter 9 SYNTHESIS'
It is not clear which particular components of male colour pattern

are important for sex recognition and territorial defence. Nor is it
clear to what extent signals in wavelengths of light visible to
humans, versus signals in the near ultraviolet (see Fleishman, Loew
& Leal 1993; Bennett, Cuthill, Partridge & Maier 1996) are
important. The eyes of both male and female Peninsula Dragons have
four types of cones (Johnston & Locket, unpublished data), similar
to the pattern found in anoline lizards (Fleishmann et al. 1993). This
suggests that they may be able to see in the near ultraviolet.
A good deal of attention has been paid to the role of gular

colouration as a signal in other lizard species (Rand & Williams
1970). It may play a role in species recognition (Williams & Rand
1977; Fitch & Hillis 1984; Losos 1985) and as a signal of social
status among males (Thompson & Moore 1991). Colouration on
other parts of a lizard's body may also signal social status (Olsson
1994a; Zucker 1994) or allow individual recognition (Olsson 1994c).
Many lizard species are able to modify their colouration naturally
(Cooper & Greenberg 1992), and the fitness of each colour variant
may easily be assessed in the context of male combat. Lizards may be
good model organisms for studies of multiple (Johnstone 1996) and
modifiable (Zucker 1994) social signals.
For colour patterns to play a role in individual recognition they

must be variable and complex (Olsson 1994c). The colour
patterns on the throat and lateral surfaces of the body in male
263
Chapter 9 SYNTHESIS
Peninsula Dragons show considerable variation and may be under

the influence of disruptive selection for individual recognition.
Components of the colour pattern that indicate status are usually
monochrome patches of colour, referred to as 'badges' (Dawkins
& Krebs 1978). Badges must be costly to produce if they are to
act as reliable indicators of status (Stamp-Dawkins & Guilford

1991). Carotenoids and melanins are two classes of pigments
which are known to be costly to produce, and which may produce
reliable signals of status (Lozano 1994). Colour, size, or
brightness of badges are known to be correlated with male
fighting ability in other lizard species (Inoue & Inoue 1977;
Thompson & Moore 1991; Olsson 1994a; Zucker 1994). The
black chest and dorsal colour of male Peninsula Dragons may act
as a signal of status. These aspects of male colouration were not
investigated in the present study. Future studies might attempt to
determine the function of each component of the colour pattern in
male Peninsula Dragons.
Body size and head size were also important determinants of whether -
a male could win a fight, maintain a territory, and thus had access to
females (Chapter 7). Large body size is co=only associated with an
increased likelihood of winning contests in other lizard species
(Tokarz 1985; Hews 1990; Olsson 1992). Large relative head size
has also been associated with the an increased likelihood of winning
contests (Hews 1990), and thus gaining access to mates (Ruby 1981;
Bull & Pamula 1996).
264
Chapter 9 SYNTHESIS
The selection analysis (Chapter 7) and field manipulation of colour

pattern (Chapter 8) demonstrated that body size, head size and
colour pattern were all subject to sexual selection in male Peninsula
Dragons. However, it was not clear from the results, whether these
traits were exhibiting a response to this selection (ie. were evolving).
The response to selection (R) of a trait may be predicted by the
following equation:
R = s.h 2
where s is the coefficient of selection acting on a trait, and h 2 is the

heritability of the trait (Falconer 1981). To determine whether a trait
is evolving one must either estimate heritability, or demonstrate a
shift in trait values between generations. I could not detect a change
in the values of any trait between generations. Small sample sizes
reduced statistical power, so this analysis was probably quite
unreliable. I was also unable to raise offspring from enough, females
to estimate the heritability of any of the traits identified as targets of
selection. Although patterns of inheritance for some-traits are
relatively well documented in some snakes (see Johnston 1996 'and
references therein), I am aware of only one attempt to estimate the
heritability of any of the traits I studied in lizards. Dorsal colour
pattern in the iguanid Uta stansburiana did not show any detectable
heritability (Gueniher, Ferguson, Snell & Snell 1993). The
heritability of the traits identified here as targets of selection should
be estimated. This is an important ne:i<t step .in determining whether
265
Chapter 9 SYNTHESIS
sexual selection has caused the evolution of sexually dimorphic traits

in Peninsula Dragons.
Estimating heritability of body size in a species which exhibits

indeterminate growth, would be particularly challenging. Similarly
the measurement of heritability of relative head size is not without
its difficulties. Recent studies on the evolution of allometric relations
of sexually selected horns in beetles (Emlen 1996) may provide a
model for such studies.
There were marked differences in the patterns of sexual selection

acting on the two cohorts studied (Chapter 7). In the younger cohort
there was significant selection on body size and head size. In the
'
older cohort no traits were identified as targets of selection. The
intensity of sexual selection varied annually in a population of
adders, Vipera berus (Madsen & 'Shine 1993). This variation in the
intf.lnsity of selection acting on reptiles may be a result of
environmental, physiological and demographic constraints (Halliday
1992). Being ectotherms, environmental constraints may have an
important effect of the direction and intensity of selection (Willmer
1991). Appart from the adder example (Madsen & Shine 1993),
published analyses of sexual selection in reptiles appear to have
ignored among year or among cohort variation in the selective regime
(see Andersson 1994). Changes in food availability may change the
costs and benefits of territoriality, and thus affect the mating system
in territorial lizards (Simon 1975) .. Determining the importance of
266
Chapter. 9 SYNTHESIS
such constraints and temporal changes may considerably alter our

view of the importance of different sources of selection.
This study only considered morphological traits. However, olfactory

and behavioural signals are also important for communication among
iguanian lizards (Alberts, Pratt & Phillips 1992; Fleishman 1992).
For example, the frequency of head bobs was correlated with
reproductive success in the iguanian, Sce/oporus jarrovi (Ruby
1981). Olfactory signals derived from the femoral gland secretions
play a role in determining dominance relationships in Iguana iguana
(Alberts et al. 1992). There are sexual differences in the chemical
components of the femoral pore secretions in Ctenophorus fionni (La
Nafie 1995). Determining (1) the extent to which behavioural and
olfactory traits are sexually dimorphic, (2) their interrelationship
with morphological traits, and (3) the evolutionary forces giving rise
to these differences would improve our understanding of lizard
behavioural ecology considerably.
267
REFERENCES CITED
AI-Johany, AM. (1995). The ecology ofAgama yemenensis Klausewjtz

(Lacertilia, Agamidae) in south-western Arabia. Journal ofArid Environments 29,
495-503.
Alatalo, R. V., Lundberg, A & Glynn, C. (1986). Female pied flycatchers choose
territory quality and not male characteristics. Nature 323, 152-153.
Alberts, AC. (1992). Density-dependent scent gland activity in desert:iguanas.

Animal Behaviour 44, 774-776.
Alberts, AC., Pratt, N.C. & Phillips, J.A (1992). Seasonal productivity of lizard
femoral glands: relationship to social dominance and androgen levels. l'hysiology
and Behaviour 51, 729-733.
Alexander, R.D., Hoogland, J.L., Howard, R., Noonan, KM. & Sherman, P. W.
(1979). Sexual dimotphisms and breeding systems in pinn.ipeds, ungulates and
humans. In Evolutionary Biology and Human Social Behaviour: An
Anthropological Perspective (Chagnon, N.A and Irons, W.D., eds.), Duxbury
Press, North Scituate, MA, pp. 402-435.
Allison, A (1982). Distribution and ecology ofNew Guinea Iiz..ards. In

Biogeography and Ecology ofNew Guinea (Gressitt, J.L., ed.), Dr W. Junk, The
Hague, Netherlands, pp. 803-813.
Altmann, J. (1974). Observational study ofbehaviour: sampling methods.

Behaviour 49, 227-267.
Andersson, M.B. (1982). Female choice selects for extreme tail length in a widow
bird. Nature 299, 818-820.
REFERENCES
Andersson, M.B. (1994). Sexual Selection. Princeton University Pres~: Princeton,

New Jersey
Andersson, M. & Norberg, R.A (1981) Evoluition of reversed sexual size

dimorphism and role partitioning among predatory birds, with a size scaling of
flight performance. Biological Journalof the Linnean Society 15, lOS-130.
Andren, C. (1986). Courtship, mating and agonistic behaviour in a free-living

population of adders, Vipera berus. Amphibia-Reptilia 7, 353-383.
Andrews, R.M. (1971). Structural habitat and time budget ofa tropicalAilo/is
I
lizard. Ecology 52, 262-270.
Andrews, R.M. (1982). Patterns of growth in Reptiles. In Biology of ~he reptilia,

Volume 13. Physiology D:Physiological Ecology (Gans, C. and Pough, F.R,
I
eds.), Academic Press: New York, New York, pp. 273-322.
Andrews, R.M. (1985). Mate choice by females ofthe,lizard, Anolis carolinensis.

Journal ofHerpetology 19, 284-289.
Andrews, RM. & Stamps, J.A (1994). Temporal variation in sexual size
dtmorphism ofAno/is lamifrons in Panama. Copeia 1994, 613-622.
Arak, A. ( 1988). Sexual dimorphism in body size: a model and a test. Evolution
42, 820-825.
Arnold, S.J. (1983). Sexual selection: the interfilce between theory and empiricism.
To.Mate Choice (Bateson, P., ed.), Cambridge University Press: Cambridge, United
Kingdom, pp. 67-107.
Arnold, S.J. & Duvall, D. (1994). Animal mating systems: a synthesis l>ased on
~lection theory. American Naturalist 143, 317-348.
Arnold, S.J. & Wade, M.J. (1984a). On the measurement ofnatural and sexual
selection: Theory. Evolution 38, 709-719.
269
REFERENCES
Arnold, S.J. & Wade, M.J. (1984b). On the measurement of natural and sexual
selection: Application. Evolution 38, 720-734.
Badham, J.A. (1971). Albumin formation in eggs of the agamid Amphibo/urus

barbatus. Copeia 1971, 543-545.
Baird, T.A, Acree, M.A. & Sloan, C.L. (1996). Age and gender-related
differences in the social behaviour and mating success offre&-living Collared
Lizards, Crotaphytus col/aris. Copeia 1996, 336-347.
Baker, R.R. & Parker, G.A. (1979). The evolution ofbird colouration.
Philosophical Transactions of the Raya/ Society B 287, 63-130.
Bakken, G.S. & Gates, D.M. (1975). Heat-transfer analysis of animals: some
implications for field ecology, physiology, and evolution. In Perspectives in
Biophysical Ecology (Gates, D.M. and Schmerl., R.B., eds.), Springer-Verlag,
New York, New York, pp. 255-290.
Bateman, A.J. (1948). Intra-sexual selection in Drosophila. Heredity 2, 349-368.
Bateson, P. (Ed.) (1983). Mate Choice. Cambridge ll"niversity Press: Cambridge,

United Kingdom,
Baverstock, P.R. (1972). The evolution and maintenance ofgenetic variability in

growth rates in the lizard, Ampln'bolurus omatus. Unpublished Ph.D. Thesis
Department of Zoology, University ofWestem Australia,
Baverstock, P.R. (1979). A three year study of the mammals and lizards ofBilliat
Conservation Park in tlie Murray mallee, South Australia. South Australian
Naturalist 53, 52-58.
Bellairs, AD. & Bryant, S. V. ( 1985). Autotomy and Regeneration in Reptiles. In

Biology of the Reptilia, Volume 15. Development 8. (Gans, C. and Billett, F.,
eds.), John Wiley& Sons, New York, NewYork,pp. 301-410.
270
REFERENCES
Benford, F., Lloyd, G.P. & Scharz, S. (1948). Coefficients of reflection of

magnesium oxide and magnesium carbonate. Journal of the Optical Society of
America 38, 445-447.
Bennett, A, Cuthill, L, Partridge, J.C. & Maier, E.J. (1996). mtraviolet vision and
mate choice in z.ebra finches. }fature 380, 433-435.
Bergland, A, Rosenqvist, G. & Svensson, I. (1986). Mate choice, fecundity and

sexual dimorphism in two pipefish species (Sygnathidae). Behavioural Ecology
and Sociobiology 19, 301-307.
Berk, M.L. & Heath, J.E. (1975). An analysis ofbehavioural thermoregulation in

the lizard Dipsosaurus dorsalis. Journal o/Thermal Biology 1, 15-22.
Berry, J.F. & Shine, R. (1980). Sexual size dimorphism and sexual selection in
turtles (Order Testudines). Oecologia 44, 185-191.
Berry, KR (1974). The ecology and social behaviour ofthe Cuckwalla,

Sa111'0malus obesus obesus Baird. University of California Publications in
Zoology 101, 1-60.
Bildstein, KL., McDowell, S.G. & Brisbin, I.L. (1989). Consequences of sexual
.dimorphism in sand fiddler crabs, Uca pugilator; differential vulnerability to avian
predation. Animal Behaviour 37, 133-139.
Blair, W.F. (1960). The Rusty Lizard: A Population Study. University of Texas
Press Austin, Texas,
Booth, C.L. (1990). Evolutionary significance of ontogenetic colour change in

animals. Biological Journal of the Linnean Society 40, 125-163.
Borgia, G. ( 1987). A critical review of sexual selection models. In Sexual

Selection: Testing the Alternatives (Bradbury, J.W. and Andersson, M.B., eds.),
Wiley, Chichester, U.K, pp. 55-66.
Bradbury, J.W. & Andersson, M.B. (Eds) (1987). Sexual Selection: Testing the
Alternatives. Wiley: Chichester, U.K
271
REFERENCES
Bradbury, J.W. & Davies, N.B. (1987). Relative roles of intra- and intersexual
selection. In Sexual Selection: Testing the Alternatives (Bradbury, J.W. and
Andersson, M.B., eds.), Wiley, Chichester, U.K, pp. 143-163.
Bradbury, J.W. & Vehrencamp, S.L. (1977). Social organisation and foraging in
emballonurid bats. m. Mating. systems. Behavioural Ecology and Sociobiology 2,
1-17.
Bradshaw, S.D. (1971). Growth and mortality in a field population of

Amphibolurus lizards exposed to seasonal cold and aridity.. Journal of Zoology
165, 1-25.
Bradshaw, S.D. (1981). Ecophysiology of Australian desert lizards: studies on the

genus Amphibolurus. In Ecological Biogeography efAustralia (Keast, A, ed.),
Dr W. Junk, The Hague, pp. 1394-1434.
Bradshaw, S.D. (1986). Ecophysiology efDesert Reptiles. Academic Press

Sydney, Australia,
Bradshaw, S.D. & Main, AR. (1968). Behavioural attitudes and regulation of
temperature inAmphibolurus lizards. Journal o/Zoology 154, 193-221.
Bradshaw, S.D. & Rice, G.E. (1981). Changes in dermal reflectance and
vascularity and their effects on thennoregulation in Amphibolums nuchalis
(Reptilia: Agamidae). Journal a/Comparative Physiology 135, 139-146.
Bradshaw, S.D., Saint Girons, R & Bradshaw, F.i (1991). Patterns ofbreeding in
two species of agamid lizards in the arid subtropical Pilbara region ofWestem
Australia. General and Comparative Endocrinology 82, 407-424.
Brodie, E.D., III, Moore, AJ. & Janzen, F.J. (1995). Visualizing and quantifying
natural selection. Trends in Ecology and Evolution 10, 313-318.
Brown, J.L. ( 1964). The evolution of diversity in avian territorial systems. Wilson
Bulletin 76, 160-169.
272
REFERENCES
Bull, C.M. (1988). Mate fidelity in an Australian lizard Trachydosaurus rugosus.

Behavioural Ecology and Sociobiology 23, 45-49.
Bull, C.M. ( 1994). Population dynamics and pair fidelity in sleepy lizards. In
Lizard Ecology: Historical and Experimental Perspectives (Vitt, L.J. and Pianka,
E.R., eds.), Princeton University Press, Princeton, New Jersey, pp. 57-174.
Bull, C.M., McNally, A & Dubas, G. (1991). Asynchronous seasonal activity of

male and female Sleepy Lizards, Tiliqua rugosa. Journal ofHerpetology 25, 436-
441.
Bull, C.M. & Pamula, Y. ( 1996). Sexually dimorphic head sizes and reproductive
success in the sleepy lizard, Tiliqua rugosa. Journal a/Zoology 240, 511-521.
Burke, T. (1989). DNA fingerprinting and other methods for the study of mating
success. Trends in Ecology and Evolution 4, 139-144.
Burley, N. (1981). Mate choice by multiple criteria in a monogamous species.

American Naturalist 117, 721-722.
Bustard, H.R. ( 1967). Activity cycle and thermoregulation in the Australian gecko,
Gehyra variegata. Copeia 1967, 753-758.
Butcher, G.S. & Rohwer, S. (1989). The evolution ofconspicoous and distinctive
coloration for comrn1mication in birds. In Current Ornithology 6 (Power, D.M.,
ed.), Plenum Press, New York, New York, pp. 51-108.
Cabanac, M., Hammel, T. & Hardy, J.D. (1967). Tifiqua scincoides: temperature--
sensitive units in lizard brain. Science 158, 1050-1051.
Campbell, B. (Ed.) ( 1972). Sexual Selection and the Descent ofMan. Aldine
Chicago, Illinois,
Carothers, J.H. (1984). Sexual selection and sexual dimorphism in some

herbivorous lizards. American Naturalist 124, 244-254.
273
REFERENCES
Carpenter, C.C. & Ferguson, G.W. (1977). Variation and evolution of stereotyped
behaviour in reptiles. In Biology of the Reptilia, Volume 7, Behavior and Ecology
(Gans, C. and Tinkle, D., eds.), Academic Press, New York, Ne\;V York, pp. 335-
554.
Caughley, G. (1977). Analysis ofvertebrate Populations. John Wiley & Sons

Brisbane, Australia,
Cheverud, J.M., Dow, M.N. & Leutenegger, W. (1985). The quantitative

assessment of phylogenetic constraints in comparative analyses: sexual dimorphism
in body weight among primates. Evolution 39, 1335-1351.
Christian, KA, Tracy, C.R. & Porter, W.P. (1983). Seasonal shifts in body
temperature and use ofmicrohabitats by the Galapagos Land Iguana. Ecology 64,
463-468.
Cleveland, W.S. (1979). Robust locally weighted regression and smoothing

scatterplots. Journal ofthe American Statistical Association 74, 829-836.
Cleveland, W.S. (1981). LOWESS: A program for smoothing scatterplots by

robust locally weighted regression. The American Statistician 35, 54
Clutton-Brock, T.H., Guiness, F.E. & Albon, S.D. (1982). Red Deer. Behaviour
and Ecology of Two Sexes. University of Chicago press Chicago, Illinois,
Clutton-Brock, T.H., Harvey, P.H. & Rudder, B. (1977). Sexual dimorphism,

socionomic sex ratio and body weight in primates. Nature 269, 797-800.
Cogger, H. G. ( 1974). Thermal relations of the malle dragon, Amphibolurus fordi

(Lacertila: Agamidae). Australian Jouma/ ofZoology 22, 319-339.
Cogger, HG. (1978). Reproductive cycles, fat body cycles and socio-sexual
behaviour in the mallee dragon, Amphibolurusfordi (Lacertilia: Agamidae).
Australian Journal ofZoology 26, 653-672.
274
REFERENCES
Cooper, W.E. (1984). Female secondary sexual coloration and sex recognition in
the keeled earless liz.ard, Holbrookia propinqua. Animal Behaviour 32, 1142-
1150.
Cooper,. W.E (1995). Foraging mode, prey chemical discrimination, and phylogeny
in liz.ards.. Anima/ Behaviour 50, 973-985.
Cooper, W.E. & Bums, N. (1987). Social sigoi.ficance ofventrolateral coloration

in the fence liz.ard, Sceloporus undulatus. Animal Behaviour 35, 526-532.
Cooper, W.E., Caffrey, C. & Vitt, L.J. (1985). diel activity patterns in the Banded
gecko, Coleonyxvariegatus. Journal ofHerpetology 19, 308-311.
Cooper, W.E. & Greenberg, N. ( 1992). Reptilian coloration and behavior. Tn

Biology ofthe Reptilia, Volume 18, Physiology E, Hormones, Brain, and
Behavior (Gans, C. and Crews, D., eds.), University of Chicago Press, Chicago,
Illinois, pp. 298-422.
Cooper, W.E. & Vrtt, L.J. (1987). Deferred agonistic behaviour in a long-lived
lizard Eumeces laticeps: field and lab<iratory data on the roles ofbody size and
residence in agonistic strategy. Oecologia 72, 321-326.
Cooper, W.E. & Vrtt, L.J. (1988). Orange head coloration of the male broad-
headed skink (Eumeces laticeps), a sexually selected social cue. Copeia 1988, 1-6.
Cooper, W.E. &. Vttt, L.J. (1989). Sexual dimorphismofhead and body size in an
iguanid liz.ard: paradoxical results. American Naturalist 133, 729-735.
Cooper, W.E. &. Vitt, L.J. (1993). Female mate choice oflarge male broad headed
skinks. Animal Behaviour 45, 683-693.
Crews, D. (1975). Effects of different components of male courtship behaviour on

environmentally induced ovarian recrudescence and mating preferences in the
lizard, Ano/is carolinensis. Animal Behaviour 23, 349-356.
Crisp, M., Cook, L.M. &. Hereward, F.V. (1979). Colour an heat balance in the
lizard Lacerta dugesii. Copeia 1979, 250.258.
275
REFERENCES
Cronin,, H ( 1991 ). The Ant and the Peacock. Cambridge University Press
Cambridge, United Kingdom,
Cuellar, 0. (1966). Oviduca1 anatomy and sperm storage structures in lizards.

Journal ofMorphalogy 119, 7-20.
Darwin, C. (1871). The Descent ofMan and Selection in Relation to Sex. John
Murray London,
Darwin, C. & Wallace, AR. (1858). On the tendency of species to form varieties;
and on the perpetuation of varieties and species by natural means of selection.
Journal of the Linnean Society ofLondon (Zoology) 3, 45-62.
Davidge, C. ( 1980). Reproduction in the herpetofimnal community of a Banksia

woodland near Perth, Western Australia. Australian Journal ofZoology 28, 435-
443.
Dawkins, R. & Krebs, J.R. (1978). Animal signals: information or manipulation. In

Behavioural Ecology: An Evolutionary Approach (Krebs, J.R. and Davies, N.B.,
eds.), Blackwell Scientific Publications, Oxford, U.K, pp. 282-309.
Davies, N.B. (1991). Mating Systems. In Behavioural Ecology: An Evolutionary

Approach (Krebs, J.R. and Davies, N.B., eds.), Blackwell Scientific Publications,
Oxford, U.K, pp. 263-300.
De Marco, V.G., Drenner, R. W. & Fergnson, G.W. (1985). Maximum prey size of
an insectivorous lizard, Sceloporus undulatus garmani.. Copeia 1985, 1077-1080.
Deslippe, R.J. & McCloskey, R. T. (1991). An experimental test of mate defence in

an ignanid lizard (Sce/oporus graciosus). Ecology 72, 1218-1224.
Diaz, J.A. (1993). Breeding coloration, mating opportunities, activity, and survival
in the lacertid liz.ardPsammodromus algirus. Canadian Journal of Zoology 71,
1104-1110.
276
REFERENCES
Dodd, C.K (1993). The effect oftoeclipping on sprint perfonnance of the liz.ard
Cnemidophorus sex/ineatus. Journal ofHerpetology 27, 209-213.
Downhower, J.F. (1976). Darwin's finches and the evolution of sexual dimorphism
in body size. Nature 263, 558-563.
Dugan, B. & Wiewandt, T.V. (1982). Socio-ecological determinants of mating

strategies in iguanine liz.ards. In Iguanas ofthe World: their behavior, ecology and
conservation (Burghardt, G.M & Rand, AS., eds.), Noyes Publications: New
Jersey, pp. 303-319.
Dumbacher, J.P., Beehler, B.M, Spande, T.F., Garraffo, HM. & Daly, J.W.
(1992). Homobatrachotoxin in the genusPitohui: Chemical defense in birds?.
Science 258, 799-801.
Dunham, A.E. (1980). An experimental study ofinterspecific competition between

the iguanid lizards Sceloporus me"iami and Urosaurus omatus. Ecological
Monographs 50, 309-330.
Dunham, A.E. (1981). Populations in a fluctuating environment: the comparative

population ecology of Sceloporus me"iami and Urosaurus ornatus.
Miscellaneous Publications ofthe University ofMichigan Museum of Zoology
158, 1-62.
Dyson, M.L. (1989). Aspects ofsocial behaviour in a caged population ofpainted

reedfrogs. Unpublished PhD Thesis University ofWitwatersrand, Johanesburg,
Dyson, M.L. &Passmore, N.L (1988a). The combined effect of intensity and the
temporal relationship of stimuli on the phonotactic responses of female painted
reed frogs (Hyperolius marmoratus). Animal Behaviour 36, 1555-1556.
Dyson, M.L. & Passmore, N.l (1988a). two-choice phonotaxis in Hyperolius

marmoratus (Anura: Hyperolidae): the effect of temporal variation on presented
stimuli Animal Behaviour 36, 648-652.
Eberhard, W.G. (1985). Sexual Selection and Animal Genitalia. Harvard

University Press Cambridge, Massuchusets,
277
REFERENCES
Ellington, J.M. (1993). Sexual selection, coloration, and sensory systems in a

diurnal gecko. Abstract of the joint meeting of the American Society of
Ichthyologists and Herpetologists, Larval Fish Co,iferense, the Herpetologist's
league and the American Elasmobranch Society
Emlen, D.J. (1996). Artificial selection on horn length-body size allometry in the
homed beetle Onthophagus acuminatus (Coleoptera: Scarabaeidae). Evolution
50, 1219-1230.
Emlen, S.T. & Oring, L.W. (1977). Ecology, sexual selection, and the evolution of
mating systems. Science 197, 215-223.
Endler, J.A (1986). Natural Selection in the Wild. Princeton University Press
Princeton, New Jersey,
Enquist, M., Leitnar, 0., Ljungberg, T., Mallner, Y. & Segerdahl, N. (1990}. A
test of the sequential assessment game: fighting in the cichlid fish Nannacara
anomala. Animal Behaviour 40, 1-14.
Estes, R., de Quiroz, K & Gauthier, J. (1988). Phylogenetic relationships within

the Squamata. Jn Phylogenetic Relationships of the Lizard Families, Essays
commemorating Charles. L. Camp (Estes, R. and Pregill, G., eds.), Stanford
University Press, Stanford, pp. 119-281.
Etheridge, R. (1967). Lizard caudal vertebrae. Copeia 1967, 699-721.
Falconer, D.S. (1981). Introduction to Quantitative Genetics. Longman New

York, New York,
Fenner, J.W. (1979). A review ofMarking Techniques for Amphibians and

reptiles. Society for the Study of Amplullians and Reptiles Oxford, Ohio,
Fink, W.L. (1982). The conceptual relationship between ontogeny and phylogeny.
Paleobiology 8, 254-264.
278
REFERENCES
Firth, B. T. & Turner, J.S. (1982}. Sensory, neural, and honnonal aspects of
thermoregulation. In Biology of the Reptilia, Volume 12: Physiology C:
Physiological ecology (Gans, C. and Pough, F.R, eds.), Academic Press, New
York, New York, pp. 213-274.
Fisher, R.A (1930). The genetical Theory ofNatural Selection. Clarendon Press
Oxfurd, United Kingdom,
Fisher, R.A (1958). The Genetical Theory ofNatural Selection. Dover New
York, New York,
Fitch, RS. (1981). Sexual size differences in reptiles. Miscellaneous Publications

ofthe Museum ofNatural History, University ofKansas 70, 1-72.
Fitch, RS. & Hillis, D.M. (1984). Te Ano/is dewlap: intraspecific variability and
morphological associations with habitat. Copeia 1984, 315-323.
Fleishman, L.J., Loew, E.R. & Leal, M. (1993). Ultraviolet vision in lizards.
Nature 365, 397
Fleming, T.R & Hooker, R.S. (1975). Ano/is cupreus: the response ofa lmird to
tropical seasonality. Ecology 56, 1243-1261.
Forsman, A (1991 ). Variation in sexual size dimorphism and maximum body size
among adder populations: effects of prey size. Journal ofAnimal Ecology 60,
253-267.
Frost, D.R. & Etheridge, R. (1989). A phylogenetic analysis and taxonomy of

iguanian lizards (Reptilia: Squamata). University ofKansas Museum ofNatural
History Miscellaneous Publications 81, 1-65.
Garland, T. (1985). Ontogenetic and individual variation in size, shape, and speed
in the Australian agarnid lizard Amphibolurus nuchalis. Journal ofZoology 207,
425-439.
Garland, T., Hankins, E. & Huey, R.B. (1990). Locomotor capacity and social
dominance in male lizards. Functional Ecology 4, 243-250.
279
REFERENCES
Ghiselin, M. T. ( 1974). The Economy ofNature and the Evolution ofSex.

University ofCalifomia Press berkeley, California,
Gibbons, J.R.H. (1979). The hind leg pushup display of the Amphibolurus decresii
species complex (Lacertilia: Agamidae). Copeia 1979, 29-40.
Gibbons, J.R.H. & Lillywhite, H.B. ( 1981). Ecological segregation, colour

matching, and speci.ation in Ji7.ards of the Amphibolurus decresii species complex
(Lacertilia: Agamidae). Ecology 62, 1573-1584.
Gibson, R.M. (1996). female choice in sage grouse: the roles of attraction and
active comparison. Behavioural Ecology and Sociobiology 39, 55-59.
Gibson, R.M. & Bradbury, J.W. (1985). Sexual selection in lekking sage grouse:
phenotypic correlates of male mating success. Behavioural Ecology and
Sociobiology 18, 117-123.
Gotmark, F. (1992). Anti-predator effect of conspicuous plumage in a male bird.
Grahn, M. (1993). Mortality in the pheasant Phasianus colchicus during the

breeding season. Behavioural Ecology and Sociobiology 32, 95-10 l.
Greenberg, B. & Noble, G.K (1944). Social behaviour of the American chameleon
(Ano/is caralinensis Voigt). Physiological Zoology 17, 392-439.
Greer, AE. (1989). The Biology and Evolution ofAustralian Lizards. Surrey
Beatty Norton, New South Wales,
Griffiths, AD. (1994). The effect ofa seasonal environment andfire on the
ecology offrillneck lizards, Chlamydosaurus kingii, in the wet-dry tropics of
northern Australia. Faculty of Science, Northern Territory University Unpublished
MSc Thesis,
Guenther, M.M., Ferguson, G.W., Snell, H.L. & Snell, H. (1993). The variation
and genetic basis of dorsal color pattern in the desert side-blotched Ji7.ard, Uta
stansburiana stejnegeri. Journal ofHerpetology 27, 199-205.
280
REFERENCES
Gwynne, D.T. (1984). Courtship feeding increases female reproductive success in

bushcrickets. Nature 307, 361-363.
Hall, W.P. (1970). Three probable cases of parthenogenesis in lizards (Agamidae,

Chamaeleontidae, Gekkonidae). F.xperientia 26, 1271-1273.
Halliday, T.R (1992). Sexual selection in amphibians and reptiles: theoretical

issues and new directions. Jn Herpetology: Current Research on the Biology of
Amphibians and Reptiles (Adler, K, ed.), Society for the Study of Amphibians and
Reptiles, Oxford, Ohio, pp. 81-95.
Halliday, T.R & Verrell, P.A (1988). Body size and age in Ampht"bians and
Reptiles. Journal ofHerpetology 22, 253-265.
Hamilton, W.D. & Zuk, M (1982). Heritable true fitness and bright birds: a role
for parasites? Science 218, 384-387.
Harris, V.A (1964). The life of the Rainbow Lizard. Hutchinson London,
Hasson, 0. (1989). Amplifiers and the handicap principle of sexual selection: a

different emphasis. Proceedings of the Royal Society ofLondon B 235, 383-406.
Hastings, P.A (1988). Female choice and male reproductive success in the angel
blenny, Cora//iozetus angelica (Teleostei: Chaenopsidae). Animal Behaviour 36,
115-124.
Hay, M. (1972). The breeding ofPhysignathus leseurii L. in captivity.

Herpetofauna 5, 2-3.
Head, G. (1995). Selection on fecundity and variation in the degree of sexual size
dimruphism among spider species (Class Araneae). Evolution 49, 776-781.
Heatwole, H. (1970). Thermal ecology of the desert dragonAmphibolurus

inermis. Ecological Monographs 40, 425-457.
281
REFERENCES
Hedrick, A.V. (1986). Female choice and the heritability of attractive male traits:
An empirical study. American Naturalist 132, 267-276.
Hedrick, AV. & Temeles, E.J. (1989). The evolution of sexual dimorphism in
animals: hypotheses and tests. Trends in Ecology and Evolution 4, 136-138.
Heideman, N.J.L. (1993). Does crypsis reduce predation pressure in agruna

lizards? Evidence from tail damage. Amphibia-Reptilia 195-197.
Hertz, P.E. (1992). Evalnating thermal resource partitioning by sympatric lizards

Ano/is cooki and Ano/is cristatellus: a field test using null hypotheses. Oecologia
90, 127-136.
Hertz, P.E., Huey, R.B. & Stevenson, R.D. (1993). Evaluating temperature
regulation by field-active ectotherms: the fallacy of the inappropriate question.
Hews, D.K. (1990). Examining hypotheses generated by field mea~s of sexual

selection on male liz.ards, Uta palmeri. Evolution 44, 1936-1966.
Hews, D.K. (1993). Food resources affect female distnlmtion and male mating
opportunities in the iguanian liard Uta palmeri. Animal Behaviour 46, 279-291.
Hews, D.K. (1996). Size and scaling of sexually-selected traits in the lizard, Uta
palmeri. Journal of Zoology 238, 743-757.
Hews, D.K. & Moore, M.C. (1995). Honnonal basis of alternative reproductive
phenotypes in male lizards. XXIV International Ethological Co,iference Abstracts
50(Abstract)
Hixon, M.A (1987). Territory area as a determinant pf mating system American

Zoologist 27, 229-247.
Holling, C.S. (1959). Some characteristics of simple types of predation and

parasitism Canadian Entomologist 91, 385-398.
282
REFERENCES
Houde, AE. (1987). Mate choice based upon naturally occurring color-pattern
variation in a guppy population. Evolution 41, 1-10.
Houston, D. & Shine, R. (1993). Sexual dimorphism and niche divergence: feeding
habits of the Arafura filesnake. Journal ofAnimal Ecology 6Z, 737-748.
Houston, T.F. (1974). Revision of the Amphibolurus decresii complex (Lacertilia:

Agamidae) of South Australia. Transactions of the Royal Society ofSouth
Australia 98, 49-60.
Houston, T.F. (1978). Dragon Lizards and Gommas ofSouth Australia. South
Australian Museum Adelaide, South Australia,
Howard, R.D. (1981). Sexual dimoxphism in Bullfrogs. Ecology 6Z, 303-310.
Hubbell, S.P. & Johnson, L.K (1987). Environmental variance in lifetime mating
success, mate choice and sexual selection. American Naturalist 130, 91-112.
Huck, W. & Banks, E.M. (1982). Male dominance status, female choice and
mating success in the brown lemming, Lemmus trimucronatus. Animal Behaviour
30, 665-675.
Hudson, P.J. ( 1977). An account of egg laying by the Thorny Devil, Moloch
horridus (Gray). Herpetofauna 9, 23-24.
Huey, R.B., Dunham, AE., Overall, KA & Newman, R.A (1990). Variation in
locomotor performance in demographically known population ofthe lizard
Sceloporus merriami. Physiological Zoology 63, 845-872.
Huey, R.B., Pianka, E.R. & Schoener, T.W. (Eds) (1983). Lizard Ecology: Studies
of a Model Organism. Cambridge, Massachusetts Harvard University Press,
Huey, R.B. & Slatkin, M. (1976). Costs and benefits oflizard thermoregulation.
Quarterly Reviews in Biology 51, 363-384.
Huxley, J.S. (J.932). Problems ofRelative Growth. Dial New York, New York,
283
REFERENCES
Imms, R.A ( 1988). Spatial clumping of sexually receptive females induces space
sharing among male voles. Nature, 335, 541-543.
Inger, R.F. & Greenberg, B. (1966). Annual reproductive patterns of lizards from a
Bomean rain forest. Ecology 47, 1007-1021.
Inoue, S. & Inoue, Z. (1977). Colour changes induced by pairing and painting in
the male rainbow lizard, Agama agama agama. Experientia 33, 1443-1444.
James, C.D. & Shine, R. (1985). The seasonal timing of reproduction; a tropical-
temperate comparison in Australian lizards. Oecologia 67, 464-474.
Jennrich, R.I. & Turner, F.B. (1969). Measurement of non-circular home ranges.
Journal of Theoretical Biology 22, 227-237.
Jenssen, T.A ( 1970). The ethoecology of Ano/is nebulosus (Sauria, Jguanidae).

Jownal ofHerpetology 4, 1-38.
Jenssen, T.A, Greenberg, N. & Hovde, KA (1995). Behavioural profile offree-

ranging male lizards, Ano/is carolinensis, across breeding and post breeding
seasons. Herpetological Monographs 9, 41-62.
Jolmston, G.R. (1979). The eggs, incubation and ymmg of the bearded dragon
Amphibolurus vitticeps Ahl Herpetofauna 11, 5-8.
Jolmston, G.R. (1982). The herpetofauna of the Middleback Range area, South
Australia 1. An annotated checklist. Herpetofauna 14, 52-60.
Johnston, G.R. (1992). Ctenophorus tjantja!Jra, a new dragon lizard (Lacertilia:

Agamidae) from Northern South Australia. Records of the South Australian
Museum 26, 51-59.
Jolmston, G.R. (1996). Genetic and seasonal variation in body colour of the
Australian death adder, Acanthophis antarcticus (Squamata: Elapidae). Journal
ofZoology 239, 187-196.
284
REFERENCES
Johnstone, R.A (1996). Multiple displays in animal communication - backup

signals and multiple messages. Philosophical Transactions of the Royal Society B
351, 329-338.
Jones, S'.M. & Ferguson, G.W. (1980). The effect of paint marking on mortality in
a Texas population of Sceloporus undulatus. Copeia 1980, 850-854.
Kanfinan, KW. (1981). Frtting and using growth cUIVes. Oecologia 49, 293-299.
Kimura, M. (1983). The Neutral Theory ofMolecular Evolution. Cambridge

University Press Cambridge, United Kingdom
Kingsolver, J.G. (1985). Thermoregnlatory significance of wing melan.ization in

Pieris butterflies: physics, posture and pattern. Oecologia 66, 546-551.
Kirkpatrick, M. & Ryan, M.J. (1991). The evolution of mating preferences and the
paradox of the lek. Nature 350, 33-38.
Kitzler, G.Z. (1941). Die Paarungsbiologie einiger Eidechsen. Zietschrift

Tierpsychologie 4, 335-402.
Kluge, AG. (1981). The life history, social organization, and parental behaviour of
Hy/a rosenbergl Boulenger, a nest-building gladiator frog. Miscellaneous
publications qfthe Museum o/Zoology, University ofMichogan 160, 1-170.
Kramer, G. (1937). Beobachtungen uber Paaraungsbiologie und sociales Verhalten

vonMauereidechsen. Zietschrift Morphologie 0/rol Tiere. 32,.752-783.
Krekorian, C.O. (1984). Life history of the desert iguana, Dipsosaurus dorsa/is.
Herpetologica 40, 415-424.
Krekorian, C.O. (1989). Field and laboratory observations on chemoreception in

the desert iguana, Dipsosaurus dorsalis. Journal ofHerpetology 23, 267-273.
La Nafie, N. (1995). Chemical Properties ofFemoral Gland Secretions in Dragon

Lizards. School of Chemistry, Flinders University Unpublished M.Sc. Thesis
285
REFERENCES
La Nafie, N., Prager, R.R & Johnston, G.R. (1995). Chemical characterisation of
races ofthe Penrosnla Dragon liurd. Australian Coordinating Group for
Chemistry Chemical Research Communications 3, 40-44.
Lande, R. (1980). Sexual dimorphism, sexual selection, and adaptation in

polygenic characters. Evolution 34, 292-307.
Lande, R. (1981). Models of speciation by sexual selection on polygenic traits.

Proceedings of the National Academy of Sciences of the United States ofAmerica
78, 3721-3725.
Lande, R. & Arnold, S.J. (1983). The measurement of selection on correlated

characters. Evolution 37, 1210-1226.
Lennar, 0., Austad, S. & Enquist, M. (1991). A test of the sequential assessment
game: fighting in the bowl and doily spider Frontinella pyramitela. Evolution 45,
862-874.
Licht, P. (1967). Interaction ofprolactin and gonadotropins on appetite, growth,

and tail regeneration in the liurd, Ano/ls carolinensis. General and Comparative
Endocrinology 9, 49-63.
Lin, J. & Lu, K (1982). Population ecology ofthe liurd Japalura swinlwnis
formosensis (Samia: Agamidae) in Taiwan. Copeia 1982, 425-434.
Livezey, B.C. & Humphrey, P.S. (1985). Territoriality and interspecific aggression
in steamer-ducks. Condor 87, 154-157.
Livezey, B.C. &Humphrey, P.S. (1986). Flightlessaess in Steamer-Ducks

(Auatidae: Trachyeres): its morphological bases and probable evolution. Evolution
40, 540-558.
Loffredo, C.A. & Borgia, G. (1986). Sexual selection, mating systems, and the
evolution of avian acoustical displays. American Naturalist 128, 773-794.
Losos, J.B. (1985). Au experimental demonstration of the species-recognition role

ofAnolis dewlap color. Copeia 1985, 905-910.
286
REFERENCES
Losos, J.B. ( 1988). Thermoregulatory correlates of escape behaviour by a desert

lizard, Ctenophorus isolepis. Journal ofHerpetology 22, 353-356.
Lowe, P.R (1928). A description ofAtlantisia rogerst, the diminutive and

flightless rail oflnaccesst'ble Island (southem Atlantic), with some notes on
flightless rails. Ibis 70, 99-131.
Louno, G.A (1994). Carotenoids, parasites, and sexual selection. Oikos 70, 309-
311.
Ludbrook, N.H. (1980). A Guide to the Geology and Mineral Resources ofSouth
Australia. Government Printer Adelaide, South Australia,
Madsen, T. (1988). Reproductive success, mortality and sexual size dimorphism in

the adder, Vipera berus. Ho/arctic Ecology 11, 77-80.
Madsen, T. &Loman, J. (1987). On the role of colour display in the social and
spatial organization of male rainbow lizards (Agama agama). Amphibia-Reptilia
8, 365-372.
Madsen, T. & Shine, R (1992). A rapidly, sexually selected shift in mean body size
in a population of snakes. Evolution 46, 1220-1224.
Madsen, T. & Shine, R. (1993). Temporal variability in sexual selection on

reproductive tactics and body size in male snakes. American Naturalist 141, 167-
171.
Madsen, T., Shine, R., Loman, J. & Hakansson, T. (1992), Why do female adders
copulate so frequently? Nature 355, 440-441.
Magnuson, J.J., Crowder, L.B. & Medvick, P.A (1979). Temperature as an

ecological resource. American Zoologist 19, 331-343.
Manning, A (1991). Notes on nesting, incubation and hatching of the Southern

Angle-headed Dragon, Hypsi/urus spinipes (Squamata: Agaminae). Herpetofauna
21, 15-19.
287
REFERENCES
Manzur, M.I. & Fuentes, E.R. (1979). Polygyny and agonistic behaviour in the
tree-dwelling lizard Liolaemus tenuis (Iguanidae ). Behavioural Ecology and
Marler, C.A & Moore, M. C. ( 1988). Evolutionazy costs of aggression revealed by

testosterone manipulations in free-living male lizards. Behavioural Ecology and
Martins, E.P. (1994). Phylogenetic perspectives on the evolution of lizard

territoriality. In Lizard ecology: Historical and Experimental Perspectives (Vitt,
L.J. and Pianka, E.R., eds.), Princeton University Press, Princeton, New Jersey,
Mawson, P.M. ( 1972). The nematode genus Maxvachonia (Oxyurata:

Cosmocercidae) in Australian reptiles and frogs. Transactions of the Royal
Society ofSouth Australia 96, 101-108.
Maynard Smith, J. (1987). Sexual selection- a classification of models. In Sexual

Selection: testing the Alternatives (Bradbury, J.W. and Andersson, M.B., eds.),
Wiley, Chichester, United Kingdom, pp. 9-20.
Mayr, E. (1963). Animal Species and Evolution. Belknap press Cambridge,

Massachusetts,
McCloskey, R. T., Baia, KA & Russell, R W. (1987). Defense of mates: a

territory departure rule for male tree lizards following sex-ratio manipulation.
Oecologia 73, 28-31.
Mccloskey, R. T., Deslippe, R.J., Szpak, C.P. & Baia, KA (1990). Ecological
correlates of the variable mating system of an iguanid lizard. Oikos 59, 63-69.
McCloskey, RT., Szpak, C.P. & Deslippe, R.J. (1990). Experimental assessment
of factors affecting the distnlmtion of adult female tree lizards. Oikos 59, 183-18 8.
McComb, K & Clutton-Brock, T.R (1994). Is mate choice copying or

aggregation responsi'ble for skewed distnlmtions of females at leks?. Proceedings
ofthe Royal Society ofLondon B 255, 13-19.
288
REFERENCES
McGillivray, W.B. & Johnston, R.F. (1987). Differences in sexual size dimorphism
and body proportions between aduh and subaduh sparrows in North America. Auk
104, 681-687.
McNamara, KJ. (1995). Sexual dimorphism: the role ofheterochrony. In

Evolutionary Change and Heterochrony (McNamara, KJ., ed.), John Wiley &
Sons, Chichester, United Kingdom, pp. 65-90.
Mitchell, F.J. (1973). Studies on the ecology of the agamid lizardAmphibolurus

maculosus. Transactions ofthe Royal Society ofSouth Australia 97, 47-76.
Mitchell-Olds, T. & Shaw, R.G. (1987). Regression analysisofnatural selection:

statistical inference and biological interpretation. Evolution 41, 1149-1161.
Moller, AP. & Pomiankowski, A (1993). Why have birds got muhiple sexual
omaments? Behavioural Ecology and Sociobiology 3:2, 167-176. ·
Moody, S.M. (1980). Phylogenetic and historical biogeographical relationships

ofthe genera in the family Agamiclae (Reptilia: Lacertilia). Unpublished PhD
Thesis Department of Zoology, University ofMichigan,
Moore, AJ. (1988) Female preferences, male social status, and sexual selection in
Nauphoeta cinerea. Animal Behaviour 36, 303-305.
Moore, AJ. (1990). The evolution of sexual dimorphism by sexual selection: the
separate effects ofintrasexual and intersexual selection. Evolution 44, 315-331.
Moore, AJ. & Breed, M.D. (1986). Mate assessment in a cockroach, Nauphoeta
cinerea. Animal Behaviour 34, 1160-1165.
Moore, M.C. (1987). Castration affects territorial and sexual behaviour offree-
living male lizards, Sceloporus jarrovi. Animal Behaviour 35, 1193-1199.
289
Rl'!FERENCES
Moore, M.C. (1988). Testosterone control of territorial behaviour: tonic-release

implants fully restore seasonal and short-term aggressive responses in free-living
castrated lu.ards. General and Comparative Endocrinology 70, 450-459.
Mori, A. & Hikida, T. (1994). Field observations on the social behaviour of the
Flying Lu.ard, Draco volans sumatranus, in Borneo. Copeia 1994, 124-130.
Niewiarowski, P.H ( 1995). Effects of supplemental feeding and thermal

environment on growth rates of eastern fence lu.ards, Sceloporus undulatus.
Nisbet, I.C. T. (1973). Courtship-feeding, egg-size and breeding success in

common terns. Nature 241, 141-142.
Noble, G.K (1934). Experimenting with the courtship oflu.ards. Natural History
34, 5-15.
Noble, G.K & Bradley, HT. ( 1933). The mating behaviour oflu.ards; its bearing
on the theory of sexual selection. Annals of the New York Academy ofScience 33,
25-100.
Norris, KS. (1967). Color adaptation in desert reptiles and its thermal
relationships. lnLizardEcology: A Symposium (Milstead, W.W.,.ed.), University
of Missouri Press, Columbia, Missouri, pp. 169-229.
Olsson, M. (1992a). Contest success in relation to size and residency in male sand
lu.ards Lacerta agilis. Animal Behaviour 44, 386-388.
Olsson, M. (1993a). Nuptial coloration and predation risk in model sand lu.ards,
Lacerta agilis. Animal Behaviour 46, 410-412.
Olsson, M. ( 1993b). Contest success and mate guarding in male sand lu.ards,
·Lacerta agilis. Animal Behaviour 46, 408-409.
290
REFERENCES
Olsson, M. (1993c). Male preference for large females and assortative mating for
body size in the sand lizard (Lacerta agilis). Behavioural Ecology and
Olsson, M. (1994a). Nuptial colouration in the sand lizard, Lacerta agilis: an intra-
sexually selected cue to fighting ability. Animal Behaviour 48, 607-613.
Olsson, M. (1994b). Why are sand lizard males (Lacerta agilis) not equally green?.
Behavioural &ology and Sociobiology 35, 169-173.
Olsson, M. (1994c). Rival recognition affects male contest behaviour in sand

lizards (Lacerta agilis). Behavioural Ecology and Sociobiology 35, 249-252.
Olsson, M (1995). Forced copulation and costly female resistance behaviour in the
Lake Eyre Dragon, Ctenophorus macu/osus. Herpetologica 51, 19-24.
Olsson, M., Gullberg, A, Tegelstrom, R, Madsen, T. & Shine, R. (1994). Can

female adders multiply?. Nature 369, 528
Olsson, M. & Madsen, T. (1995). Female choice and male quantitative traits in
lizards- Why is it so rare?. Behavioural Ecology and Sociobiology 36, 179-184.
r
Owen-Smith, N. (1993). Comparative mortality rates of male and female kudus:
the costs of sexual dimorphism. Journal of'Animal &o/ogy 62, 428-440.
Parker, AJ., Fanning, C.M & Flint, R.B. (1985). Geology. In Natural History of
Eyre Peninsula (Twidale, C.R., Tyler, M.J. and Davies, M., eds.), Royal Society of
South Australia, Adelaide, South Australia, pp. 21-45.
Parker, G.A (1983). Mate quality and mating decisions. In Mate Choice (Bateson,
P., ed.), Cambridge University Press, Cambrldge,U.K, pp. 141-166.
Parker, W.S. & Pianka, E.R. (1973). Notes on the ecology of the iguanid lizard
Sceloporus magister. Herpetologica 29, 143-152.
291
REFERENCES
Patterson, J.W. & Davies, P.M.C. (1978). Preferred body temperature: seasonal
and sexual differences in the lizard Lacerta vivipara. Jounwl of Themzal Biology
3, 39-41.
Patton, J.L. & Brylski, P.V. (1987). Pocket gophers in alfalfa fields: causes and
consequences of habitat-related body size variation. American Naturalist 130,
493-506.
Payne, R.B. (1979). Sexual selection and intersexual differences in variance of

breeding success. American Naturalist 114, 447-452.
Payne, R.B. (1984). Sexual selection, lek and arena behaviour, and sexual size
dimorphism in birds. Ornithological Monographs 33, 1-53.
Pengilley, R. (1982). Note on the reproductive biology ofthe ring-tailed dragon

(Chelo.sania brunnea). Northern Territory Naturalist 5, 6
Perezi-Mellado, V. &. De La Riva, I. (1993). Sexual size dimorphism arid ecology:

the case of a tropical lizard, Tropidurus me/anopleurus (Sauria: Tropiduridae).
Copeia 1993, 969-976.
Petrie, M. (1983a). Female moorhens compete for small fat males. Science 220,
413-415.
Petrie, M. (1983b). Mate choice in role-reversed species. In Mate Choice

(Bateson, P., ed.), Cambridge University Press, Cambridge, United Kingdom, pp.
167-180.
Petrie, M., Halliday, T.R. &. Sanders, C. (1991). Peahens prefer peacocks with
elaborate trains. Animal Behaviour 41, 323-331.
Petrov, 0. V. (1996). Sexual dimorphism in the skull ofMustela emzinea L..

VestnikLeningrad Univ. 15, 41-56.
Pianka, E.R. (1967). Lizard species diversity. Ecology 48, 338-351.
292
REFERENCES
Pianka, E.R. (1970). Comparative autecology of the li7.ard Cnemidophorus tigris

in different parts of its geographic range. Ecology 51, 703-720.
Pianka, E.R. ( 1971 ). Ecology of the agamid li7.ard Amphibolurus isolepis in

Western Australia. Copeia 1971, 527-536.
Pianka, E.R. (1973). The structure oflizard communities.. Annual Revi<!Ws of

Ecology and Systematics 4, 53-74.
Pianka, E.R. (1986). Ecology and natural history ofdesert lizards: analysis of the
ecological niche and community structure. Princeton University Press Princeton,
New Jersey,
Pianka, E.R. & Pianka, RD. (1970). The ecology ofMoloch horridus (Lacertilia:
Agamidae) in Western Australia. Copeia 1970, 90-103.
Pietruszki, R.D. (1986). Search tactics of desert li7.ards: how polarised are they?.
Platt, J.R. (1964). Strong inference. Science 146, 347-353.
Porter, W.P. & James, F.C. (1979). Behavioral implications ofmechanistic ecology
II. the Aftican Rainbow Liard, Agama agama. Copeia 1979, 594-619.
Porter, W.P., Mitchell,_ J.W., Beckman, W.A & De Wrtt, C.B. (1973). Behavioural
implications of mechanistic ecology. Thermal and behavioural modelling of desert
ectothenns and their microenvironment. Oecologia 13, 1-54.
Porter, W.P. & Norris, KS. (1969). Lizard reflectivity change and its effect on
light transmission through the body wall. Science 163, 482-484.
Pounds, J.A & Jackson, J.F. (1983). Utilization of perch sites by sex and size
classes of Sceloporus undulatus undulatus. Journal ofHerpetology 11, 287-289.
Prestt, I. (1971). An ecological study of the viper, Vipera berus, in southern

Britain. Journal of Zoology 164, 373-418.
293
REFERENCES
Price, T.D. (1984). Sexual selection on body size, territocy and plumage variables
in a population of Darwin's finches. Evolution 38, 327-341.
Procter, J.B. (1923). On new and rare reptiles and batracbians from the Australian
region. Proceedings of the Zoological Society ofLondon 1923, 1069-1077.
Queller, D.C., Strassmann, J.E. & Hughes, C.R. (1993). Microsatellites and
kinship. Trends in Ecology and Evolution 8, 285-288.
Quinn, J.F. & Dunham, AE. (1983). On hypothesis testing in ecology and
evolution. American Naturalist 122, 602-617.
Ralls, K (1977). Sexual dimorphism in mammals' Avian models and unanswered

questions. American Naturalist 111, 917-937.
Ralls, K & Harvey, P.H (1985). Geographic variation in size and sexual
dimorphism of North American weasels. Biological Journal of the Linnean
Society 25, 119-167.
Rand, AS. ( 1954). Variation and predator pressure in an island and a mainland
population ofliz.ards. Copeia 1954, 260-262.
Rand, AS. (1967). Ecology and social organisation of the iguanid liz.ardAno/is
/ineatopus. Proceedings of the United States National Museum 122, 1-79.
Rand, AS. & Williams, E.E. (1970). An estimation of redundancy and information
content of anole dewlaps. American Naturalist 104, 99-103.
Regal, P.J. (1983). Adaptive zone and behaviour of lizards.. In Lizard Ecology:
studies ofa model organism. (Huey, R.B .., Pianka, E.R. and Schoener, T.W., eds.),
Harvard University Press, Cambridge, Massachusetts, pp. 105-118.
Reyment, R.A, Blackith, R.E. & Campbell, N.A (1984). Multivariate

Morphometrics. Academic Press London, United Kingdom,
Reichert, S.E. (1978). Games spiders play: Behavioral variability in territorial

disputes. Behavioural Ecology and Sociobiology 3, 135-162.
294
REFERENCES
Reichert, S,E. (1984). Games spiders play. ill: Cues underlying context-associated
changes in agonistic behaviour. Animal Behaviour 32, 1-15.
Robinson, S.K (1986). Benefits, costs, and.determinants of dominance in a

polygynous oriole. Animal Behaviour 34, 241-255.
Robinson, M.D. (1990). Summer field energetics of the Namib sesert dune lizard
Aporosaura anchietae (Lacertidae), and its relation to reproduction. Journal of
Arid Environments 18, 207-215.
Robson, G.M. & Lambert, M.R.K (1980). Observation on the insects food of
some semi-desert lizards in southem Morocco. Journal ofArid Environments 3,
141-151.
Rohwer, S. (1982). The evolution of reliable and unreliable badges of fighting

ability. American Zoolog,st 22, 531-546.
Rohwer, S., Fretwell, S.D. & Niles, D.M. ( 1980). Delayed maturation in passerine
phnnages and the deceptive aquisition of resources. American Naturalist 115,
400-437.
Rohwer, S. & Roskraft, E. (1989). Results of dyeing male yellow-headed

blackbirds solid black: implications for arbitrary identity badge hypothesis.
Behavioural Ecology and Sociobiology 25, 39-48.
Rose, B. (1982). Lizard home ranges: methodology and functions. Journal of

Herpetology 16, 253-269.
Roskraft, E. & Rohwer, S. (1987). An experimental study of the function ofthe

red epaulettes and the black body colour of male red-winged blackbirds. Animal
Behaviour 35, 1070-1077.
Roughgarden, J. (1995). Ano/is.Lizards qfthe Caribbean: Ecology, Evolution,

and Plate Tectonics. Oxfurd University Press Oxford, United Kingdom,
295
REFERENCES
Roughgarden, J., Porter, W. & Hecke4 D. ( 1981 ). Resource partitioning of space

and its relationship to body temperatures in A,wlis lizard populations.. Oecologia
50, 256-264.
Ruby, D.E. (1981). Phenotypic correlates of male reproductive success in the

lizard Sceloporus jarrovi.. In Natural Selection and Social Behaviour: Recent
Research and New Theory. (Alexander, R.D. and Tinkle, D. W., eds.), Chiron
Press, New York, New York, pp. 96-107.
Ruby, D.E. (1984). Male breeding success and differential access to females in
Ano/is carolinensis.. Herpetologica 40, 272-280.
Runnne4 J.D. & Roughgarden, J. (1985). Effects ofreducingperch-height

separation on competition between two An.olis lizards. Ecology 66, 430-444.
Rutowski, R.L. (1982). Epigamic selection by males as evidenced by courtship

partner preferences in the checkered white butterfly (Pieris protodice). Animal
Ryan, M.J. (1985). The Tungara Frog. Chicago University Press Chicago, Illinois,
Ryan, M.J. (1990). Sexual selection, sensory systems and sensory exploitation.
Oxford Surveys in Evolutionary Biology 7, 157-195.
Ryan, M.J., Fox, J.H, Wilczynski, W. & Rand, A.S. (1990). Sexual selection for
sensory exploitation in the frog Physalaemus pustulosus. Nature 343, 66-67.
Ryan, M.J. & Rand, A.S. {1990}. The sensory basis of sexual selection for complex
calls in the tungara frog, Physalaemus pustulosus (sexual selection for sensory
exploitation). Evolution 44, 305-314.
SandeR M. (1989). Ecological energetics, optimal body size and sexual

dimorphism: a model applied to the stoat, Mustela errninea L.. Functional
Ecology 3, 315-324.
Schaeffer, W.H (1934). Diagnosis of sex in snakes. Copeia 1934, 182
296
REFERENCES
Schwarzkopf"; L. & Shine, R (1991). Thermal biology of reproduction in

viviparous skinks, Eulamprus tympanum: Why do gravid females bask more?.
Oecologia 88, 562-569.
Schwarzkopt: L. & Shine, R (1992). Costs of reproduction in lizards: escape

tactics and susceptibility to predation. Oecologia 31, 17-25.
Schluter, D. ( 1988). Estimating the form of natural selection on a quantitative trait.

Evolution 42, 849-861.
Schluter, D. & Smith, J.N.M. (1986). Genetic and phenotypic correlations in a

natural population of song sparrows. Biological Journal of the Linnean Society
29, 23-36.
Schmale, M. C. ( 1981 ). Sexual selection and reproductive success in males of the

bicolor damselfish, Eupomacentrus partitus (Pisces: Pomacentridae). Animal
Behaviour 29, 1172-1184.
Schoener, T.W. (1967). The ecological significance of sexual dimorphism in size in

the lizardAnolis conspersus. Science 155, 474-577.
Schoener, T,W. (1971). Theory of feeding strategies.. Annual Reviews ofEcology

and Systematics 2, 369-404.
Schoener, T.W. & Schoener, A (1978). Estimating and interpreting body-size

growth in some Ano/is lizards. Copeia 1978, 390..405.
Schoener, T.W. & Schoener, A (1980). Densities, sex ratios and population
structure in four species of Bahamian Anolis lizards. Journal ofAnimal &ology
49, 19-53.
Schoener, T.W., Slade, J.B. & Stinson, C.H (1982). Diet and sexual dimorphism
in the vecy catholic lizard genus, Leiocephalus of the Bahamas. Oecologia 53,
160-169.
297
REFERENCES
Schwaner, T.D. (1989). A field study ofthermoregulation in Black Tiger Snakes

(Notechis ater niger: Elapidae) on the Franklin Islands, South Australia.
Schwartz, A ( 1970). A systematic review of Amieva auberi Cocteau (Reptilia,

Teiidae) in Cuba and the Bahamas. Annals of the Carnegie Museum 41, 45-168.
Searcy, W.A (1982). The evolutionary effects of mate selection. Annual Reviews
ofEcology and Systematics 13, 57-85.
Selander, R.K (1966). Sexual dimorphism and differential niche utilization in birds.
Condor 68, 113-151.
Selander, R.K (1972). Sexual selection and dimorphism in birds. In Sexual

Selection and the Descent ofMan (Campbell, B., ed.), Aldine, Chicago, Illinois,
pp. 180-230.
Shapiro, L.E. & Dewsbury, D.A (1986). Male dominance, femal choice and male
copulatory behaviour in two species of voles (Microtus ochrogaster and Microtus
montanus). Behavioural Ecology and Sociobiology 18, 267-274.
Sharrad, R.D. (1969). Studies ofrapid colour changes in some members of the
genus Amphibolurus: Unpublished BSc(Hons) thesis Zoology Department:
University ofAdelaide,
Shea, B. T. (1983). Allometry and heterochrony in the Afiican apes. American

Journal ofPhysical Anthropology 62, 275-289.
Shea, G., Husband, G. & Weigel, J. (1991). Nesting of the Southern Rainforest
Dragon, Hypsilurus spinipes (Squamata: Agaminae}. Herpetofauna 21, 7-10.
Sherman, P.W. & Morton, M.L. (1984). Demography ofBelding's ground

squirrels. Ecology 65, 1617-1628.
Sherman, P.W. & Morton, M.L. (1988). Extra-pair fertilizations in mountain

white-crowned sparrows. Behavioural Ecology and Sociobiology 22, 413-420.
298
REFERENCES
Shine, R. (1978). Sexual dimorphism and male combat in snakes. Oecologia 33,.
269-277.
Shine, R. (1980). 'Costs' of reproduction in reptiles. Oecologia 46, 92-100.
Shine, R. ( 1986). Sexual differences in morphology and niche utilization in an

aquatic snake, Acrochordus arafarae.. Oecologia 69, 260-267.
Shine, R. (1988a). Parental care in Reptiles. Jn Biology of the r~ptilia. Volume 16.
Ecology B. defense and Lifehistory (Gans, C. and Huey, R.B., eds.), Alan R. Liss,
Jnc., New York, New York, pp. 275-330.
Shine, R. (1988b). Tue evolution oflarge body size in females: a critique of

Darwin's "fecuodity advantage" model American Naturalist 131, 124-131.
Shine, R. (1989). Ecological causes for the evolution of sexual dimorphism: a

review. Quarterly Revie.vs in Biology 64, 419-461.
Shine, R. (1990). proximate determinants of sexual differences in adult body size.

Shine, R. (1991 ). Why do larger snakes eat larger prey items?. Functional
Ecology 5, 493-502.
Shine, R. (1994). Sexual dimorphis.min snakes revisited. Copeia 1994, 326-346.
Shine, R. & Fttzgerald, M. (1996). Variation in mating systems and sexual size
dimorphism between populatons of the Australian.python Morella spiiota
(Serpentes: Pythonidae). Oecologia 103, 490-498.
Shine, R. & Lambeck, R. (1985). A radiotelemetric study of movements,

thermoregulation and habitat utilization of Arafura filesnakes (Serpentes,
Acrochordidae). Herpetologica 41, 351-361.
Shine, R. & Madsen, T. (1994). Sexual dichromatism in snakes ofthe genus

Vipera: a review and a new hypothesis. Journal ofHerpetology 28, 114-117.
299
REFERENCBS
Sievert, L.M. & Hutchinson, V.I! (1989). Influences of season, time of day, light
and sex on the thermoregulatocy behaviour of Crotaphytus collaris. Journal of
Thermal Biology 14, 159-165.
Sigmund, W.R. ( 1983). Female preference for Ano/is carolinensis males as a

function of dewlap color and background coloration. Journal ofHerpetology 17,
137-143.
Silverman, RB. & Dunbar, M.J. (1980). Aggressive tusk use by the narwhal
(Monodon manoceros L.). Nature 284, 57-58.
Simon, C.A (1975). The influence fo food abundance on territry size in the iguanid
liz.ard Sceloporus jarrovi. Ecology 56, 993-998.
Simon, C.A & Bissinger, B.R (1983). Paint marking liz.ards: Does the color atfect
SUIVivorship?. Journal ofHerpetology 17, 184-186.
Sinervo, R & Adolph, S.C. (1989}. Thermal sensitivity of growth in hatchling

Sce/oporus liz.ards: Environmental, behavioural and genetic aspects. Oecologta
78, 411-419.
Slatkin, M. (1934). Ecological causes of sexual dimorphism.. Evolution 38, 622-

630.
Smith, J. (197.M. Hatching bearded dragon eggs. South Australian Herpetologist

2, 12
Smith, M.A (1935). The Fauna ofBritish India, including Ceylon and Burma.
Reptilia and Amphibia. II. Sauria.. Taylor ~ Francis London, United Kingdom,
Smith, R. (Ed.) (1984). Sperm Competition and the Evolution ofAnimal Mating
Systems. Academic Press Orlando, Florida,
Smyth, M. & Smith, M.J. (1968). Obligatory sperm storage in the skinkHemiergis
peronii. Science 161, 575-576.
300
REFERENCES
Sokal, R.R. & Rohlf; F.J. (1981). Biometry. W.I-L Freeman San Francisco,
California,
Sorenson, L.G. & Derrickson, S.R. (1994). Sexual selection in the northern pintail
(Anus acuta): the importance of female choice versus male-male competition in the
evolution of sexually-selected traits. Behavioural Ecology and Sociobiology 35,
389-400.
Sporn, C.C. (1955). The breeding of the mountain devil in captivity. Westem
Australian Naturalist 5, 1-5.
Spotila, J.R., Lommen, P.W., Bakken, G.S. & Gates, D.M. (1973). A
mathematical model for body temperature of large reptiles: implications for
dinosaur ecology. American Naturalist 107, 391-404.
Stamp-Dawkins, M. & Guilford, T. (1991). The corruption ofhonest signalling.

Stamps, J.A (1977). The function of the survey posture in Ano/is lizards. Copeia
1977, 756-758.
Stamps, J:A. (1983). Sexual selection, sexual dimorphism and territoriality. In

Lizard Ecology: studies ofa model organism (Huey, RB., Pianka, E.R. and
Schoener, T. W., eds.), Harvard University Press, Cambridge, Massachusetts, pp.
169-204.
Stamps, J.A (1993). Sexual size dimorphism in species with asymptotic growth
after maturity. Biological Journal of the Linnean Society 50, 123-145.
Stamps, J.A ( 1995). Using growth-based models to study behavioural factors

affecting sexual size dimorphism Herpetological Monographs 9, 75-87.
Stamps, J.A., Krishnan, V.V. & Andrews, R.M. (1994). Analyses of sexual size
dimorphism using null growth-based models. Copeia 1994, 598-613.
Stamps, J.A. & Tanaka, S. (1981). The influence of food and water on growth
rates in a tropical lizard (Ano/is aeneus). Ecology 62, 33-40.
301
REFERENCES
Steams, S.C. (1992). The Evolution ofLife Histories. Oxford University Press
Oxford, United kingdom,
Stevenson, R.D. (1985). Body size and limits to daily range ofbody temperature in
terrestrial ectothenns. American Naturalist 125, 102-117.
Storr, G.M. (1965). The Amphibolurus maculatus species-group (Lacertilia,

Agamidae) in Western Australia. Journal ofthe Royal Society of Western
Australia 48, 45-54.
Storr, G.M. (1967). Geographic races ofthe agamid lizards Amphibo/urus

caudicinctus. Journal of the Royal Society of Western Australia 50, 49-56.
Storr, G.M., Smith, L.A & Johnstone, R.E. (1983). Lizards of Western Australia.
fl Dragon Lizards and Goannas. Western Australian Museum Nedlands, Western
Australia,
Sutherland, W.J. (1985). Measures of sexual selection. Oxford Surveys in

Evolutionary Biology 1, 90-101.
Telford, S.R. &Dyson, ML. (1988). Some determinants ofthemating system in a

population ofpainted reed frogs (Hyperolius marmoratus). Behaviour 106, 265-
278.
Thompson, C.W. & Moore, M.C. (1991). Throat colour reliably signals status in
male tree lizards, Urosaurus ornatus. Animal Behaviour 42, 745-753.
Thompson, M.B. (1993). Estimate of the population structure of the Eastern

Water Dragon, Physignathus lesueurii (Reptilia: Agamidae), along riverside
habitat. Wildlife Research 20, 613-619.
Thornhill, R. (1976). Sexual selection and paternal investment in insects.

Thornhill, R. (1976). Sexual selection and nuptial feeding behaviour in Bittacus

apicalis (Insecta: Mecoptera). American Naturalist 110, 529-548.
302
REFERENCES
Thornhill, R ( 1980). Competitive, charming males and choosy females: Was

Darwin correct? Florida Entomologist 63, 5-30.
Thornhill, R & Alcock, J. ( 1983). The Evolution ofInsect Mating Systems.

Harvard University Press Cambridge, Massachusetts,
Tinkle, D.W. (1967). The life and demography of the side-blotched lizard, Uta
stansburiana. Miscellaneous publications of the Museum ofZoology, University
o/Michagan 132, 1-182.
Tinkle, D.W. & Ballinger, RE. (1972). Sceloporus undulatus: a study of the
intraspecific comparative demography of a lizard. Ecology 53, 570-584.
Toft, C.A (1985). Resource partitioning in amphtoians and reptiles.. Copeia 1985,
1-21.
Tokarz, RR (1985). Body size as a factor determining dominance in staged

agonistic encounters between male brown anoles (Ano/is sagrei). Animal
Tokarz, RR (1992). Male mating preference for nnfiuniliar females in the lizards,
Ano/is sagrei. Animal Behaviour 44, 843-849.
Tokarz., R.R. (1995). Mate choice in lizards: a review. Herpetologicat

Monographs 9, 17-40.
Tracy, C.R. (1982). Biophysical modeling in reptilian physiology and ecology. In

Biology of the Reptilia Volume I 2. Physiology C: Physiological Ecology. (Gans,
C. and Pough, F.H., eds.), Academic Press, New York, New York, pp. 275-315.
Tracy, C.R. & Christian, K.A (1986). Ecological relations among space, time and
thermal niche axes. Ecology 67, 609-615.
Trillmich, K.G.K. (1983). The mating system of the marine iguana (Amblyrhynchus
cristatus). Zietschrift Tierpsychologie 63, 141-172.
303
REFERENCES
Trivers, R.L. (1972). Parental Investment and Sexual Selection. In Sexual

Selection and the Descent ofMan 1871-197I (Campbell, B., ed.), Aldine,
Chicago, Illinois, pp. 136-.179.
Trivers, R.L. (1976). Sexual selection and resource-accruing abilities in Ano/is

garmani. Evolution 30, 2530269.
Tyler, M.J. (1960). Observations on the diet and size variation ofAmphibolurus
adelaidensis (Gray)(Reptilia: Agamidae) on the NuJlarbor Plain. Transactions of
the Royal Society of South Australia 83, 111-117.
Van Sluys, M. (1993). Food habits of the lizard Tropidurus itambere

(Tropiduridae) in southeastern Brazil.. Journal ofHerpetology 27, 347-351.
Viai J.L. & Stewart, J.R. (1989). The manifestation and significance of sexual .
dimorphism in anguid lizards: a case study of Barisia montirola. Canadian
Journal a/Zoology 67, 68-72.
Vttt, L.J. ( 1983). Reproduction and sexual dimorphism in the tropical teiid lizard
Cnemidophorus ocellifer. Copeia 1983, 359-366.
Vitt, L.J., Congdon, J.D., Hulse, AC. & Platz, I.E. (1974). Territorial aggressive
encounters and tail breaks in the lizard Sceloporus magister. Copeia 1974, 990-
993.
Vitt, L.J. & Cooper, W.E. (1985). The evolution of sexual dimorphism in the skink
Eumeces laticeps: an example of sexual selection. Canadian Journal of Zoology
63, 995-1002.
Vitt, L.J. & Zani, P.A (1996). Ecology of the elusive tropical lizard Tropidurus (=
Uracentron)jlaviceps (Tropiduridae) in the lowland rainforest of Ecuador.
Herpetologica52, 121-132.
Vollrath, F. & Parker, G.A (1992). Sexual dimorphism and distorted sex ratios in
spiders. Nature 360, 156-159.
304
REFERENCES
Von Schantz, T., Goransson, G., Andersson, G., Froberg, I., Grahn, M., Helgee,
A & Wittzell, H. (1989). Female choice selects for a viability-based male trait in
pheasants. Nature 337, 166-169.
Wade, M.J. & Arnold, S.J. (1980). The intensity of sexual selection in relation to
male sexual behavionr, fem.ale choice and sperm precedence. Animal Behaviour
28, 446-461.
Wade, MJ. & Kal.w; S. (1990). The causes ofnatnral selection. Evolution 44,
1947-1955.
Wallace, A.R. (1889). Danvinism. MacMillan London, United kingdom,
Warburg, M.R. (1966). Water economy of several Australian geckos, agarnids and
skinks. Copeia 1966, 230-235.
Watkins, G. G. ( 1996). Proximate caises of sexual size dimo:rphism in the Iguanian

lizardMicrolophus occipita/is. Ecology 11, 1473-1482.
Wells, KD. (1978). Territoriality in the green frog (Rana climitans): vocaliz.aions
and agonistic behaviour. Animal Behaviour 26, 1051-1063.
Werner, D.I. ( 1978). On the biology of Tropidurus delanonis Baur (lguanidae).

Zietschrift Tierpsychologie 41, 337-395.
Westneat, D.F. (1987). Extra-pair fertilizations in a predominantly monogamous

bird: genetic evidence. Animal Behaviour 35, 877-886.
Whitfield, D.P. (1986). Plumage variability and territoriality in breeding turnstone

Arenaria interpres: status signalling or individual recognition?. Animal Behaviour
34, 1471-1482.
Wikelsld, M., Carbone, C. & Trillmch, F. (1996). Lekking in marine iguanas:

female grouping and male reproductive strategies. Animal Behaviour 52, 581-596.
\Viley, R.H ( 1974). Evolution of social organisation and life history patterns
among grouse. Quarterly Reviews in Biology 49, 201-227.
305
REFERENCES
Williams, E.E. & Rand, AS. ( 1977). Species recognition, dewlap function and
fauna! size. American Zoologist 17, 261-270.
Willmer, P. (1991). Thermal biology and mate acquisition in ectotherms. Trends in

Ecology and Evolution 6, 396-399.
Witten, G.J. (1974). Population movements of the agamid lizardAmphibolurus

nobbi. Australian Zoologist 18, 129-132.
Witten, G.J. & Coventzy, A.J. ( 1984). A new lizard of the genus Amphibolurus
(Agamidae) from southern Australia. Proceedings of the Royal Society of Victoria
96, 155-159.
Woolbright, L.L. (1983). Sexual selection and size dimorphism in anuran ampb.t"bia.
American Naturalist 121, 11()..119.
Woolbright, L.L. (1989). Sexual dimorphisminEleutherodactylus coqui: selection

pressures and growth rates. Herpetologica 45, 68-74.
Wright, S. ( 1931 ). Evolution in Mendelian populations. Genetics 16, 97-159.
Zahavi, A (1975). Mate selection - a selection for a handicap. Journal of

Theoretical Biology 53, 205-214.
Zahavi, A (1977). The cost ofhonesty (further remarks on the handicap principle).
Journal ofTheorettcal Biology 61, 603-605.
Zeh, D. (1987). Aggression, density, and sexual dimorphism in chemetid

pseudoscorpions (Arachnida: Pseudoscorpionidae). Evolution 41, 1072-1087.
Znari, M. & El Monden, E. (in press). Seasonal changes in the diet of adult and
juvenile Agama impalearis (Sauria, Agamidae) in the central Jbilet Mountains of
Morocco.. Herpetological Journal
306
REFERENCES
Zucker, N. (1986). Perch height preferences of male and female tree lizards,
Urosaurus ornatus: a matter of food competition or social role?. Journal of
Herpetology 20, 547-553.
Zucker, N. (1989). Dorsal darkening and territoriality in a wild population of the

tree lizard, (Urosauros ornatus). Journal ofHerpetology 23, 389-398.
Zucker, N. (1994). Social influence on the use of a modifiable status signal

307
APPENDICES
APPENDIXA
Abstracts published during my tenure at Flinders University.
I. Jolmston G.R. & N.A Locket 1992. An undescnoed Urodacus (Scorpionida) from
Eyre Peninsula, South Australia. XII International Congress qfArachno/ogy,
Queensland Museum, Queensland
2. Jolmston G.R. & Bennett M.J. 1992. Facultative brood reduction in the Australian
Pelican (Pe/ecanus conspicil/atus). Ecological Society ofAustralia Conference,
University ofAdelaide,, South Australia.
3. Bennett M.J. & Jolmston G.R. 1992. Breeding biology of the Black-faced Cormorant
(Leucocarbo fuscescens). Ecological Society ofAustralia Conference, University of
Adelaide,, South Australia.
4. Jolmston G.R. 1994. Sexual dbnorphism and mating system in the agamid lizard
Ctenophorus jionni. Second World Congress ofHerpetology, University ofAdelaide,
South Australia.
5. Jolmston G.R. 1994. Sexual selection in the iz.ard Ctenophorusjionni: the role of mate
choice and male combat. Second World Congress ofHerpetology. University of
Adelaide, South Australia.
6. Jolmston G.R. & Richards S.J. 1994. Community structure of anuran amph.toiaos in the
Ok Tedi region of Papua New Guinea. Second World Congress ofHerpetology,
University ofAdelaide, South Australia.
7. LaNafie N., Johnston G.R. & Prager R.H. 1994. Chemical properties of femoral gland
secretions among populations of the agamid lizard Ctenophorus fiom1i. Second World
Congress ofHerpetology, University ofAdelaide, South Australia.
8. LaNafie N., Johnston G.R. & Prager R.H. 1994. Chemical properties of femoral gland
secretions among populations of the agamid lizard Ctenophorusfionni. Australian
Society ofOrganic Chemistry, University of Woolongong, New South Wales. ·
9. Johnston G.R. 1995. Sequential assessment and male conflict in the tenitorial lizard
Ctenophorus ftonni. XXIV International Ethological Conference, University aj
Hawaii, Hawaii.
10. Johnston G.R. 1996. Tenitoriality and Sexual Selection in the Peninsula Dragon
Lizard. lntemational Behavioural Ecology Congress, Australian National University,
Australian Capital Territory.
11. Johnston G.R. 1996. Sexual dimorphism in the New Guinean Hylid Frog, Litoria
havina. lnternational Behavioural Ecology Congress, Australian National University,
Australian Capital Territory.
APPENDIXB
Papers published during my tenure at Flinders University.
1. Johnston, G.R. 1992. Ctenophorus tjantjalka, a new dragon lizard (Lacertilia:

Agamidae) from northern South Australia. RR.cords of the South Australian Museum
26:51-59.
2. Johnston G.R. 1992. Relictual population of Tiliqua scincoides ( Sauria: Scincidae) in

north western South Australia. Transactions of the Royal Society of South Australia
116:149-150.
3. Richards S.J., Johnston G.R. & Burton, T.C. 1992. A new species ofmicrohylid frog
(genus Cophixalus) from the Star Mountains, New Guinea. Science in New Guinea.
18:141-145.
4. Johnston G.R. & Richards S.J. 1993. Obsei:vations on the breeding biology of a
rnicrohylid frog (genus Oreophryne) from New Guinea. Transactions of the Royal
Society of South Australia 117: 105-107.
5. Gregory P.A & Johnston G.R. 1993. Birds of the cold tropics: Dokfoma, Star
Mountains. Bulletin of the British Ornithological Club 113:139-145.
6. Richards S.J. & Johnston G.R. 1993. A new species of Nyctimystes (Anura: Hylidae)
from the Star Mountains, Papua New Guinea. Memoirs of the Queensland Museum
33:73-76
7. Johnston G.R. 1994. Seasonal changes in dorsal reflectance oftwo species of
Australian elapid snake. Journal ofHerpetology 28: 109-112
8. Johnston G.R. & Richards S.J. 1994. Bird obseIVations from the Western Province,
July 1993. Murok 6:9.
9. Peterson M., Shea G.M., Johnston G.R. & Miller, B. 1994. Notes on the morphology
and biology ofCtenophorus mckenziei (Storr 19SlXLacertilia: Chamaeleonidae:
Agamidae). Transactions of the Royal Society ofSouth Australia 118:237-244.
IO. Richards S.J., Johnston G.R. & Burton T.C. 1994. A remarkable new species of
microhylid frog (genus Asterophrys) from the Star Mountains of Papua New Guinea.
Memoirs of the Queensland Museum 34:281-286.
11. Johnston G.R. & Richards S.J. 1994. A new species ofLitoria (Anura: Hylidae) from
the Star Mountains ofNew Guinea, with a redefinition ofLitoria leucova: Memoirs of
the Queensland Museum 34:273-279.
12. La Nafie N., Prager R.H. &Johnston G.R. 1995. Chemical characterisation of races of
the peninsnla dragon lizard, Ctenophorus ftonni. Asian Coordinating Group for
Chemistry Chemical Research Communications 3:40-44.
13. Johnston G.R. 1996. Genetic and seasonal variation in body colour ofthe death adder,
Acanthophis antarcticus (Squamata: Elapidae). Journal ofZoology (London)
239:187-196.
UNEMPLOYED at last!
Scientifically, such a contingency can never have befallen of

itself. According to one theory of the Universe, the momentum of
Original Impress has been tending toward this far-off, divine event
ever since a scrap of fire-mist ftew from the aolar cen,tre to form our
planet. Not this event alone, of course; hut every occurrence, past
and present, from the fall of captured Troy to the fall of a captured
insect. According to another theory, I hold an independent diploma
as one of the architects of our Social System, with a commission to
·use my own judgment, and take my own risks, like any other unit
of humanity. This theory, unlike the first, entails frequent hitches
and cross-purposes; and to some malign operation of these I ahould
owe my present holiday. ,
SuchisLife
Joseph Furphy

Behavioural Ecology of The Peninsula Dragon Lizard, Ctenophorus Fionni

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BEHAVIOURAL ECOLOGY OF

THE PENINSULA DRAGON LIZARD

School of Biological Sciences

A thesis submitted for the degree of Doctor of Philosophy

For Barbara Johnston (nee Baker)

I certify that this thesis does not incorporate without

I believe that this thesis is properly presented and conforms to the

A mature male Peninsula Dragon, Ctenophorus fionni

2. Sexual Dimorphism ........... ................... ... ......... ... .... 37

3. Reproduction . . ... . .. . .. . . . .. . . .. . .. .. . .. . .. . ... .. . . .. .. . .. .. . . . . .. . . . 70

4. Growth and Survivorship: Proximate causes of SexuaJ

5. Ecological Differences between Males and Females ...

6. ].\,fating System and the Opportunity for Sexual

7. Targets of Sexual Selection in Males ........................ 210

8. Mate Choice or Male Competition ? . .. . ... .. .. .. . . . .. ... .. . 240

9. Synthesis ............................................................... 258

References ....................................................................... 268

This project was formally supervised by Associate Pr'ofessor C.

Michael Bull, and was funded by an Australian Postgraduate

Research Scholarship, a Flinders University Scholatship and an Ethel

Mats Olsson unwittingly provided the inspiration for this project,

long ago, in a little caravan, somewhere near Lake Eyre.

Errol, Elizabeth & Mark Carmody allowed me to work on their

property. Andrew & Vicki Gohl provided welcome showers,

conversation, meals in Cowell whenever I got sick of staring at

bloody lizards during field trips. The Whyalla Amatuer Reptile

Keepers Association, Ben Baghurst, Anne Brow, Paula Dempsey,

Richard Evans, Rachel Furbank, Michelle Hale, Brett Johnston, Colin

Kranz, Cra,ig Martins, Paul Mazdon, and Elizabeth Yeatman helped

Jon Havenhand, Peter Hudson, Mark Jerisevic, Greg Kirby, Louise

Lawson, Duncan Mackay, Jim Mitchell, Yvonne Pamula, Rolf Prager,

and Tatia Zubrinich contributed to the project in various ways.

Grant Fleming and Wendy Sysouphat provided office and laboratory

facilities at the Whyalla Campus of the University of South Australia.

Colin Kranz helped in many ways to establish the basic reproductive

biology of Peninsula Dragons.

Martins for keeping open houses to me and miscellaneous hangers

Dempsey and Jeremy Roberston read several chapters each. Paula

also provided support in many other ways.

This thesis aimed to explain the evolution of sexually dimorphic

Peninsula Dragons mate and reproduce during the austral spring.

The pattern of sexual size dimorphism in Peninsula Dragons appears

There is no evidence for the hypothesis that sexually dimorphic traits

The mating system of Peninsula Dragons is best described as

A selection analysis identified body size and head width as

An experimental manipulation in the field showed that there was

In conclusion, sexual selection is the only hypothesis that appears to

Explaining the evolution of sexual dimorphic traits has. been a major

Examples are sexual differences in body size in Elephant Seals

The extent to which sexually dimorphic traits require a separate

EVOLUTION, COMPONENTS OF FITNESS AND

Evolution refers to changes in gene frequencies between generations.

Selection occurs if there is a relationship between the expression of a

Traits which show an evolutionary response to selection may be

Variation in fitness due to differential mating success is referred to

A key reason for distinguishing between these three types of

To understand the evolution of any trait completely it is necessary to

offspring, and offer a fecundity advantage. Thus body size may be a

Formal selection analyses using the approach developed by Arnold

Measures of phenotypic selection are mosi: useful when combined

marmoratus) have shown: a clear preference among females for male

EXPLANATIONS FOR SEXUALLY DIMORPHIC

Sexual selection is the pre-eminent explanation used for sexually