BREVIORA

Museum of Comparative Loology

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US ISSN 0006-9698

CAMBRIDGE, MAss. 7 SEPTEMBER 2012 NUMBER 532

ECOLOGY OF THE FLAP-NECKED CHAMELEON CHAMAELEO DILEPIS IN
SOUTHERN AFRICA

LEEANN T. REANEY,! STEPHEN YEE,” JONATHAN B. Losos,? AND MArTIN J. WHITING*

ABSTRACT. We quantified sexual size dimorphism, reproduction, and diet in the flap-necked chameleon,
Chamaeleo dilepis, using museum specimens. Females were larger than males in both snout-vent length (SVL) and
pelvic width. The smallest sexually reproductive female was 80 mm SVL, whereas the smallest mature male was
60 mm. Female body size also correlated with clutch size (mean: 44.2, range: 19-74) and volume, suggesting the
female-biased size dimorphism may be the product of fecundity selection. Males and females have slightly

asynchronous reproductive cycles but breed during spring—-summer.
but their diet is dominated both numerically and volumetrically by orthopterans,

arthropods,
coleopterans.

KEY WORDS:

Despite their fascinating characteristics,
the life history and ecology of chameleons
has been little explored, leading to a signif-
icant gap in our understanding of lizard

'School of Biological and Chemical Sciences, Queen
Mary, University of London, Mile End Road, London
E3 5QX, U.K.; e-mail: leeann.reaney@gmx.com.
*Plant and Microbial Biology, University of California,
Berkeley, California 94720, U.S.A.; e-mail: sfyee@
berkeley.edu

> Department of Organismic and Evolutionary Biology,
Harvard University, Cambridge, Massachusetts 02138,
U.S.A.; e-mail: jlosos@oeb.harvard.edu.

*Department of Biological Sciences, Macquarie Univer-
sity, Sydney, NSW 2109, Australia; e-mail: martin.
whiting@mq.edu.au.

Chamaeleo dilepis feeds on a range of
followed by

Chamaeleo dilepis; sexual dimorphism; reproduction; diet; flap-necked chameleon

ecology. Chameleons have evolved a unique
set of traits that allow them to occupy a
highly specialized niche. Typical chameleons
have telescopic eyes, fusion of some digits, a
prehensile tail, and a “ballistic” tongue that
can exceed the length of their body, and they
are capable of rapid color change (Tilbury,
2010). These features allow chameleons to
hunt with stealth and may enable them to be
less conspicuous to predators during prey
capture.

While the unique form and function of
chameleons has attracted considerable atten-
tion, we still have only a cursory grasp of their
ecology and life history, although consider-

© The President and Fellows of Harvard College 2012.

i)

able work has been done on reproduction and
embryonic development in Chamaeleo calyp-
tratus (e.g., Andrews, 2005, 2007, 2008), and
we know the female reproductive cycle for
Chamaeleo chamaeleon (Cuadrado and Lo-
man, 1999). We also know the foraging mode
of two species from different genera. Brady-
podion pumilum has been classified as a
“cruise” forager because it spends a high
enough proportion of its time moving (21%)
to qualify as an active forager, but it makes
very few moves per minute (Butler, 2005).
Similarly, an invasive population of Jackson’s
chameleon (Chamaeleo jacksoni xanthopholis)
in Hawaii spends 19.7% of its activity time in
movement, with few moves per minute (¥ =
0.24; Hagey et al., 2010). These results are
surprising given the highly specialized mor-
phological features consistent with ambush
foraging and suggest that chameleons employ
a foraging mode quite distinct from those
of most other lizards. In terms of sexual
dimorphism, the males of many species have
horns or protrusions that function as either
armaments or ornaments, and males can be
larger than females (Necas, 2001; Karsten ef
al., 2009). In other species, including most of
the dwarf chameleons (Bradypodion), females
are larger than males, most likely as a result of
fecundity selection (Stuart-Fox, 2009).

Here, we describe the ecology of the flap-
necked chameleon, Chamaeleo dilepis, with
particular reference to sexual dimorphism,
reproduction, and diet using specimens from
the Ditsong National Museum of Natural
History (formerly the Transvaal Museum) in
South Africa. Chamaeleo dilepis is a large
chameleon native to savannah woodland and
sometimes coastal forest in sub-Saharan
Africa (FitzSimons, 1943; Brain, 1961; Pie-
naar, 1978; Branch, 1998; Tolley and Burger,
2007). This species is typically green or
brown and usually possesses a pale stripe
on the lower flanks and one to three pale
patches higher on the flanks (Fig. 1; Tilbury,

BREVIORA

No. 532

2010). Interestingly, in the dry season in
Kenya males tend to be brown and use
defoliated vegetation, whereas females are
mostly green and occupy leafy foliage. This
difference falls away in the wet season when
most individuals are green and occupy leafy
vegetation (Hebrard and Madsen, 1984).
Females are reported to be larger than males,
although this has never been quantified
(Brain, 1961; Necas, 2001; Tilbury, 2010).

Chamaeleo dilepis in the wild lay their eggs
in late summer in a tunnel 150-300 mm deep,
excavated in damp soil. The eggs can take up
to 377 days to hatch depending on environ-
mental conditions (Brain, 1961; Wager, 1983;
Branch, 1998). After egg laying, females are
emaciated and generally vulnerable to a wide
variety of predators that include various
snakes, mongooses, raptors, hornbills, and
monkeys (Pienaar, 1978; Branch, 1998; Van
Wyk and Els, 2004; Cunningham and Adank,
2005; Tilbury, 2010). Chamaeleo dilepis has a
broad distribution, and while a number of
subspecies are recognized, their systematic
and taxonomic status is uncertain (see Tilbury
[2010] for a detailed discussion of their
taxonomy). We examined only material from
the southern African subregion (south and
east of Kunene River separating Namibia
from Angola and including southern Mozam-
bique; see Appendix 1) and provide the first
quantitative assessment of diet, reproduction
and morphology.

MATERIALS AND METHODS

Dimorphism

To quantify sexual size dimorphism, we
measured morphological traits from a subset
of mature individuals (80 of the 251 speci-
mens; see below for maturity criteria). Snout-
vent length (SVL) was measured from the tip
of the snout to the anterior edge of the cloaca

2012

ECOLOGY OF CHAMAELEO DILEPSIS 3

Figure 1.
and a row of raised triangular tubercles extending from the tip of the snout down the ventral midline. The southern
African forms look identical. Photo by M. J. Whiting.

and tail length from the posterior edge of the
cloaca to the tip of the tail. Head height was
measured as the distance between the center
of the parietal bone to the bottom of the
articular. Head width was measured at the
widest point of the head. Head length was
measured from the tip of the snout to the
back of the skull. The femur, tibia, humerus,
and ulna were measured from the proximal
to distal ends of the bones. Pelvic width was
measured between the outer edges of the iliac
bones. SVL and tail length were measured
with a ruler to the nearest | mm, whereas
digital calipers were used to measure the
remaining morphological measurements to

A typical Chamaeleo dilepis from Chuka, Kenya, with characteristic pale flank patches, occipital flaps,

the nearest 0.01 mm. Before testing for
differences in sexual dimorphism, we re-
moved the effect of size by regressing each
morphological measurement on SVL. We
made pairwise plots to determine if there
were any outliers and removed one specimen
from the analysis because it was an outlier
in the majority of the plots. A principal
components analysis was then performed on
SVL and the residuals resulting from the
regression of each of the measured characters
on SVL. We retained the first three principal
components because they explained the ma-
jority of the variation. We then used a
multivariate analysis of variance (MANOVA)

4 BREVIORA

on the first three principal components to
determine whether trait variation among
individuals of C. dilepis could be explained
significantly by sex.

Reproduction

We made a mid-ventral incision to expose
the gonads and score sexual maturity. Males
were recorded as sexually mature if they
contained enlarged testes and convoluted
epididymides. We measured the right testis
for length and width to the nearest 0.01 mm
using digital calipers. Testis volume (right
testis) was then determined using the formula
for-a prolate spheroid (Vitt et al, 1993; Vitt
and Zani, 2005):

volume =4/3z(length/2) « (width /2)

We scored females as sexually mature
if they contained oviductal eggs, enlarged
vitellogenic follicles or a convoluted oviduct.
For gravid females (sexually active), oviductal
eggs were counted and the size (length and
width) of each egg was measured to the
nearest 0.01 mm using digital calipers. Ovi-
ductal egg volume was then calculated using
the formula for a prolate spheroid (as above).

Diet

We removed stomach contents and spread
them on a Petri dish for identification to
order. The length and width of all intact prey
items were measured to the nearest 0.01 mm
using digital calipers. Prey volume was cal-
culated using the formula for a prolate
spheroid (as above).

We estimated niche breadth using the
reciprocal of Simpson’s (1949) diversity
measure,

No. 532

where 7 = resource category, p = proportion
of resource category 7, and n = total number
of categories. Values vary from 1 (exclusive
use of one prey type) to n (even use of all
prey types). Prey categories were the arthro-
pod orders of prey items found in the
stomach contents.

All data are reported as ¥ + 1 SE and
differences were considered significant when
alpha was < 0.05.

RESULTS

Dimorphism

The first three principal components (PCs)
explained 97% of the variation (Table 1),
with PCI explaining 79.5% of the variation,
PC2 14.5%, and PC3 3.2%. PC1 loads only
for body size (as represented by SVL); PC2
correlates most strongly with tail length, and
to a lesser extent by head length and ulna
length; and PC3 correlates primarily with
tibia length, femur length, humerus length,
ulna length, head height, and head length.

The sexes differ in position in the multi-
variate space defined by these axes (MAN-
OVA F375 a 3.899, P= 0.012), with the first
principal component being highly significant
(Fi 75 = 7.613, P = 0.007), the second
component only tending toward significance
(F, 75 = 3.300, P = 0.073), and the third
being nonsignificant (F,75 = 0.783, P =
0.38). Females were significantly larger than
males, with the largest female measuring
162 mm SVL, whereas the largest male was
only 126 mm (Table 2).

Reproduction

The smallest sexually mature female was
80 mm SVL (Fig. 2). Clutch size (n = 25)
averaged 44.2 + 14.7 (range, 19-74). Female
SVL was significantly positively correlated
with both clutch size (7? = 0.472, Fy24 =

2012 ECOLOGY OF CHAMAELEO DILEPSIS 5

TABLE 1. FACTOR LOADINGS OF THE FIRST THREE PRINCIPAL
COMPONENTS ON SNOUT-VENT LENGTH (SVL) AND THE
RESIDUALS RESULTING FROM THE REGRESSION OF EACH OF THE
OTHER 13 TRAITS ON SVL MEASURED IN MALE AND FEMALE
C. DILEPSIS. TRAITS WITH THE HIGHEST LOADINGS ARE
IN ITALICS.

Factor I Factor II Factor III

Snout—vent

length 1.000 0 0
Head length 0 0.508 0.673
Head width 0 0:275 0.600
Head height 0 0.362 0.698
Tail length 0 0.990 — 2839
Pelvis width 0 0.122 0.374
Left ulna 0 0.571 0.635
Left humerus 0 0.386 0.617
Right ulna 0 0.561 0.592
Right humerus 0 0.471 Us
Left tibia 0 0.324 0.720
Left femur 0 0.415 0.610
Right tibia 0 0.406 0.726
Right femur 0 0.296 0.597
Eigenvalue 353.681 64.411 14.001
Explained

variance (%) 195 14.5 32

20.587, P < 0.001) and total egg volume (r?
= 0.194, F; 54 = 5.53, P = 0.028), but not the
largest egg measured (7? = 0.015, Fy>.4 =
0.359, P = 0.55). Seasonal variation in

ovarian volume of the largest follicle showed
that females start to produce eggs in
November, tapering off in May. A single
female contained a clutch of developed eggs
in August (Fig. 3A).

The smallest sexually mature male mea-
sured 60 mm SVL (Fig. 2). Log testicular
volume increased with log SVL (r? = 0.349,
Ei =. 57.361, PP < 0.001). Testicular
volume was largest from September to
February (spring and summer), with a few
males showing enlarged testes in May, June,
and July (Fig. 3B).

Diet

We identified 13 prey categories in the
stomachs of 206 C. dilepsis (the remaining 45
were empty; Table 3, Fig. 4A, B). Although
the majority of prey items consumed were
arthropods, a snail and two reptiles (a skink
and a gecko) were found in three individuals
(Fig. 4A). Most of the arthropods consumed
were insects (85.4%). Orthopterans dominat-
ed the diet, making up 36.4% numerically
(Fig. 4A) and 68.9% volumetrically (Fig. 4B),
as well as being consumed by most chame-
leons (37.5%). Coleopterans were also an

TABLE 2. SUMMARY OF SEXUAL SIZE DIMORPHISM IN 14 MORPHOLOGICAL TRAITS IN FEMALE AND MALE C. DILEPIS.

Trait (mm)

Snout—vent length

Tail length
Head height
Head width
Head length
Right femur
Right tibia
Left femur
Left tibia
Right humerus
Right ulna
Left humerus
Left ulna
Pelvis width

Adult Females (n = 41)

paee NS)

LD 3 AN
S759 253.20
2031222 0769
[S49 20.59
PIE = 50.89.
Das: 0.83
18.48 + 0.72
PASO ee) 8.
LS:40;=.0°72
19:03-270:05
18.16 + 0.64
P9132: 0.66
18.32 + 0.64
moF = 0.29

Adult Males (n = 39)

Range Xo SH Range
50.00—140.00 i ee Rees one ea) 58.00—142.00
48 .00—141.00 86.03 + 3.00 57.00—141.00
11.83—32.06 19.74 + 0.60 14.37-29.18

9.04-23.26 14.67 + 0.51 10.10—22.26
17.63-42.89 28.38 = 0.92 20.44-43.58
10.98—36.84 AN BS Rue 13.65—-34.98
9.08—28.26 17.10 + 0.60 10.52—27.89
10.54—-35.50 20.04 + 0.73 13.28-32.34
9.26-28.75 7.24 = 0.57 11.48—27.00
9.87-28.21 13.317 =50:62 11.70—28.32
9.94-27.77 Ki5 2) 0.62 11.51-28.62
10.87—28.01 18.56 + 0.66 12.16—30.24
9.61-28.11 7 SE +O265 11.26—29.87
4.08—-10.78 GO i: a eR | 9 4.78-11.41

6 BREVIORA No. 532

Females

Frequency

Snout-vent length (mm)

Figure 2. Frequency distribution of SVL of female and male Chamaeleo dilepis examined in this study. Dashed
lines represent the smallest SVL at sexual maturity. See Table 2 for X + SE of all morphological traits measured.

2012 ECOLOGY OF CHAMAELEO DILEPSIS d.

Volume (mm?) of largest egg/follicle

Testicular volume (mm7°)

8
°
2
°
°
8
8
°
°

Month

Figure 3. Egg/follicle volume (A) and testicular volume (of right testes) (B) in relation to the time of year.

important prey item, making up 25.5% of the number and prey volume indicate that al-
diet numerically and 10.6% volumetrically. though prey items made up similar compo-
After orthopterans, coleopterans were also nents of C. dilepsis’ diet numerically, prey
consumed by the most individual chameleons volume was dominated by _ orthopterans
(25.3%). Niche breadths of proportion prey (Table 3).

8 BREVIORA

TABLE 3.

No. 532

COMPOSITION OF THE DIET OF 206 C. DILEPIS FROM SOUTHERN AFRICA. INSECT PREY WERE IDENTIFIED TO ORDER.

WE EXCLUDED 59 UNIDENTIFIED PREY ITEMS WITH A TOTAL VOLUME OF 4,899.32 MM3 FROM THE SUMMARY. DIET IS REPORTED

AS THE TOTAL NUMBER OF PREY ITEMS FOUND IN STOMACHS (N) WITH THE ASSOCIATED PERCENTAGE; VOLUME WAS CALCULATED

USING THE FORMULA FOR A PROLATE SPHEROID (SEE TEXT), AND FREQUENCY REFERS TO THE NUMBER OF LIZARDS CONTAINING A
PARTICULAR PREY ITEM.

Prey Type N Jo N Volume (mm?) % Volume Frequency % Frequency
Orthoptera 180 36.44 S308) 75 68.9] Be 37.30
Coleoptera 126 S55) 7,864.30 10.56 1 25.34
Diptera 34 6.88 1,718.00 73 18 6.08
Mantodea 10 2.02 729 10 OTe W) Y) 3.04
Hymenoptera 4] 8.30 1,213;20 1.63 24 8.1]
Hemiptera DD 4.45 2,166.64 Zo 12 4.05
Phasmatodea 2) 0.40 366.71 0.49 2 0.68
Odonata 7 1.42 626.63 0.84 6 1.74
Diplopoda y) 1.82 1,744.73 2.34 9 3.04
Larvae/pupa 54 hO93 3,703.43 4.97 2h 7.09
Spiders 6 2 736.02 0.99 6 2.03
Mollusks | 0.20 14.06 0.02 I 0.34
Vertebrates 2 0.40 1,264.40 1.70 2 0.68
Totals 494 100.00 74,456.58 100.00 296 100.00
Niche breadths 4.45 2.03

When controlling for SVL, there was no
effect of sex on the number (F; 295 = 3.161,
P = 0.077) or volume (Fy. 145 = 2.00, P =
0.159) of prey items consumed, SVE and
total prey volume were significantly and
positively correlated (7? = 0.456, Fy 445 =
2.667, P = 01001. Fic. 5) bur se eandsrhe
total number of prey consumed were not
significantly related (r? = 0.092, Fy 295 =
1.242, P = 0.137). Gape size (head length x
head width) was positively correlated with
the number of prey consumed (7? = 0.114,
Fi64 = 8.115, P = 0.006) and marginally
significantly correlated with the size of the
largest prey item (7? = 0.083, Fy45 = 4.008,
P = 0.051).

DISCUSSION

Of the 14 morphological variables we
measured in C. dilepis, only SVL and pelvic
width differed significantly between the
sexes. Females were significantly larger than
males in both SVL and pelvic width. In
squamate reptiles, the degree of sexual

dimorphism depends on the relative strength
of natural and sexual selection on both sexes
(Fitch, 1981) and disentangling these effects
can be difficult. Among 21 lineages of the
viviparous dwarf chameleons (Bradypodion),
females were significantly larger in 15 (71%)
cases (Stuart-Fox, 2009). Chameleons also
tend to have relatively large clutches (max-
imum, 74 in this study) and as_ such,
fecundity selection might favor larger fe-
males relative to any selective pressures that
may be acting on male body size. Further-
more, female SVL was positively correlated
with both clutch size and total clutch
volume, suggesting that larger SVL and
pelvic width in females is an adaptation to
increased female fecundity.

Male and female reproductive cycles
did not appear to be highly synchronized.
Females appear to be seasonal, commencing
vitellogenesis in summer (November) and
containing fully developed eggs throughout
the warmer months of the year (November—
May). Males have winter and summer peaks
in testes volume and appear to commence

2012

ECOLOGY OF CHAMAELEO DILEPSIS 9

Number of prey items consumed (%)

Volume of prey items consumed (%)

on
WY

g F é 2
= E 3 £
o a e >
=

a 3

z
ge &
—- =
eT:

Figure 4. Comparison of the diet of female (dark bars) and male (clear bars) Chamaeleo dilepis. Prey number (A)
and volume (B) are presented as percentages.

spermatogenesis one to two months earlier in distribution of fully developed eggs raises
spring (September). Although we were not this possibility. Female C. dilepis invest
able to determine if individual females lay heavily in reproduction by producing large
multiple clutches in a season, the temporal clutches of small eggs. Previous reports of

10 BREVIORA

Total prey volume (mm?)

© ~w00¢6

No. 532

8 &
Ong ae

\oTe) CO
re) 088 GOO fa) re) =
re)
oP Roam 0 Soo Ph SES oo

°o

°

Snout-vent length (mm)

ictinems:

clutch sizes have been up to 50 (FitzSimons,
1943) or 77 eggs (Jacobsen, 1989), although
one individual from East Africa that was
dissected had 95 enlarged follicles in its
oviducts (Fitch, 1970). Of the 25 gravid
females we examined, the mean was 44, and
the largest clutch was 74. These are high
values for any lizard and suggest that
chameleons are typical capital breeders that
invest heavily in a single reproductive event
(see Bonnet et al, 1998). However, in the
similar sized but distantly related Trioceros
[Chamaeleo| montium from Cameroon, clutch
size 1s considerably smaller: 3-12 (mean, 6.5),
and mean relative clutch mass was 40%
(Herrmann and Herrmann, 2005). Andrews
and Karsten (2010) provide an excellent
review of the evolution of reproductive mode,
developmental rate, and body size in chame-
leons. In the context of their review, C. dilepis
is a relatively large sized, derived chame-
leon that develops comparatively slowly.

Relationship between SVL (mm) and total prey volume (mm+) in Chamaeleo dilepsis.

Furthermore, this species has embryonic
diapause because the eggs overwinter under-
ground. This reproductive strategy is thought
to have evolved in habitats experiencing dry,
seasonal climates, thereby increasing the
likelihood that the eggs hatch during favor-
able conditions the following season (An-
drews and Karsten, 2010).

The majority of prey consumed by C.
dilepis, both volumetrically and numerically,
were orthopterans, followed by coleopterans.
Prey volume was a function of body size,
with larger individuals consuming larger
prey volumes. However, SVL and _ prey
number were not significantly correlated,
possibly because most individuals contained
relatively few prey items. Gape size was
correlated with both prey number and the
largest prey item consumed. A relatively
large number of individuals contained prey
items in their stomach (82%), suggesting that
C. dilepsis feed frequently. This agrees with

2012

the patterns reported by Huey ef al. (2001)
that most diurnal lizards maintain a positive
energy balance. Nine individuals were also
found to contain a single millipede, a prey
item usually avoided by many reptiles because
of their toxicity (Wapstra and Swain, 1996).
However, the presence of noxious millipedes
has been found in the diet of two other
African species (African tree agama, Acantho-
cercus a. atricollis, Reaney and Whiting, 2002;
Wahlberg’s velvet gecko, Homopholis wahl-
bergii, Whiting et al., 2007). Together with
numerous anecdotal reports of miullipede
consumption (Branch, 1998), these studies
suggest that they are at least occasional prey
items of African lizards either because they
are less toxic than their North American
counterparts or because African lizards are
better adapted to deal with their toxicity.

In addition to insects, six chameleons had
eaten spiders, one had eaten a mollusk, and
two had eaten lizards. These constitute a
very small proportion (< 4%) of C. dilepis
specimens that contained food items. Only a
few instances of vertebrate predation by C.
dilepis have been reported in the literature,
including two geckos and a case of canni-
balism (Tilbury, 2010). Therefore, although
C. dilepis may on occasion take noninsect
prey, their diet is dominated by a relatively
narrow range of insect prey. Our results
largely confirm previous anecdotal reports.
Chamaeleo dilepis 1s reported to feed mainly
on grasshoppers and beetles (Branch, 1998;
Tilbury, 2010), but also to include occasional
butterflies, dragonflies, winged termites,
myriapods, various flying insects, and inver-
tebrates such as snails and spiders (Fitz-
Simons, 1943; Brain, 1961; Pienaar, 1978).
The similarly sized Chamaeleo namaquensis,
which is terrestrial, has a similar diet, ex-

ECOLOGY OF CHAMAELEO DILEPSIS I]

cept that tenebrionid beetles are their most
commonly consumed (and probably encoun-
tered) prey (Burrage, 1973). Three species
of rainforest chameleons (Chamaeleo) from
Cameroon were relatively opportunistic,
feeding on mainly coleopterans, heteropter-
ans, hymenopterans, dipterans, and spiders
(Hofer et al, 2003). Similarly, the common
chameleon (Chamaeleo chamaeleon) from the
Iberian Peninsula feeds on Diptera, Hyme-
noptera, Orthoptera, and Heteroptera with
little seasonal variation and no differences
between males and females (Pleguezuelos
et al., 1999).

In summary, our study provides the first
quantitative baseline ecological data for C.
dilepis, one of few species of chameleons to
occur in a savanna biome. Chamaeleo dilepis
is an insectivore that consumes mostly
orthopterans and coleopterans; they follow
a typical spring-summer reproductive cycle,
and females are larger than males and invest
heavily in egg production. Chameleons have
long been underrepresented in ecological
studies, but given the substantial investment
of females in egg production, their unique
morphology, and their wide distribution
across habitats of varying rainfall and tem-
perature, they promise to be an interesting
model group for future studies.

ACKNOWLEDGMENTS

We thank the Ditsong National Museum
of Natural History for the loan of this
material and, in particular, Wulf Haacke,
Lauretta Mahlangu, and Lemmy Mashini
for assistance. We also thank Luke Mahler
for sharing his chameleon reference database
and for general assistance and advice during
data collection.

12 BREVIORA No. 532

APPENDIX |. SPECIMENS EXAMINED FROM THE DITSONG NATIONAL MUSEUM OF NATURAL HISTORY.

Catalog No. Location (quarter degree grid square) Country
14469 2027Cc BOTSWANA
14473 Gabarone BOTSWANA
30655 1922Ca BOTSWANA
30656 1922Ca BOTSWANA
30657 1922Ca BOTSWANA
30863 1921Ca BOTSWANA
31099 1821Bd BOTSWANA
45699 1923Cb BOTSWANA
14470 1923Cd BOTSWANA
1447] 1923Cd BOTSWANA
14472 Makgadikagi BOTSWANA
26965 Kube pan BOTSWANA
30633 Nokaneng BOTSWANA
30658 1922Ca BOTSWANA
30659 1922Ca BOTSWANA
31094 1821Ba BOTSWANA
41463 Botswana BOTSWANA
2471 2355 59:-32308 MOZAMBIQUE
3934 2532Ba MOZAMBIQUE
29258 2233D¢ MOZAMBIQUE
29325 2233Dc MOZAMBIQUE
29371 2433Cd MOZAMBIQUE
29372 2433Cd MOZAMBIQUE
29375 2433Cd MOZAMBIQUE
29380 2433Cd MOZAMBIQUE
29401 2233Cd MOZAMBIQUE
80934 1659S 3843E MOZAMBIQUE
80968 1703S 3844E MOZAMBIQUE
80969 1703S 3844E MOZAMBIQUE
29156 2600S 3255E MOZAMBIQUE
29314 2233Cd MOZAMBIQUE
29377 2433Cd MOZAMBIQUE
17063 1724S 1553E NAMIBIA
17065 1724S 1553E NAMIBIA
23988 2015Ab NAMIBIA
25090 1920Da NAMIBIA
31193 2311S 1629E NAMIBIA |
36359 2308S 1628E NAMIBIA
39187 1724Ad NAMIBIA
39207 1724Ad NAMIBIA
39354 1724Ad NAMIBIA
40061 1816Dd NAMIBIA
52784 1914Dd NAMIBIA
17066 1724S 1553E NAMIBIA
22618 1724Ad NAMIBIA
43710 1724Cb NAMIBIA
57411 1920Dd NAMIBIA
68751 1952S 1354E NAMIBIA
118 2506S 3027E SOUTH AFRICA

2012

Catalog No.

1263
2663
4570
7386
7387
9479
11053
11635
13926
14216
14219
24722
26062
26079
43973
45757
47430
50679
52149
92533
61772
61775
61777
61779
61780
61781
61793
61794
61799
61800
61801
61802
61804
61805
61806
61807
67621
82096
61797A
61797B
1214
1215
1216
1388
1901
1985
26053
28872
30211

ECOLOGY OF CHAMAELEO DILEPSIS

APPENDIX |. CONTINUED.

Location (quarter degree grid square)

2329CD
TVL

2328CB
2330DC
2330DC
‘PVE

2328CB

2435S 3104E

TVL
TVE
TVL
TVL
2528CA
2528CA
2428CB
222 IIe
2528CB
2929 ANC
2528CB
2526CB
2627CD
2428CC
2330AB
2230DA
2429BB
2328DB
2328CD
2431CA

DS28EA

22ICE
2627CB
2230DA
2330DB
2229DA
TVE

2430DA

2438S 3128E

2528AD
2528BA
2230DA
2330AA
2330AA
2330AA
[VE

2345S 3015E

2528CA

2506S 3027E

KNP
2330CD

Country

SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTELAPRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA

Catalog No.

33554
35669
47855
50230
61774
61776
61778
61782
61783
61784
61785
61786
61787
61789
61791
61792
61795
61796
80135
80
9
132
133
140
149
643
719
720
721
1080
1134
199
1254
1264
1702
2035
2061
2364
2486
pI \ I
Behe)
3374
Bes
4500
11056
13023
13267
13296
13563

BREVIORA

APPENDIX |. CONTINUED.

Location (quarter degree grid square)

2329DD
2330AB
2628AA
2431DD
2427BA
2427BB
2428CD
2627CA
2230AC
252.7€B
2630DD
2528BC
2329DB
2526CB
2426DC
2431CB
2528BD
2329CA
2627AD
2329Cd
2506S 3027E
2017Ad
2017Ad
2430Ba
2528Ca
TVD
2531'Ge
Barberton
Barberton
2528Ca
2627Ba
2528Ca
2527 DC
2329Cd
2528S 3058E
2528Ca
Pietersburg
Vygeboomspt
2628Aa
2550S 2813E
2530Dc
2530Dc
2528Ca
Barberton
2429Aa
2409S 3015E
2546S 2814E
Zeerust
Pietersburg

Country

SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA

No. 532

2012

Catalog No.

14366
31144
33261
34616
34617
64331
81
120
12)
134
142
148
165
670
1132
1899
1948
2047
2048
2060
2062
2365
2488
2489
2540
Say
3767
3956
3976
4279
4451
4501
4730
12509
13561
14215
14367
33824
33895
57314
64243
65728
65935
66166
66167
67273
67470
67619
13769

ECOLOGY OF CHAMAELEO DILEPSIS

APPENDIX |. CONTINUED.

Location (quarter degree grid square)

Vaal Dam
22548 2931E
2523Ca
2453S 2817E
2453S 2817E
2508S 2840E
2329Cd
2506S 3027E
2506S 3027E
2017Ad
2430Ba
2528Ca
2528Ca
2ZIoIGE
2528Ca
2329Dd
2528Ca
2527Cc
2527Ca
2329Cd
2329Cd
2446S 2826E
2628Aa
2628Aa
Leysdorp
2329Cd
2428Ad
Modderaek
2528Ca
Nylstroom
Koster
Barberton
Nylstroom
2435S 3104E
2329Cd
Tshakoma
Hope

2453S 2817E
2453S 2817E
2536S 2824E
2349S 3003E
2442S 3131E
2532S 3059E
2306S 2902E
2306S 2902E
2438S 3128E
2502S 3113E
2438S 3128E
Mahaba

Country

SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH. AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTEH-APRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA

16

Catalog No.

L307.
13778
14075
37995
45715
45733
46026
47941
48272
50728
51689
51706
55148
56484
65231
65260
66577
66578
66579
66581
67271
68048
69051
79356
[3779
30221
SOF
52392
54579
55154
57528
62756
62859
62866
64175
66576
66580
67185
67307
67369
67583
71881
82099
16202
16204
18555
18729
22398
34459

BREVIORA

APPENDIX 1. CONTINUED.

Location (quarter degree grid square)

Mangusi
Mangusi
28128 3228E
Sodwana
2732Ba
2732Bc
2132BC
2732Bd
2152 Be
Pif[2N\@)0)
2930Cb
3030Bc
3058S 3017E
2930Dd
3043S 3010E
2850S 2959E
2832Ad
2832Ad
2832Ad
2832Ad
2832Ad
28265 32312
2832Ad
3030Cd

Illovo Beach
3030Be
2831Cd
WSE52S. 32226
2930S 3014E
3030Cb
2832Ba
2832Ad
2832Ad
2823S 3225E
2832Ad
2832Ad
2832Ab
2904S 2904E
2847S 3205E
282713532 268
2743S 2955E
2832Dc
2032Bec
2032Bc
2032Ab

1832Bd

Country

SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
SOUTH AFRICA
ZIMBABWE

ZIMBABWE

ZIMBABWE

ZIMBABWE

ZIMBABWE

ZIMBABWE

No. 532

2012 ECOLOGY OF CHAMAELEO DILEPSIS 17

APPENDIX |. CONTINUED.

Catalog No. Location (quarter degree grid square) Country
SLE2 Kariba area ZIMBABWE
83516 Mbalabala ZIMBABWE
18558 2032Bc ZIMBABWE
18559 2032Bc ZIMBABWE
21581 1730Cb ZIMBABWE
45962 1826Dd ZIMBABWE

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a
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