Diet of the Invasive Lizard Chamaeleo jacksonii(Squamata: Chamaeleonidae) at a Wet-Forest Site in Hawai‘i1
Fred Kraus2,3 and David Preston2
Abstract: We assessed diet from a sample of Jackson’s chameleons inhabiting
wet forest in Volcano Village, Hawai‘i Island, Hawai‘i. Most lizards contained
food in their gastrointestinal tract, diet was broad, and large numbers of invertebrates, mostly insects, were consumed. Native insects composed the largest
component of dietary items, and these spanned most of the body-size range
found in the native invertebrate community. Taxonomic composition of diets
does not conform to expectations for iguanian lizards inasmuch as hymenopterans form only a small portion of consumed items. Insect chemical or stinging
defenses did not preclude consumption by lizards. The study site is a mixture ofnative and alien vegetation. Chameleons collected from alien plants containedlarger numbers of native prey items than did those lizards removed from nativeplants, suggesting that lizard residency on non-native plants will not reduce theirpredation impact on native invertebrates in mixed forest. Our results confirm thefinding of a similar study in dry-forest habitat on Maui that Jackson’s chameleons are likely to pose a substantial threat to a broad range of native invertebratebiodiversity in Hawai‘i.
Chamaeleo jacksonii Boulenger, 1896, is amedium-sized (snout – vent length up to ∼160mm) lizard native to humid and wet forests of
Kenya and Tanzania. It was introduced into
Hawai‘i via the pet trade in the early 1970s
and, with considerable assistance from chameleon fanciers, has spread widely throughout the state in wet, mesic, and dry forests.
Two different modelling approaches haveshown that this species is physiologically capable of colonizing most nonalpine habitats
within Hawai‘i (Rodder et al. 2011; Kraus
et al. 2012); hence, it seems poised to invade
most native habitats in the state. These chameleons also occur at high population densities and occupy a novel predatory niche, so

1
Financial support provided by the Hawai‘i Invasive
Species Council. Manuscript accepted 31 October 2011.
2
Bishop Museum, 1525 Bernice Street, Honolulu,
Hawai‘i 96817.
3
Corresponding author (fkraus@hawaii.edu).

Pacific Science (2012), vol. 66, no. 3:397 – 404
doi:10.2984/66.3.10© 2012 by University of Hawai‘i Press
All rights reserved

they seem capable of posing a severe conservation threat to many native arthropods(Kraus et al. 2012). An earlier study of animals
from native dry forest on Maui demonstrated
that C. jacksonii feeds on a wide diversity of
invertebrates, with endemic Hawaiian invertebrates composing 35% of all prey items(Kraus et al. 2012). It has also been shown onO‘ahu that these chameleons consume somespecies of endangered land snails as well as
native insects (Holland et al. 2010). The impression gained from that earlier dietary information is that these chameleons are generalist opportunistic predators that will consume
any small animal that comes within reach andattracts their notice. So as to further explore
the dietary range of this species we conducteda follow-up study of the diet of C. jacksonii
under different habitat conditions: a largely
native wet forest on Hawai‘i Island. The intent of this study was threefold: to further
assess how catholic the diet of this lizard is,
to determine whether similarly large numbersof native invertebrates are consumed acrossthese differing (wet- versus dry-forest) habitats, and to investigate whether more native
prey are consumed by lizards found occupying native versus alien plants.

397

398

PACIFIC SCIENCE · July 2012

materials and methods
We obtained a sample of 34 Chamaeleo jacksonii from Volcano Village and immediate vicinity, 1,100 – 1,200 m elevation, Hawai‘i Island. Habitat in this area comprises a mix oflargely native overstory species with several
invasive trees (primarily Morella faya) andunderstory shrubs (primarily Tibouchina urvilleana). We collected most lizards (n = 28) atnight while they were sleeping on vegetation, but one was collected in daytime while
active. For all these animals the plant species
on which they were perched was recorded so
as to ascertain whether animals perched onnative vegetation were more likely to havenative prey items in their digestive tracts.
Five additional animals were collected byothers living in or near Volcano and given tous; some of these were obtained while they
were crossing roads. Consequently, there isno perch information for any of those animals.
We fixed animals in formalin or froze
them within a few hours of collection to terminate digestion; we later fixed frozen animals in formalin. We then stored all animals
in 70% ethanol at Bishop Museum ( bpbm
21741, 21743, 35305 – 35327, 35409 – 35415,
35418 – 35419) and measured snout – ventlength (SV ) of all lizards. We removed digestive tracts from specimens, opened them, andremoved their contents. Using a binoculardissecting scope, we then sorted these contents by taxon, identified them to the lowest
taxonomic level possible, and counted their
numbers. We determined identifications of
dietary items (whole specimens [when present] or diagnostic fragments) by comparison
with authoritatively identified specimens in
the Bishop Museum entomology collections.
The bulk of removed contents comprised partially digested and disarticulated arthropods
mixed with unidentifiable debris. Because asingle individual prey item was partially or
wholly disarticulated during consumption, we
counted these arthropod parts separately anddetermined the total number of prey items
per taxon by choosing the identifiable partwith the highest count. For example, if there
were 20 pairs of elytra from species A but only

five legs, the total count of prey items for species A was 20.
For dietary items that we could identify to
species or to a set of very similar species, weestimated prey-volume by (1) measuring fromthe entomology collection at the Bishop Museum the maximum body length, width, andheight (to the nearest 0.05 mm) of 10 specimens of the consumed species; (2) averaging
these sample body measurements; (3) using
these averages to calculate an average volume
for each prey species using the equation fora scalene ellipsoid ( V = 4/3πabc, where a, b,
and c are the three body axes); and (4) multiplying those species-averaged volumes by therelevant number of dietary items identified
within each stomach to estimate total foodvolume for each lizard. For dietary items that
we could not identify to species, we measured
the dimensions of the consumed items directly. The benefit of measuring whole specimensof identified species from the entomology
collection is that it provides a more accurate
measure of prey volume than estimating thesame from partially digested fragments ofprey. Of 795 dietary items, all could be identified to family, 703 to genus, and 344 to species; only four were so fragmented by digestion that we could not identify them or
estimate their volume. Taxonomy and provenance for all identified arthropod dietary
items followed Nishida (2002). We labeled,stored in vials with 95% ethanol, and maintained in the collections of the Bishop Museum all recovered and sorted material. We
assessed statistical differences with G tests
and Mann-Whitney U tests implemented in
Minitab 14. We used means for comparisonsof prey numbers, and we used medians forcomparisons of prey volume because of theadditional order of magnitude observed in therange of those numbers. We assessed prey diversity using the Shannon-Wiener index.
results
We retrieved from our sample of lizards
a total of 795 dietary items from 48 taxa.
We identified 26 of these dietary taxa to species, 17 to genus, and five to family. This
comprised 43.3% of dietary items identified

Diet of Chamaeleo jacksonii · Kraus and Preston

to species, 45.2% to genus, and 11.6% tofamily.
All chameleons but one had food items intheir digestive tract (97.1%); all but four fooditems were arthropods. Nonarthropods included two land snails (Oxychilus alliarius) andtwo lizards (Lampropholis delicata). Numbersof food items per chameleon ranged from 0
to 77 (mean = 23.4, SD = 3.18). For stomachs
containing food, numbers of prey species per
chameleon ranged from 1 to 12 (mean = 5.6,
SD = 0.45), and prey volumes per lizard
ranged from 0.009 to 5.10 ml (mean = 0.76
ml, SD = 0.17). For stomachs containing
food, numbers of prey did not differ between
sexes (Mann-Whitney U = 168.5, nF = 11,nM = 21, P = .62), nor did prey volume (MannWhitney U = 163.0, nF = 11, nM = 21, P = .47).
Median numbers of prey for males were 22
and median numbers for females 16; medianvolumes for males were 0.38 ml and for females 0.45 ml. The largest prey items (Elimaea punctifera) were 48 mm long; however,most prey were of small size (Figure 1). Male399

and female lizards did not differ significantly
in body size (male median = 98 mm, medianfemale = 105 mm, Mann-Whitney U = 191.5,
nF = 11, nM = 21, P = .71).
Endemic Hawaiian taxa composed 46.9%
of dietary items, adventive aliens composed
31.7%, intentionally introduced aliens composed 9.8%, and taxa whose provenance could
not clearly be identified composed 11.6%.
For animals with food in their stomach, numbers of prey per lizard varied from 1 to 77(mean = 24.1, SD = 3.20), and volume varied
from 0.009 to 5.104 ml (mean = 0.758 ml,
SD = 0.169). Numbers of endemic prey items
per lizard varied from 0 to 58 (mean = 11.3,
SD = 1.99), and volumes of endemic prey
items per lizard varied from 0 to 0.405 ml(mean = 0.094 ml, SD = 0.019). Numbers ofendemic prey did not differ between sexes(Mann-Whitney U = 186.0, nF = 11, nM = 21,
P = .87), nor did volumes of endemic prey
(Mann-Whitney U = 198.5, nF = 11, nM = 21,
P = .51): median numbers of endemic preyfor males were nine and median numbers for

Figure 1. Size distribution of prey retrieved from 33 Chamaeleo jacksonii on Maui. n = 795 measurable prey items.

400

females eight; median volumes for males were0.053 ml and for females 0.082 ml. The
Shannon-Wiener index for identified prey
items was 3.01, suggesting a considerable degree of prey diversity among food items analyzed at the lower taxonomic levels (species orgenus) at which most of our prey items wereidentified. Food items per lizard that could
not have their provenance (endemic versus
alien) determined because of identification
uncertainty varied from 0 to 18. These numbers were sufficiently small that overall assessment of lizard impact on endemic prey is unlikely.
Analyzed taxonomically, homopteranscomposed the greatest numbers of prey, followed by dipterans and coleopterans (Table1). Together, these three taxa formed 74.8%
of all dietary items. The large majority ofendemic insects consumed at Volcano Village
were homopterans of the genera Oliarus andNesophrosyne, composing 27.9% of all prey
items, but several other native species acrossa diversity of orders were taken as well
(Table 2).
When analyzed by volume, however, theresults are somewhat different. In that case,dipterans and coleopterans were still important (35.2% and 17.7% of prey volume, respectively), but hymenopterans and orthopterans composed significant volumes as well
(16.9% and 13.7% of prey volume, respectively). Adventive aliens composed 59.4% offood items by volume, endemic species composed 12.5%, intentionally introduced aliens
composed 13.0%, and taxa whose provenancecould not clearly be identified composed
15.1%.
The one lizard without food items in its digestive tract was collected in daytime from alime tree (Citrus aurantifolia). Of the remaining 27 specimens for which perch species was
identified, 21 were collected on native plant
species (12 on Metrosideros polymorpha, 4 on
Acacia koa, 2 on Coprosma ochracea, 2 on Dicranopteris linearis, 1 on Vaccinium calycinum)and 6 were found in alien species (4 on
Tibouchina urvilleana, 2 on Morella faya). Unexpectedly, lizards found on alien plants had ahigher percentage of endemic prey items intheir digestive tract than did animals collected

PACIFIC SCIENCE · July 2012

on native plants (57.6% versus 45.5%, G =4.4877, df = 1, P = .034).
discussion
Few studies have been done on chameleon diets, with much of this information involving
alien Chamaeleo chamaeleon in Spain (Blasco
et al. 1985, Pleguezuelos et al. 1999) andMalta (Luiselli and Rugiero 1996) and C. africanus in Greece (Dimaki et al. 2001). Relatively few species have been studied in theirnative range (Burrage 1973, Wild 1994, Hofer
et al. 2003, Keren-Rotem et al. 2006). Noneof these studies identified dietary items below
the level of insect order. A prior study of C.
jacksonii from a native dry forest on southern
Maui (Kraus et al. 2012), however, identified
most dietary items to the level of species orgenus, and several interesting features about
the diets of these lizards were discovered.
First, these dry-forest chameleons consumed
a large number of small prey items, with amean of 41.8 dietary items per stomach and arange of 0 – 352. Second, dietary breadth waswide, with a Shannon-Wiener index of diversity at the genus/species level of 2.56. Dipterans, heteropterans, and homopterans composed the predominant prey items. Third,
ants, other hymenopterans, and beetles composed small proportions of the chameleons’diets; this is unusual for iguanian lizards,
which are largely ambush predators reliant on
visual cues for prey detection and prey preferentially on those taxa. It was, however, consistent with other evidence suggesting that,among iguanians, chameleons are unusual inbeing cruise foragers (Hagey et al. 2010).
Should this foraging strategy prove applicable
to chameleons as a whole, it would suggestthat chameleon diets may have shifted as theirforaging strategy changed. Directly relevant
to conservation concerns are that (1) a large
number of native insects (35.2% of total) was
consumed, and (2) cropping rates seem likely
to be high because of the large numbers of
prey items in the lizards’ digestive tract andbecause chameleons have been reported totake 3 – 15 meals per day and to have high digestion rates that quickly clear their digestive
tracts (Burrage 1973).

TABLE 1
Composition of Dietary Items in Gastrointestinal Tract of 33 Chamaeleo jacksonii from Volcano Village,
Hawai‘i Island
Order
Squamata
AraneaeBlattodea
Coleoptera

Dermaptera
DipteraHeteroptera

HomopteraHymenoptera

Lepidoptera

Orthoptera
JulidaPolydesmidae
IsopodaGastropoda
Total

FamilyScincidae
Tetragnathidae
Thomicidae
Blattelidae
Cerambicidae
Chrysomelidae
Coccinellidae
Curculionidae
Elateridae
Forficulidae
Calliphoridae
Drosophilidae
Stratiomyidae
Syrphidae
Lygaeidae
MiridaeNabidaePentatomidae
Cicadellidae
CixiidaePsylidae
ApidaeIchneumonidae
Sphaecidae
VespidaeCrambidae
Cosmopterygidae
Geometridae
Gryllidae
Tettigoniidae
JulidaeParadoxosomatidae
Porcellionidae
Zonitidae

No. Prey Items% Prey Items% Prey Volume

2
2
11
9
2
3
3
128
3
1
38
71
15
1
1
185
99
1
37
48
35
2
14
17
2
182
103
130
49
32
2
5
1
24
38
8
13
17
29
25
4
40
40
2
2
5
5
2
2
795

0.25
0.25
1.38
1.13
0.25
0.38
0.38
16.10
0.38
0.13
4.78
8.93
1.89
0.13
0.13
23.27
12.45
0.13
4.65
6.04
4.40
0.25
1.76
2.14
0.25
35.47
12.96
16.35
6.16
4.03
0.25
0.63
0.13
3.02
4.79
1.01
1.64
2.14
3.64
3.14
0.50
5.03
5.03
0.25
0.25
0.63
0.63
0.25
0.25
100.00

3.00
3.00
0.52
0.44
0.08
0.87
0.87
17.66
0.81
0.00
5.72
8.00
3.13
0.03
0.03
35.24
26.86
0.00
5.49
2.89
3.15
0.01
0.24
0.36
2.53
4.02
1.81
1.70
0.51
16.94
1.41
0.28
0.40
14.85
3.96
0.92
0.27
2.76
13.74
2.04
11.70
0.27
0.27
0.17
0.17
0.38
0.38
0.04
0.04
100.00

402

PACIFIC SCIENCE · July 2012
TABLE 2Endemic Hawaiian Insects Composing Dietary Items in the Sample of 33 Chamaeleo jacksonii from Wet-Forest
Habitat on Hawai‘i Island Having Gastrointestinal Contents
Order
Araneae
Araneae
AraneaeColeoptera
Coleoptera
DipteraHeteroptera
Heteroptera
Heteroptera
Heteroptera
Heteroptera
Heteroptera
Heteroptera
Homoptera
Homoptera
HomopteraHymenoptera
Lepidoptera
Lepidoptera
Orthoptera

FamilySpecies

No. Prey Items% Prey Items% Prey Volume

Tetragnathidae
Tetragnathidae
Thomicidae
Cerambicidae
Cerambicidae
Drosophilidae
Lygaeidae
MiridaeMiridaeNabidaeNabidaeNabidaePentatomidae
Cicadellidae
CixiidaePsylidae
Ichneumonidae
Cosmopterygidae
Geometridae
Gryllidae

Tetragnatha sp. 1
Tetragnatha sp. 2
Mecaphesa sp.
Plagithmysus sp. 1
Plagithmysus sp. 2
Drosophila sp.
Nysius sp.
Sarona sp.
Hyalopeplus pellucidus
Nabis blackburni
Nabis lusciosus
Nabis sp. nr. blackburni
Coleoticus blackburniae
Nesophrosyne sp.
Oliarus sp.
Megatrioza sp
Enicospilus sp.
Hyposmocoma sp.
Eupithecia sp.
Laupala sp.

6
3
2
2
1
1
2
6
5
6
1
10
2
92
130
49
1
13
16
25

0.75
0.38
0.25
0.25
0.13
0.13
0.25
0.75
0.63
0.75
0.13
1.26
0.25
11.57
16.35
6.16
0.13
1.64
2.01
3.14

0.26
0.18
0.08
0.72
0.09
0.00
0.01
0.06
0.18
0.13
0.04
0.20
2.53
1.60
1.70
0.51
0.10
0.27
1.82
2.04

This study of Chamaeleo jacksonii, from adifferent form of native vegetation in Hawai‘i(wet versus dry forest), supports several ofthese findings and expands on others. Chameleons investigated in this study contained
fewer food items on average than did animals
from the earlier dry-forest study (23.4 versus41.8), and the numerical range of items (0 – 77versus 0 – 352) and standard deviation of samples (3.18 versus 10.41) were both lower, but
numbers of consumed items were still large.
Size distribution of prey items, however, wasvirtually identical with that in the earlier study
from dry-forest habitat, with (1) the large majority of dietary items being less than 9 mm inlength (Figure 1), and (2) larger prey items
being consumed at lower frequencies. Preydiversity was even higher in this study than inthat on Maui (Shannon-Wiener index 3.01
versus 2.56), perhaps reflecting the greater
structural and community diversity of vegetation at the wet-forest study site or its closer
proximity to human activity, either of which
could be expected to result in a larger community of invertebrate species. It is not explained by larger sizes of the Volcano lizards,

which do not differ significantly from thosecollected earlier on Maui (for males, MannWhitney U = 392.0, nVolcano = 21, nMaui = 21,
P = .18; for females, Mann-Whitney U =392.0, nVolcano = 11, nMaui = 12, P = .81).
The vast majority of prey in this study werearthropods, but land snails and lizards werealso consumed. The latter confirms the observation of Holland et al. (2010), although
the snails obtained in our study are alien introductions. Both the lizard (Lampropholis
delicata) and snails (Oxycheilus alliarius) are ofinterest because they are leaf-litter inhabitants and must have been consumed while thechameleons were active on the ground. Although chameleons are primarily arboreal,
they are often seen crossing roads in Hawai‘i,
and these prey items confirm their willingness to feed while crossing between patchesof arboreal habitat. As found in our earlier
study, homopterans, dipterans, and coleopterans were of greatest dietary importance,with percentages of prey items consumed of
the first and last virtually the same between
study sites (homopterans: 34.64% on Maui,35.47% on Hawai‘i Island; coleopterans:

Diet of Chamaeleo jacksonii · Kraus and Preston

14.57% on Maui, 16.10% on Hawai‘i Island)
but with far fewer flies consumed in this study(42.88% on Maui, 23.27% on Hawai‘i Island).
We presume that the reduced consumption of
dipterans on Hawai‘i Island reflects the lesser
relative abundance of flies in this wet forest.
Also, as found earlier, hymenopteransformed a very small portion of the diet in bothstudies (1.69% of prey items on Maui, 4.03%
on Hawai‘i Island), in contrast to the usual
expectations for iguanian lizards ( Vitt et al.
2003, Vitt and Pianka 2005). In this instance,
we found no ants at all in the diet, although
two honey bees (Apis mellifera), one sphecid(Ampulex compressa), five ichneumonid wasps(4 Barichneumon, 1 Enichospilus), and 24 yellow
jackets ( Vespula pensylvanica) were present.
Beetles also compose a large percentage of thediet of most iguanians ( Vitt et al. 2003, Vittand Pianka 2005) and, as already noted, composed a fair portion of chameleon diets inboth dry forest on Maui and wet forest on
Hawai‘i Island, although in neither case didthey approach constituting a majority of fooditems.
A larger percentage of prey in the diets ofVolcano chameleons consisted of native species than was found in dry forest on Maui(46.9% versus 35.2%). The unexpected resultthat chameleons taken from native plants had
fewer native prey species in their digestive
tracts could reflect that (1) chameleons movefrequently among perches, such that their
final resting places for the night do not necessary reflect where they last foraged; (2) thenative arthropods ingested are not particularly restricted to native plants; or (3) native insects are more vulnerable to predation whenon alien plants. Given what we presume to be
rapid digestion rates for these chameleons
and their generally sedentary nature, we are
inclined to view the first hypothesis as lesslikely than the other two.
What impact might be expected fromChamaeleo jacksonii on Hawai‘i’s native invertebrates? Three points are directly relevantto that question. First, as we discussed in ourearlier study, these lizards clearly tolerate adiversity of prey having noxious defenses.
In both studies, insects armed with painful
stings (Apis mellifera, Vespula pensylvanica) or

403

bites (Pheidole megacephala) were eaten in sufficient numbers that their consumption cannot readily be ascribed to accidental ingestion. Similarly, a large number of prey items
belonged to species having noxious chemical
defenses: 36 Nezara viridula and 4 Danaus
plexippus in the Maui study, 40 Cylindroiulus
latestriatus in this study. Second, active motion appears to be unnecessary to elicit feeding in these chameleons, as suggested by theirconsumption of land snails (this study, Holland et al. 2010), which are largely inactive
during daytime, and their consumption of
bird eggs in captivity (cf. Kraus et al. 2012).
Third, on both Hawai‘i Island and Maui wefound these lizards to consume species spanning virtually the entire range of body sizes
found in native arthropods in Hawai‘i. Thus,
it appears that few, if any, native Hawaiian
invertebrates will prove invulnerable to predation by these lizards. Coupled with thechameleons’ high population densities andexpected wide geographical range across
Hawai‘i (Kraus et al. 2012), the conservation
implications of this conclusion for the largest
component of Hawai‘i’s native biodiversity
should be obvious.
acknowledgments
We thank Meg Barnaby, David Foote, Dennis
LaPointe, Will Pitt, Thane and Linda Pratt,
Ezra Teodoro, and Rick Warshauer for theirgenerous assistance in collecting chameleons;Shar Hashimoto and Terri Lopez for specimen preparation; Pumehana Imada for curatorial services; Tracie Mackenzie for administrative assistance; and the Hawaii Invasive
Species Council for financial support. This iscontribution no. 2012-0003 from the Hawai‘i
Biological Survey at the Bishop Museum.
Literature Cited
Blasco, M., J. Romero, J. M. Sánchez, andE. Crespillo. 1985. La biología alimentaria
y reproductiva del camaleón común,
Chamaeleo chamaeleon (L.) (Reptilia,
Chamaeleonidae) de la Península Ibérica ynorte de Africa. Monogr. Inst. Nac. Conserv. Nat. 43:121 – 148.

404

Burrage, B. R. 1973. Comparative ecology
and behaviour of Chamaeleo pumilus pumilus (Gmelin) and C. namaquensis A. Smith(Sauria: Chamaeleonidae). Ann. S. Afr.
Mus. 61:1 – 158.
Dimaki, M., E. D. Valakos, A. Legakis, B.
Chondropoulos, and A. Bonetti. 2001.
Preliminary results on the feeding ecologyof the African chameleon Chamaeleo africanus Laurenti, 1768 from the southwestern
Peloponnese, Greece. Pages 57 – 63 in P.
Lymberakis, E. Valakos, P. Pafilis, and M.
Mylonas, eds. Herpetologia Candiana. Societas Europaea Herpetologica, Irakleio,
Greece.
Hagey, T. J., J. B. Losos, and L. J. Harmon.
2010. Cruise foraging of invasive chameleon (Chamaeleo jacksonii xantholophus) in
Hawai‘i. Breviora 519:1 – 7.
Hofer, U., H. Baur, and L.-F. Bersier. 2003.
Ecology of three sympatric species of thegenus Chamaeleo in a tropical upland forest
in Cameroon. J. Herpetol. 37:203 – 207.
Holland, B. S., S. L. Montgomery, and V.
Costello. 2010. A reptilian smoking gun:First record of invasive Jackson’s chameleon (Chamaeleo jacksonii) predation on native Hawaiian species. Biodivers. Conserv.
19:1437 – 1441.
Keren-Rotem, T., A. Bouskila, and E. Geffen.
2006. Ontogenetic habitat shift and risk ofcannibalism in the common chameleon(Chamaeleo chamaeleon). Behav. Ecol. Sociobiol. 59:723 – 731.
Kraus, F., A. Medeiros, D. Preston, C. S.

PACIFIC SCIENCE · July 2012

Jarnevich, and G. H. Rodda. 2012. Dietand conservation implications of an invasive chameleon, Chamaeleo jacksonii (Squamata: Chamaeleonidae), in Hawai‘i. Biol.
Invasions 14:579 – 593.
Luiselli, L., and L. Rugiero. 1996. Natural
history notes: Chamaeleo chamaeleon: diet.
Herpetol. Rev. 27:78 – 79.
Nishida, G. M., ed. 2002. Hawaiian terrestrial
arthropod checklist. 4th ed. Bishop Mus.
Tech. Rep. 22. http:// hbs.bishopmuseum.org/ hbsdb.html.
Pleguezuelos, J. M., J. C. Poveda, R. Monterrubio, and D. Ontiveros. 1999. Feeding
habits of the common chameleon, Chamaeleo chamaeleon (Linnaeus, 1758) in thesoutheastern Iberian Peninsula. Isr. J.
Zool. 45:267 – 276.
Rodder, D., W. Bohme, and S. Lotters. 2011.
‘Chamaliens’ on the Hawaiian Islands:
Spatial risk assessment for the invasive
Jackson’s chameleon (Chamaeleonidae).
Salamandra 47:36 – 42.
Vitt, L. J., and E. R. Pianka. 2005. Deep history impacts present-day ecology andbiodiversity. Proc. Natl. Acad. Sci. U.S.A.
102:7877 – 7881.
Vitt, L. J., E. R. Pianka, W. E. Cooper Jr., and
K. Schwenk. 2003. History and the global
ecology of squamate reptiles. Am. Nat.
162:44 – 60.
Wild, C. 1994. Ecology of the Western
Pygmy Chameleon Rhampholeon spectrum
Buchholz 1874 (Sauria: Chamaeleonidae).
Br. Herpetol. Soc. Bull. 49:29 – 35.