CSIRO PUBLISHING

www.publish.csiro.au/journals/ajz Australian Journal of Zoology, 2007, 55, 341–350

Diet and habitat preferences of the Fijian crested iguana

(Brachylophus vitiensis) on Yadua Taba, Fiji: implications
for conservation

Clare Morrison A,E, Tamara Osborne B, Peter S. Harlow C, Nunia Thomas A,

Pita Biciloa D and Jone Niukula D
A
Institute of Applied Sciences, University of the South Pacific, Private Mailbag, Suva, Fiji.
B
Biology Division, University of the South Pacific, Private Mailbag, Suva, Fiji.
C
c/o Herpetofauna Division, Taronga Zoo, PO Box 20, Mosman, NSW 2088, Australia.
D
National Trust of Fiji, 4 Ma’afu Street, Suva, Fiji.
E
Corresponding author. Email: Morrison_c@usp.ac.fj

Abstract. The Fijian crested iguana (Brachylophus vitiensis) is restricted to tropical dry forest habitat and has been
extirpated from over 80% of its original range primarily because of habitat destruction. A large population on Yadua Taba
island has been proposed as a source for iguana translocations. This study aimed to determine the dietary and habitat
requirements of the herbivorous B. vitiensis on Yadua Taba to identify essential tree species. Between September 2005 and
June 2006 we examined the diet of B. vitiensis using faecal analyses, while perch preferences were examined using field
surveys. Faecal analyses identified 26 plant species in the diet of B. vitiensis, while field surveys recorded iguanas in
33 tree species. The most common diet species largely overlapped with the most common perch species. There were no
major seasonal shifts, sex differences, or age-class variations in diet or perch preferences. These results suggest that while
B. vitiensis occurs in and consumes several plant species on Yadua Taba, it is primarily dependent on only a few species.
To improve translocation success, future translocations of B. vitiensis need to ensure that these important tree species are
present at new sites.

Introduction
Translocations are defined as the ‘deliberate and mediated
movement of wild individuals or populations from one part of
their range to another’ (IUCN 1995), and to date have been
used as a conservation management tool for several threatened
or endangered taxa (Griffith et al. 1989; Wikramanayake 1990;
Bright and Morris 1994; Komdeur 1994; Towns and Ferreira
2001). While translocations are likely to become progressively
more important in the preservation of genetically and demo-
graphically threatened populations, previous reviews of
species’ relocations and translocations (e.g. Griffith et al. 1989;
Dodd and Seigel 1991; Wolf et al. 1996; Fischer and
Lindenmayer 2000), have demonstrated that many trans-
locations are unsuccessful and expensive. This has resulted in
increased interest in the factors associated with successful
translocations.
While the factors associated with successful translocations
are, to a large extent, determined by the specific taxon in ques-
tion, several general factors contributing to successful trans-
locations of threatened or endangered taxa (as opposed to native
game species’ movements) have been identified: (1) trans-
locations into the core of a species historical range are more suc-
cessful than those to the periphery or outside the range,
(2) translocations are more successful when a large number of
individuals are released, (3) translocations are more successful
when the cause of the original decline is removed, (4) herbivore
translocations are more successful than those of carnivores or
omnivores, and (5) translocations into areas of high habitat
quality are more successful than those to areas of lower quality
(Griffith et al. 1989; Bright and Morris 1994; Fischer and
Lindenmayer 2000).
The endemic Fijian crested iguana (Brachylophus vitiensis)
is a herbivorous lizard restricted to tropical dry forest habitats
(one of the most threatened vegetation types in the Pacific) on
islands in western Fiji (Harlow et al. 2007). In the past 20 years
it has been extirpated from almost 80% of its original docu-
mented range primarily due to extensive destruction of its forest
habitat (through fire and goat grazing), and also by feral cat pre-
dation. Surveys in 2000, 2001 and 2003 have confirmed the
presence of the species on only six islands (Harlow et al. 2007).
Yadua Taba is the stronghold for the species, supporting ~98%
of all individuals (estimated to be >10000 animals), and is the
only legally protected population in Fiji (Yadua Taba is a
National Trust of Fiji reserve).
While the population of B. vitiensis on Yadua Taba is cur-
rently secure, there are obvious risks in having only a single
secure population. The accidental introduction of exotic preda-
tors, disease, wildfire, or the threat of invasive alien plant
species are potential threats, capable of decimating this popu-
lation. As such, translocations of iguanas from Yadua Taba to
other islands within the former range of B. vitiensis have been
proposed as a viable conservation-management option for the
species (IUCN/SSC Iguana Specialist Group 2007).

© CSIRO 2007 10.1071/ZO07062 0004-959X/07/060341

342 Australian Journal of Zoology
The successful translocation of B. vitiensis requires, among
other things, selecting sites within the former range of the
species with suitable habitat and the absence of threatening pro-
cesses, including predators (e.g. cats, dogs, mongoose) and
habitat-clearing activities (e.g. grazing by goats, fires).
Translocating iguanas to these islands will involve ensuring that
suitable plant species are currently present or reintroduced to
these islands before translocations. While there is some docu-
mented evidence of the plant requirements of B. vitiensis
(Gibbons and Watkins 1982; Gibbons 1984; Laurie et al. 1987;
Harlow and Biciloa 2001; Harlow et al. 2007), no data are avail-
able on seasonal shifts, sex differences, nor age-class variation
in diet or perch preferences.
The overall aim of our study was to determine the dietary and
shelter/perch requirements of B. vitiensis on Yadua Taba with a
view to improving the effectiveness of management and conser-
vation of the species throughout the Fiji Islands. Specifically,
our objectives were to (1) identify plant species in the diet of
B. vitiensis on Yadua Taba, including seasonal shifts and dif-
ferences between age–sex classes if present; (2) identify perch
plant species on Yadua Taba, including seasonal shifts and dif-
ferences between age–sex classes if present; and (3) based on
the results from (1) and (2), compile a list of plant species essen-
tial for any potential translocation sites.
Materials and methods
Study species
Brachylophus vitiensis (Gibbons 1981) is endemic to Fiji, is cur-
rently listed as Critically Endangered under IUCN (2006) crite-
ria, and is the only Fijian reptile listed as Endangered in the Fiji
National Biodiversity Strategy and Action Plan (NBSAP 1998).
It is a moderately sized iguana (adults 185–236 mm SVL,
hatchlings 83–88 mm SVL), and adult sex is easily distin-
guished by the presence of enlarged femoral pores in males. The
iguana is a diurnal, strongly arboreal, herbivorous species
(Morrison 2003). Males are territorial and often bear scars or
mutilations from aggressive encounters. Reproductive cycles
are poorly known; however, the mating season on Yadua Taba
appears to run from December to March and gravid females are
seen from January to April (S. Morrison, unpubl. data). Females
dig burrows on the forest floor, where 3–5 eggs are laid before
the burrow is back-filled. Hatchlings emerge roughly 8–9
months later (October–November) at the onset of the wet season
(S. Morrison, unpubl. data).
Site
Yadua Taba is a small (72 ha), volcanic island uninhabited by
people, located in the north-west of the Fiji archipelago
(16°50′S, 178°20′E). It reaches over 100 m in height and lies
120 m off the much larger island of Yadua (1360 ha, which is
inhabited by people). Four distinct vegetation types have been
recorded previously on the island: coastal strand (beach/
littoral), rocky cliff, dry forest, and grass/shrubland (Laurie
et al. 1987; Olson et al. 2002). Goats were introduced to Yadua
Taba in 1972, and numbered over 200 by the late 1970s
(Gibbons 1984). The tropical dry forest vegetation on the island
during this time was severely depleted by a combination of
grazing and fire used to drive goats for ease of capture. Since the
C. Morrison et al.
island was declared a sanctuary in 1981 all goats have been
removed, and fires banned. Today the forest on the island has
recovered to a remarkable extent and is currently one of the best
remaining examples of tropical dry forest and coastal strand
vegetation in the Pacific (Olson et al. 2002).
Survey habitat and dates
Tropical dry forests (TDFs) are found in seasonal dry, tropical
regions where the average annual rainfall is less than 2000 mm.
These regions also experience dry seasons that span three or
more months with monthly rainfall of less than 100 mm. The
TDF flora has multiple adaptations to dry conditions, including
drought avoidance and resistance through a variety of morpho-
logical and ecological characteristics.
On Yadua Taba the dry season generally lasts from May to
October. Air temperature ranges from a minimum of 16°C in
July–August to a maximum of 31°C in January–February
(Gibbons 1984). We carried out four 2-week-long surveys on
Yadua Taba during 12–23 September 2005 (referred to herein as
September), 28 November–9 December 2005 (December),
20 February–3 March 2006 (February) and 29 May–9 June
(May). These survey times were chosen to mirror seasonal
changes in weather and plant phenology.
September represents the mid-late dry season when there are
few flowering and fruiting species and few trees with new
leaves. December is the early wet season when many of the trees
are flowering and fruiting, while February is the mid wet season
and is characterised by heavy fruiting of most species, and occa-
sional flowering. June is the early dry season and is charac-
terised by limited late fruiting of some species and trees with
new leaves.
We established six 250-m permanent transects in September
(two each in TDF, coastal forest and Casuarina forest) with each
transect being surveyed once on each of the four trips. The tran-
sects were set up in different vegetation types to maximise the
number of plant species available for the iguanas to utilise. We
randomised the order in which they were surveyed on each trip.
Data collection
Iguana surveys
Each night four searchers with strong torches used line-
transect survey techniques to record all iguanas along a 250-m
transect line to a distance of 10 m either side of the line. We
recorded the tree species in which each iguana was found, the
iguana’s sex, age class, and height above ground. We were able
to sex most adult iguanas that were perched less than 15 m from
the ground. Those that could not be sexed were recorded as
adults. All animals less than 160 mm SVL without defined
femoral pores were classified as juveniles. We surveyed one
transect per night, and surveys took 2–5 h depending on the
number of iguanas found.
Due to their dentition, herbivorous iguanid lizards tend to
bite and tear off pieces of food rather than chew them, resulting
in many leaves, flowers and fruit seeds passing relatively undi-
gested through the gut and being identifiable to species
(Throckmorton 1976; Laurie et al. 1987). We sampled iguana
diet by analysing scats collected from several randomly cap-
tured iguanas on each transect (Table 1).
Diet and habitat of Fijian crested iguanas Australian Journal of Zoology 343

Iguanas were captured by collecting off the tree (individuals
less than 2 m above ground), climbing the tree (2–4 m) or using
a modified extendable fishing pole (>4 m). The fishing pole had
a small piece of wood, ~30 × 6 cm attached to the end. This was
used to lightly prod the sleeping iguanas from their perch. The
iguana would then clasp onto the wooden end of the fishing pole,
and could easily be brought down to the ground and captured by
hand. Captured iguanas were usually less than 8 m above ground,
with the mean height of captured iguanas significantly lower
than other iguanas that were sighted but not collected (Table 1).
The location of each captured iguana along transects was
marked by tying a labelled tag to the transect string at the point
of capture. Each captured lizard was kept in a clean, uniquely
numbered cloth bag in a cool, shady location for 1–3 days
(average of 2.1 days), after which it was returned to its point of
capture. Most lizards produced one large scat or occasionally
several small (usually two) scats per day in these bags. The scats
for each lizard were stored in 75% alcohol in individually
labelled ‘click seal’ or ‘Zip lock’ plastic bags and taken back to
the laboratory.
Plant surveys
In total, 147 plant species have been recorded from Yadua
Taba, including trees, shrubs, ferns, vines and grasses (Keppel
and Tuiwawa 2007). In September 2005 we recorded the number
of all tree and shrub species 5 m either side of the central tran-
sect line along all six transects to obtain a measure of the density
of each plant species on the transect. We also recorded the
phenology (e.g. flowering, fruiting, etc.) of all tree species along
the six transects on each subsequent survey to identify seasonal
variation in the availability of food sources for the iguanas.
Identification of dietary items
In the laboratory we gently washed iguana scats under flowing
water through a sieve to isolate plant material. Each sample was
sieved twice through successively finer sieves to ensure that all
of the plant material in the scat was collected for identification.
The washed plant material was stored in 75% alcohol until iden-
tified. We identified leaves, seeds, flowers and buds under a
low-power dissecting microscope by comparing them to refer-
ence samples of plant species from Yadua Taba.
Statistical analyses
We used two-factor ANOVAs to examine (1) plant species utili-
sation and seasonal variation in iguana diet, and (2) plant
species utilisation and seasonal variation in tree species used as
night perches. We used only the six most numerous plant species
in the diet analysis as there were insufficient data to make sea-
sonal comparisons for the remaining species. Only the 10 most
commonly used plant species were included in the perch
analysis for the same reason. A Student’s t-test was used to
compare the perch heights of (1) adult and juvenile iguanas and
(2) male and female iguanas.
We used Chi-square analyses to test for age class and sex dif-
ferences in diet and perch tree species. Only the top eight plant
species (frequency of occurrence >2%) found in the scats, and
the eleven most common perch plants (frequency of use >2%)
were used in these analyses. A Student’s t-test was used to
compare the number of plant species found in the scats in indi-
vidual adult and juvenile iguanas.
We used simple linear regressions to examine the relation-
ship between abundance of common plant species and utilisa-
tion by iguanas for both diet and perches. SPSS ver. 12.0 (SPSS
Inc., Chicago, IL) was used for all analyses, with α set at 0.05.
Results
Diet
Twenty-six plant species were identified in scat samples, with a
mean of 1.7 ± 0.66 species per iguana (Table 2). In all, 50% of
scat samples contained only one species, 40% two, 9% three and
<1% four or five species. The most common species detected in
the scats were Vavaea amicorum, Micromelum minutum, Dio-
spyros elliptica, Jasminum didymum, Casuarina equesetifolia
and Hibiscus tiliaceus. Due to digestion of some plant material
before collection, 42% of iguanas had the remains of unidenti-
fied species in their scats; however, only 8% had solely uniden-
tified species in their scats. There was no difference between the
number of unidentified plant species in the scats from adults
versus juveniles (Student’s t-test: t67 = 1.59, P = 0.16)
Of the 26 plant species recorded, the leaves of 21 species
were eaten, as were the fruits of 14 species and the flowers of
6 species (Table 2). Ten species provided more than one food

Table 1. Number of Brachylophus vitiensis sighted and captured on surveys of Yadua Taba (all transects combined)
M, male; F, female; A, adult (sex undetermined); J, juvenile. The t-tests compare the heights of sighted and captured iguanas on each trip

Survey date No. of iguanas Mean heightB (m) of iguanas t d.f. P

Sighted CapturedA Sighted Captured

September 2005 311 102 5.39 ± 3.36 3.95 ± 2.32 4.33 266 <0.001
(M 126, F 106, A 49, J 30) (73) (0.5–18) (1–11)
December 2005 384 79 5.30 ± 3.38 3.33 ± 1.67 7.30 253 <0.001
(M 148, F 128, A 53, J 55) (42) (0–25) (1–10)
February 2006 362 77 5.24 ± 2.35 3.03 ± 1.32 10.73 218 <0.001
(M 131, F 124, A 46, J 61) (63) (1–17) (0.8–7)
May 2006 368 65 4.99 ± 2.44 2.33 ± 0.92 14.68 265 <0.001
(M 164, F 98, A 34, J 72) (55) (0.5–14) (0.5–5)
Total 1425 317
(M 569, F 456, A 182, J 218) (233)
A
Numbers in parentheses represent number of captured iguanas producing scats in captivity.
B
Mean height (±1 s.d.) and range of captured and sighted B. vitiensis are shown.
344 Australian Journal of Zoology C. Morrison et al.

Table 2. Plant species identified in 233 Brachylophus vitiensis scat samples

Parts of plants eaten: Fr, fruit; Fl, flowers; L, leaves. Other plant species were primarily specimens that
could not be identified due to partial digestion before sample collection

Plant species Parts eaten No. of scats in which plant was detected
Sept. 05 Dec. 05 Feb. 06 May 06 Total

Vavaea amicorum Fr, Fl, L 34 8 41 42 125

Micromelum minutum Fr, Fl, L 10 8 5 13 36
Diospyros elliptica Fr, Fl, L 14 5 10 1 30
Jasminum didymum Fr, Fl, L 4 9 4 7 24
Casuarina equisetifolia L 5 1 1 5 12
Hibiscus tiliaceus L 2 1 0 6 9
Grass (unknown species) L 4 0 2 2 8
Cordia subcordata Fr, L 3 2 2 0 7
Psydrex odorata Fr, L 3 1 0 3 7
Antirhea inconspicua Fr 2 0 1 3 6
Capparis quintifolia Fr 5 0 0 0 5
Eugenia reinwardtiana Fr, L 2 1 0 1 4
Guettarda speciosa Fr, L 1 3 0 0 4
Maesa persicifolia Fl, L 2 0 2 0 4
Pteris ensiformis L 0 1 0 3 4
Derris trifoliata L 0 1 2 0 3
Dichapetalum vitiense Fr, L 2 0 1 0 3
Passiflora foetida Fr, L 2 0 0 0 2
Psychotria sp. Fr 1 0 1 0 2
Alyxia stellata Fr 0 1 0 0 1
Croton metallicus L 1 0 0 0 1
Cynometra insularis L 0 1 0 0 1
Dendrolobium umbellatum Fr 0 0 0 1 1
Glochidion vitiense L 0 1 0 0 1
Mallotus tiliifolius L 0 0 1 0 1
Pongamia pinnata L 0 1 0 0 1
Other (unidentifiable species) – 34 27 22 23 106
Total no. of identified species 18 16 13 12 26

category (leaves, fruits, flowers), whereas four species provided

all three categories: V. amicorum, M. minutum, D. elliptica and
J. didymum.
60.00
The strong leaf and fruit pulse on Yadua Taba caused by cli-
matic seasonality was reflected in the plant component compo-
50.00 sition of B. vitiensis scats throughout the year. Leaves were the
most common item found in scats (found in 41–56% of scats)
regardless of season (χ2 = 36.52, P < 0.001) (Fig. 1). The pro-
Proportion of diet (%)

40.00 portion of iguanas that had consumed fruit ranged from 15 to

38% and was lowest in the mid-late dry season (September),
while the proportion of iguana scats containing flowers ranged
30.00
from 9 to 24% and was highest in May. In addition, there was
seasonal variation in the number of plant species consumed,
20.00 ranging from 12 to 18 species for our four sampling trips
(Table 2).
Within the six most common diet species (see above),
10.00 iguanas consumed mostly V. amicorum (present in 62% of
scats), a result that did not vary with season (2-way ANOVA:
Pmodel < 0.001, Pspecies < 0.001, Pseason = 0.090, Pseason × site =
0.00
0.252) (Fig. 2).
Sep. 2005 Dec. 2005 Feb. 2006 May 2006
There was no significant difference in the total number of
Trip date
plant species consumed by adult males and females (χ2 = 5.87,
Fig. 1. Seasonal variation in the proportion of different plant parts con-
P > 0.05), nor was there a difference between adults and juve-
sumed by B. vitiensis on Yadua Taba. Black bars, leaves; white bars, flowers; niles (χ2 = 13.39, P > 0.05). However, individual adult iguanas
striped bars, fruit; and grey bars, other parts (e.g. twigs). did have a greater number of plant species in their scats than
Diet and habitat of Fijian crested iguanas
individual juveniles (mean adults = 2.12 ± 0.85 (maximum 5),
mean juveniles = 1.35 ± 0.41 (maximum 3); Student’s t-test: t67
= 1.55, P = 0.04).
Perch sites
The 1425 iguanas recorded during nocturnal transect
searches were found in 33 tree species (Table 3). The majority
(70%) were found in six species: V. amicorum, D. elliptica,
Mallotus tilifolius, H. tiliaceus, Kingiodendrum platycarpum
and M. minutum.
In general, iguanas mainly utilised V. amicorum (33% of all
records) and D. elliptica (22%), a result that did not vary with
season (2-way ANOVA: Pmodel < 0.001, Pspecies < 0.001, Pseason
= 0.711, Pseason × site = 0.999) (Fig. 3). Juveniles utilised
Pongamia pinnata, Gyrocarpus americanus and K. platycarpum
as perches more than adults (χ2 = 35.65, P < 0.001) (Fig. 4).
There was no difference in utilisation of perch trees between
adult males and females (χ2 = 13.33, P > 0.05). There were also
no differences in perch heights of observed adult and juvenile
iguanas (Student’s t-test: t1433 = 1.19, P > 0.05), or observed
male and female iguanas (Student’s t-test: t1023 = 1.39, P > 0.05).
Floral composition and utilisation
We recorded 43 tree and shrub species on the six transects. The
most abundant plant species were the tree species V. amicorum
(37% of all plants), D. elliptica (21%) and M. minutum (19%).
50
40
Number of samples
30
20
10
0
V. amicorum D. elliptica
Fig. 2. Seasonal variation in common diet species found in scats of B. vitiensis on Yadua Taba. Black bars,
September 2005; striped bars, December 2005; white bars, February 2006; and grey bars, May 2006.
Australian Journal of Zoology 345
The densities of the primary plant species (in terms of both diet
and perches) ranged from 0 to 1400 plants ha–1, with most species
distributed over much of the island (e.g. V. amicorum), while a
few were more restricted (e.g. Cynometra insularis) (Table 4).
Most plant species appeared in the diet or as perches in
roughly the same proportion that they were found in the field.
There was a significant positive relationship between plant
abundance and iguana diet for D. elliptica (linear regression;
r2 = 0.759, P = 0.024) and H. tiliaceus (r2 = 0.991, P < 0.001),
and there were strong relationships between plant abundance
and perch utilisation in D. elliptica (r2 = 0.851, P = 0.009),
V. amicorum (r2 = 0.819, P = 0.013), M. tilifolius (r2 = 0.746,
P = 0.027), K. platycarpum (r2 = 0.829, P = 0.012), and H. tili-
aceus (r2 = 0.932, P = 0.002).
Discussion
Tree density versus iguana utilisation
Abundant tree species were generally more common in iguana
diet and as perches than less abundant plant species. There were
strong positive relationships between the abundance of plant
species and their utilisation by iguanas for D. elliptica (diet and
perch), H. tiliaceus (diet and perch), V. amicorum (perch),
M. tilifolius (perch) and K. platycarpum (perch). In a similar
study, Auffenberg (1982) found no correlation between food
plant density and utilisation in diet by the iguana Cyclura cari-
nata. He did, however, find a correlation between food plant
M. minutum J. didymum H. tiliaceus C. equisetifolia
Plant species
346 Australian Journal of Zoology C. Morrison et al.

density and iguana density, with areas of greater abundance of
food plants having more iguanas than areas with fewer food
plant species.
Important plant species
Our results are very similar to those in previous studies of
B. vitiensis that identified 13 plant species in the diet of the
iguanas from faecal or gut content analyses (Gibbons and
Watkins 1982; Gibbons 1984; Laurie et al. 1987; Harlow et al.
2007). The most common species in these studies were the fruit
of Passiflora suberosa and leaves of V. amicorum, C. insularis,
D. elliptica, M. tilifolius and H. tiliaceus. Previous studies
reported that 86% of iguanas had identifiable remains of more
than one species in their scats (P. S. Harlow, unpubl. data).
Previous diet results demonstrate that V. amicorum, D. elliptica,
H. tiliaceus and M. tilifolius account for more than 70% of
observations, as in our study (Gibbons and Watkins 1982;
Gibbons 1984; Laurie et al. 1987; Harlow and Biciloa 2001).
In the present study, the two most common species in both
diet and perch utilisation were V. amicorum and D. elliptica.
These were found in 62% and 15% of the scats, respectively, and
were two of the five plant species in which the leaves, flowers
and fruit were consumed by the iguanas. In addition, more than
55% of iguanas were found perching in one of these two species
across all seasons. These two species are also two of the most
abundant trees on Yadua Taba, and were recorded on all transects.
The tree M. minutum and the vine J. didymum are also impor-
tant food species for B. vitiensis, and were found in 16% and
10% of the scats, respectively. While we did not estimate the
abundance of J. didymum, M. minutum was the third most abun-
dant plant on our transects and was the perch site for 8% of
iguanas.
The native hibiscus (H. tiliaceus) was recorded in only 4% of
scats and was the perch tree for 6% of iguanas. It has previously
been recorded as an important food tree for B. vitiensis
(Gibbons and Watkins 1982; Gibbons 1984; Laurie et al. 1987;
Harlow and Biciloa 2001; Harlow et al. 2007), and almost 80%
of H. tiliaceus trees on our transects had at least one iguana. This
tree is common only on the southern end of the island, and was
recorded on only two transects.

Table 3. Tree species in which Brachylophus vitiensis were recorded during nocturnal transect searches
Total number and percentage of total lizards perched in each plant species are presented

Plant species Number of iguanas (% of total)

Sept. 2005 Dec. 2005 Feb. 2006 May 2006

Vavaea amicorum 93 (30) 116 (30) 126 (35) 144 (39)

Diospyros elliptica 84 (27) 85 (22) 79 (22) 63 (17)
Mallotus tilifolius 11 (4) 26 (7) 23 (6) 28 (8)
Hibiscus tiliaceus 12 (4) 21 (5) 21 (6) 21 (6)
Kingiodendrum platycarpum 19 (6) 23 (6) 12 (3) 6 (1)
Micromelum minutum 2 (<1) 11 (3) 18 (5) 22 (6)
Cynometra insularis 7 (2) 17 (4) 7 (2) 6 (1)
Gyrocarpus americanus 8 (3) 4 (1) 12 (3) 9 (2)
Psydrax odorata 7 (2) 13 (3) 5 (1) 6 (1)
Pogamia pinnata 6 (2) 1 (<1) 10 (3) 9 (2)
Evratamia orientalis 8 (3) 6 (2) 4 (1) 7 (1)
Descaspermum fruticosum 3 (<1) 5 (1) 8 (2) 8 (2)
Caesaria richii 6 (2) 10 (3) 5 (1) 3 (<1)
Eugenia rariflora 3 (<1) 7 (2) 1 (<1) 10 (3)
Pittosporum brackenridgei 4 (1) 4 (1) 2 (<1) 6 (1)
Derris trifoliate 4 (1) 4 (<1) 6 (2) 2 (<1)
Thesesia populnea 6 (2) 4 (1) 4 (1) 1 (<1)
Glochidion vitiense 5 (2) 4 (1) 3 (<1) 3 (<1)
Cordia subcordata 3 (<1) 3 (<1) 3 (<1) 4 (1)
Casuarina equisetifolia 3 (<1) 1 (<1) 1 (<1) 2 (<1)
Ficus barclayana 2 (<1) 1 (<1) 2 (<1) 2 (<1)
Terminalia littoralis 2 (<1) 2 (<1) 1 (<1) –
Maesa persicifolia 1 (<1) 3 (<1) – 1 (<1)
Planchonella grayana 3 (<1) – 2 (<1) –
Premna protrusa 2 (<1) – 2 <1) 1 (<1)
Antirhea incospicua 1 (<1) 2 (<1) – 1 (<1)
Melocia degneriana 2 (<1) – 1 (<1) –
Lantana camara – 1 (<1) 1 (<1) –
Drypetes vitiensis 2 (<1) – – –
Ficus oblique 1 (<1) – – –
Calophyllum inophyllum – – – 1 (<1)
Cordyline fruticosa – 1 (<1) – –
Other – dead trees, ground, etc. 3 (1) 6 (2) 2 (<1) 1 (<1)
Unknown species 3 (1) 6 (2) 2 (<1) 1 (<1)
Total iguanas 3 (1) 6 (2) 2 (<1) 1 (<1)
Diet and habitat of Fijian crested iguanas Australian Journal of Zoology 347

Seasonal variation
In highly seasonal tropical habitats, food may be limited during
part of the year (Iverson 1982; Blázquez and Rodríguez-Estrella
2007). During times of harsher climate (e.g. colder, drier, etc.)
food items tend to be restricted, and, in the case of herbivores,
food items tend to be more difficult to digest (e.g. primarily
leaves) and are of lower caloric content than at other times of the
year (Iverson 1982; Blázquez and Rodríguez-Estrella 2007). In
turn, this is thought to lead to seasonal variation in the diet of
herbivorous species in these habitats.
While there was little seasonal variation in the proportion of
primary plant species eaten by B. vitiensis (whether leaves, fruit,
flowers, etc.), our data suggest that there is seasonal variation in
the consumption rates of some of the less abundant but poten-
tially ‘important’ species, in particular the fruiting vine species
such as J. didymum. When J. didymum fruit is available it is
selected by iguanas. At other times of the year the leaves and
flowers comprise a much smaller proportion of the diet.
Seasonal opportunistic consumption was also observed for the
vine species Capparis quintifolia, Psychotria tephrosantha and
Alyxia stellata, for which only the fruit was consumed. Similar
observations in seasonal consumption of fruit have been made
for other iguana species (Auffenberg 1982; Durtsche 2000;
Blázquez and Rodríguez-Estrella 2007) and lizard species
(Schoener et al. 1982; Hodar et al. 1996).
In addition, our data suggest that there is seasonal variation
in the number of plant species consumed. In the drier months
when there is limited flowering and fruiting, iguanas consume a
wider variety of plant species (primarily leaves), and when there
is heavy fruiting and flowering they are more specific and focus
their efforts on fewer fruiting species (presumably those more
‘profitable’ in energy terms) (Blázquez and Rodríguez-Estrella
2007).
Age–class and sex variation
While several previous studies have examined diet in other
iguana species (Auffenberg 1982; Iverson 1982, Christian et al.
1984; Durtsche 2000; Blázquez and Rodríguez-Estrella 2007),
few have investigated differences between adult males and
females, and between adults and juveniles (most studies that have
done this have compared captive specimens, e.g. Auffenberg
1982). Our results demonstrated no significant sex or age–class
differences in the diversity of plants consumed by B. vitiensis.
This suggests that, on average, all iguanas are eating the same
plant species. In a similar study, Auffenberg (1982) found no sig-
nificant difference in the numbers of species or quantity of food
consumed by male versus female Cyclura carinata.
Although there was no significant difference in the overall
diversity of plants consumed by adults and juveniles, we did
find that individual juveniles appeared to consume fewer
species, on average, than individual adults. Similar results have
been recorded in C. carinata (Auffenberg 1982) and Lacerta
lepida (Castilla et al. 1991), with the differences being
attributed to the differences in body and gape size between
adults and juveniles.

150

120
Total number of iguanas

90

60

30

0
D

P.
M

K.

P.
G

E.
V.

H
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.t
.a

.t
.m

pi
pl

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ilif

ilia
llip

nn
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ar
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ar
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an

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Plant species

Fig. 3. Seasonal variation in common perch trees for B. vitiensis on Yadua Taba. Black bars, September 2005;
striped bars, December 2005; white bars, February 2006; and grey bars, May 2006.
348 Australian Journal of Zoology C. Morrison et al.

350

300

250
Total number of iguanas

200

150

100

50

0
D

P.
M

K.

P.
G

E.
V.

H
.e

.t
.a

.t
.m

pi
pl

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ilia
llip

nn
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Plant species

Fig. 4. Common perch trees for adult (black bars) and juvenile (white bars) B. vitiensis on Yadua Taba.

There were some differences between adult and juvenile
perch tree species, with juveniles utilising P. pinnata, G. ameri-
canus and K. platycarpum more than adults. There are two
potential reasons why this may occur. First, these trees are much
taller with less lower branching than the other common trees,
and are therefore more difficult for rats (Rattus exulans) to
climb and prey upon juvenile B. vitiensis. Second, these trees are
common in areas where nesting occurs, so the higher abundance
of juveniles in these tree species may be an artefact of the
greater abundance of juveniles in the area.
Limitations of the data
A more complete record of diet would have been obtained by a
study of stomach contents as leaves and flowers of some plants
are almost completely digested, making identification difficult
(Auffenberg 1982). However, due to the threatened status of
B. vitiensis, gut content analyses were not possible in this study.
Seeds and flowers were the most useful and reliable compo-
nents for identifying dietary species, as most of the forest plants
on Yadua Taba have distinct fruit and flowers; however, flowers
tended to disintegrate during digestion. Also, as plants exhibit

Table 4. Density of major food and perch tree species on each of the six transects
(250 m × 10 m)

Plant species Density of plants per transect (plants ha–1)

1 2 3 4 5 6

Vavaea amicorum 248 1108 1400 1348 904 1024

Diospyros elliptica 1036 36 588 836 168 748
Micromelum minutum 188 300 784 480 1044 588
Cynometra insualris 488 0 0 0 0 0
Psydrax odorata 0 16 492 8 116 92
Mallotus tilifolius 108 0 172 180 360 156
Gyrocaprus americanus 80 48 56 128 108 192
Eugenia reinwardtiana 176 0 36 36 8 8
Hibiscus tiliaceus 0 0 0 92 28 0
Kingiodendrum platycarpum 0 0 76 136 8 0
Pogamia pinnata 36 0 0 76 0 136
Diet and habitat of Fijian crested iguanas
considerable variety in their resistance to digestion, with some
being slightly altered and others completely, our results may be
biased due to greater sample integrity (less digested) of some of
the species with thicker or more durable leaves. This potential
bias is expected to be small given that our results are similar to
those of previous dietary studies of B. vitiensis, and because
iguanas were never observed consuming plant species that were
not recorded in the scats.
In this study we did not survey the abundance of vine or
creeper species on transects, which prevented us from making
density comparisons with diet. As some of the primary food
species were vines (e.g. J. didymum), this is an issue that should
be included in future studies.
Implications for translocations of B. vitiensis
Before any species is reintroduced or translocated to an area, it
is critical that the cause of the original decline(s) is known and
abated, and that there is good knowledge of biology, habitat,
demography, population genetics, social structure and potential
diseases (Dodd and Seigel 1991). In the case of B. vitiensis,
there must be no exotic predators (cats, dogs, mongoose) or
destructive habitat practices (grazing by goats, fire) in potential
sites, and the TDF vegetation must be intact or have recovered
significantly from previous disturbance. There are several small
uninhabited islands in the west of Fiji that are potential trans-
location sites where the removal of threatening processes is fea-
sible (using the same techniques used on Yadua Taba) if
permission is granted by local landowning communities. The
TDF forest on most of these islands, however, has been greatly
degraded and it will take time for it to regenerate to a suitable
degree.
In addition to being an intact forest, a suitable mix of species
must be present to provide the dietary and other ecological needs
of the species. Overall, the iguanas utilised roughly 50% of the
tree and shrub species found on transects for diet or shelter but
primarily utilised only 11 species (22%), indicating that they are
not broad generalists. Similar results have been found in other
iguanids (Lichtenbelt and Van 1993; Durtsche 2000; Blázquez
and Rodríguez-Estrella 2007). This suggests that translocation
success of B. vitiensis and other species will be improved if
wildlife managers not only have knowledge of the preferred
habitat of the focal species, but are also aware of, and take into
consideration, specific diet and habitat requirements.
The TDF species that we regard as essential to B. vitiensis on
Yadua Taba (based on diet and perch preference) are V. amico-
rum (Yadua name cevua), D. elliptica (kau loa), M. tilifolius
(yagata), M. minutum (qiqila), H. tiliaceus (vau), C. insularis
(cibicibi), P. pinnata (vesiwai), J. didymum (gasau cebucebu)
and K. platycarpum (moivi). Provided that all threatening pro-
cesses previously mentioned have been eliminated, potential
sites with these plant species should be given priority for
translocations of B. vitiensis within the western Fiji islands.
Acknowledgments
Timaleti Vavuke, Eroni Matatia, Isaac Rounds, Alma Nacuva, Eleazar
O’Connor, Taina Waqatabilai, William Thomas, Pita Qarau, William
Waqavakatoga, Kalisi Fa’anunu, Suzie Morrison and Zac Pierce provided
valuable help in the field. Isaac Rounds and Gunnar Keppel provided iden-
tifications of plant specimens. Luke Shoo, Suzie Morrison and Nick
Australian Journal of Zoology 349
Clemann provided valuable comments on earlier drafts of this manuscript.
We also thank the National Trust of Fiji for permission to work on Yadua
Taba and the villagers of Denimanu for their generous hospitality. This
project was funded by an Australia and Pacific Scientific Foundation
(ASPF) grant to the senior author.
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Manuscript received 28 November 2007, accepted 1 April 2008