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ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN
1175-5334
(online edition)
Copyright © 2017 Magnolia Press
Zootaxa 4273 (3): 407
–
422
http://www.mapress.com/j/zt/
Article
407
https://doi.org/10.11646/zootaxa.4273.3.5
http://zoobank.org/urn:lsid:zoobank.org:pub:775B7652-3A6E-49A6-A297-C69B411D64DD
A new species of iguana Brachylophus Cuvier 1829 (Sauria: Iguania: Iguanidae)
from Gau Island, Fiji Islands
ROBERT N. FISHER
1,5
, JONE NIUKULA
2
, DICK WATLING
3
& PETER S. HARLOW
4
1
U.S. Geological Survey, Western Ecological Research Center, San Diego Field Station, 4165 Spruance Road, Suite 200, San Diego,
CA 92101-0812, USA. E-mail: rfisher@usgs.gov
2
The National Trust of Fiji, PO Box 2089, Government Buildings, Suva, FIJI ISLANDS. E-mail: jniukula@nationaltrust.org.fj
3
NatureFiji-MareqetiViti, P.O. Box 2041, Government Buildings, Suva, FIJI ISLANDS. E-mail: watling@naturefiji.org
4
Taronga Conservation Society Australia, PO Box 20, Mosman, NSW, AUSTRALIA. E-mail: pharlow@zoo.nsw.gov.au
5
Corresponding author. E-mail: Robert Fisher rfisher@usgs.gov
Abstract
The south Pacific iguanas (Brachylophus) currently have three recognized living species in Fiji. Recent surveys have un-
covered more specific variation (morphological and genetic) within the genus and have better defined the geographic rang-
es of the named species. One of these recent discoveries is a strikingly different iguana from all other island populations
in Fiji which is restricted to Gau Island of the Lomaiviti Province. Gau is the fifth largest island in Fiji and maintains
excellent upland forests in the higher elevations. We describe this population from Gau Island as a new species, Brachy-
lophus gau sp. nov., in recognition of its type locality.
Key words: Pacific islands, endemism, conservation, Iguanian, Brachylophus, Gau Island
Introduction
The genus Brachylophus currently consists of three living (B. bulabula, B. fasciatus, and B. vitiensis) and one
extinct (B. gibbonsi) species of iguanas from Fiji and Tonga in the South Pacific (Pregill & Steadman 2004; Keogh
et al. 2008). Additionally the extinct monotypic iguana genus Lapitiguana was also known only from Fiji and was
twice the length of the largest living Brachylophus (Pregill & Worthy 2003). Iguanas have had a deep history in the
South Pacific and these three extant species are the only true iguanidae remaining in the South Pacific. These
iguanas have possibly been present there for 40 million years and their closest relatives occur in the New World in
the North American southwest deserts (Townsend et al. 2011). Keogh et al. (2008) recently reviewed
Brachylophus using the available genetic and morphological data, resulting in the author’s description of the new
species Brachylophus bulabula. All three living species were restricted taxonomically to only the islands where
either of these data sets was available in that study. This avoided the issue of over predicting the range of these
species from islands where no actual data was collected. This was a conservative approach as the various Fijian
iguanas have been mapped as occurring on many more islands in various papers (Gibbons 1981, 1985; Gibbons &
Watkins 1982; Zug 1991; Morrison 2003) but since there are no vouchers from most of these islands they were left
as species uncertain in this recent taxonomic treatise (see Figure 1 in Keogh et al. 2008). Since 2009, we have been
visiting many of the islands where iguanas or their habitat are still known, but from which no tissue samples,
morphological measurements, or specimens exist. Our goal is to better define the taxonomic diversity within the
genus and identify manageable conservation units.
Iguanas have been known from Gau Island for decades but there had been no specimens previously identified
in museum collections for analysis (Watling 1986). Recent conservation work on Gau for the Fiji Petrel has
brought renewed field surveys, which have included several detections of these iguanas from around the island.
This area of Fiji is poorly collected and poorly known herpetologically (Zug 1991), so it is not surprising that this
population of Brachylophus might exhibit some distinctive characteristics. These characteristics uniquely
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distinguish it from the other three described species and we describe this population as a new species. Additionally
we revisited historic museum specimens to determine if any match this new species as recent reassessments of
historical specimens have found specimens of systematic significance (Ineich & Fisher 2016).
FIGURE 1. Map showing location of Gau Island, the only known location for this iguana species. No further details mapped
due to sensitive nature of the records.
Materials and Methods
Morphological measurements and color pattern descriptions are based on six different live specimens, five
preserved specimens (some of which were also observed in life), and photos of seven additional individuals
collected on Gau Island. All recent preserved specimens are housed at the University of South Pacific Herpetology
Collection, Suva, Fiji (SUVA H). A total of 232 specimens of other species, including possible undescribed
species, of Brachylophus were examined for comparison representing 38 other island populations. Few voucher
specimens of Brachylophus exist in museums, and most of these have no specific locality information, but we
searched for any that might represent this new species. Previously the majority of these museum vouchers were
studied resulting, in part, in the description of Brachylophus bulabula (Keogh et al. 2008) and the rediscovery of
the type of B. fasciatus (Ineich & Fisher 2016). In this study most of the new Brachylophus specimens analyzed
were captured live and measured in the field, and photographs were taken to document additional color pattern
characteristics. These live photos are critical as the color and color patterns typically fade rapidly in preservative.
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FISHER NEW FIJI IGUANA
The following list provides the character names and definitions, some of which are straightforward.
Previously in Keogh et al. (2008), many of these were utilized but not necessarily defined in the text. Some
changes in how they were recorded then have occurred recently as we discovered a greater phenotypic diversity in
Fijian iguana populations, and thus many new characters were added. Not all characters are discussed in the text as
they were not found to vary much within our current Brachylophus dataset but might be found variable later as we
add more island populations. Expanded definitions of all characters are presented in Appendix 1.
Measurements: Width at widest point of body bands; width at dorsal crest of bands; dorsal crest spine
maximum height; dorsal crest spine maximum width; ear diameter; eye diameter; forearm length; head height;
head length; head width; internaral distance; crus knee to heel; naris-eye length; width of nuchal band; width of
pelvic band; tail height; tail length; trunk length, between limbs; snout length, edge of eye to end of snout; snout-
vent length; mass for live specimens.
Scalation: Femoral pores, total of left and right side, exuding only; number of points on third digit combs; toe
combs with fused scales; postmental paired; total postmentals; mental single or divided; parietal eye size; head
scale size, shape; labial-nasal contact; rostral-nasal contact; nasal color and intensity; number of scales encircling
nasals, including labials; postnasal enlarged and bordering entire posterior portion of nasal; internasals total;
number of scales encircling rostral, including labials; size of postnasal/preocular scales; dorsal crest spines; nostril
placement; nostril shape; number combs digit three; number combs digit two.
Coloration: number dorsal bands; eye color; nuchal pattern. Additionally notes are taken on throat patterns and
color, nuchal pattern, tail color and banding.
Brachylophus gau sp. nov.
Gau Iguana (pronounced Ngau)
Figs. 2–7
Brachylophus fasciatus Gibbons 1984 (first map showing island record); Zug 1991; Morrison 2003 (on maps as literature
record).
Brachylophus bulabula Fisher, Harlow, Edwards, and Keogh 2008.
Holotype. SUVA H 0264; collected in forest patch 1 km behind Nukuloa Village, Gau Island, Republic of Fiji
(18°2'46.68"S; 179°18'11.41"E, datum WGS84); 243 meters in elevation; collected by Robert Fisher, Peter Harlow,
Tuverea Tuamotu, Joeli Vadada, Maleli Biciloa, Mark O’Brien, Poasa Qalo, 1 July 2013.
Paratypes. SUVA H 0265 female collected same date, collectors, and locality as holotype (Figure 1). SUVA
H 0266–0267 were collected at Nalaqere Creek (18°2'25.91"S; 179°17'12.05"E) on 2 July 2013, by the same
collectors as the type. SUVA H 0273 was collected at Waitabua Hill (18°2'S; 179°18'E, WGS84), Gau Island, 439
meters in elevation, on 10 July 2013 by Mark Fraser and Poasa Qalo. BMNH 55.8.16.1-2 collected at Sawaieke
(17°59'14"S; 179°15'12"E), Gau Island, between 12 and 27 September 1854 by John MacGillivray.
Diagnosis. This species has a unique combination of color pattern characters that distinguish it from all other
species of described Brachylophus (Figures 2–7, Table 1). It also does not match any museum specimens we have
previously seen in collections, except two from the BMNH that previously lacked specific locality info. These
specimens are included in the type series as we were able to determine their provenance as Gau Island (see below
in Provenance of Historic Specimens section). Although added to the type series, their measurements are left out
of comparisons, but included in Table 2. We found that there are no locality specific museum vouchers that could
represent extinct island populations of this species. Otherwise, this species differs from B. vitiensis by having
sexually dichromatism, with banded males and females that are either unicolor or with small spots, and a maximum
snout vent length for both sexes of 153 mm, versus 255 mm for B. vitiensis. It differs from B. bulabula and
fasciatus in that males and females have green throats, whereas in these species males have white solid or white
with green/grey spots or blotches, as do the females in most populations. Although in certain populations the
females in B. bulabula and fasciatus the throat will be unicolor green, but never the males. It differs from the other
three species by having a primarily green colored nasal scale. The mean snout vent length is 149.2 and max size for
B. gau is 153 mm (n = 7), versus adult B. fasciatus with a mean 154.5 mm SVL and a maximum 176 mm SVL (n =
57) and B. bulabula with a mean 156.6 mm and a maximum 195 mm SVL (n = 23) (Table 1).
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Description of holotype. Holotype is an adult male (Figure 2–3). Meristics are as follows: snout vent length
150 mm, tail length regenerated, live weight 80 gm, head length 31.4 mm, head width 19.3 mm, head height 18.1
mm, and jaw length 24.4 mm. It has 16 elongated/enlarged upper labials (9 left, 7 right) and 17 elongated/enlarged
lower labials (8 left, 9 right). Rostral enlarged and triangular wide at base, apparently split on the left side and
contacts with nasal scale only on the right side. Mental is incised half way anteriorly, with three postmental scales
between the labial scales. Tympanum is translucent and unpigmented, vertical and oval in shape with less height (5
mm) then the eye is wide (6.4 mm). Nostril scale squarish with egg shaped opening raised dorsally within the
scale, pale orange in color within the opening, fades to light-green over the rest of the scale. Parietal eye small and
scale containing parietal eye similar in size to surrounding parietal. Supraocular head scales smallest, other head
scales larger and polygonal shaped, a fused subocular scale on the right side, enlarged subocular scales on the left
side. Dewlap is small in size, slightly rounded and unicolor with the throat. Poorly defined gular pouch covered
with smaller scales.
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FISHER NEW FIJI IGUANA
FIGURE 2. Illustration of the holotype (SUVA H 0264; left) and paratype female (SUVA H 0265; right) of Brachylophus gau
sp. nov. The painting is from photographs of these specimens and others from life. Measurements to scale within illustration.
The holotype is missing later 2/3 of tail, but tail drawn here based on photos from other males. Painting by Cindy Hitchcock.
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FIGURE 3. Preserved holotype of Brachylophus gau sp. nov. (SUVA H 0264): A) dorsal view; and B) ventral view. Note
distinctive striping on rear legs and diffuse banding on dorsal, and lack of differentiation in throat color from ventrum and no
spotting, also tail lacking any distinctive ventral banding. Photo by Baravi Thaman.
Two nuchal bands on right side, one originating at tympanum and continuing posteriorly, reaching nuchal
crest, the other above shoulder, left side contains one band with some spotting; two dorsal body bands, both with a
dorsal bifurcation leading to green infusion within the bands similar to a saddle at the dorsum, one axial band, and
all bands not completely colored, with some background diffusion. Tail regenerated and three tail bands, banding
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FISHER NEW FIJI IGUANA
fades posteriorly into background tail coloration. Nuchal crest poorly defined, 67 modified scales, up to 1.6 mm in
height and 2.3 mm in width on the nuchal region. Dorsal scales small and conical shaped. Ventral scales much
larger then dorsal ones, elongated and strongly keeled, with posterior ends pointed and elevated.
FIGURE 4. Photo of a live male Brachylophus gau sp. nov., illustrating the green throat and diffusion within the dorsal bands.
Photo by Joerg Kretzschmar.
Limbs long and thin covered with keeled scales similar in size to ventral ones; forearm length 24.7 mm, crus-
knee to heal 29 mm. Digits elongated, palms covered with small keeled scales. Third digit of hind feet contains a
denticulate comb on proximate phalynx with some fusion of scales. Toenails very long and not worn. Total of 28
enlarged exuding femoral pores (15 left; 13 right). Tail laterally compressed in cross section, weakly crested for
first 30% then undifferentiated. Anterior dorsal scales on tail smaller than ventral ones. All tail scales keeled and
posteriorly all scales become similar in size and larger.
Color in life. Head is emerald green with a white stripe lining the lower jaw just at margin. No blue on face.
Eye is copper or coppery brown, and a light orange rim around the interior of the nostril. Throat color is unicolor
green but hints of some grey bars. No color break but a subtle transition between throat and venter. Dorsal body
background color is an avocado green and transitions to a sulfurous green ventrally. There are two wide bands
across the back of a blue/grey green color, but diffused with background color and head color via individual scales.
The bands do not cross to the ventral side and break into saddles at dorsal crest. Tail with three bands colored as
body, but the last two very faded, this is first third of tail then brown bands towards the anterior. The tail maintains
a white lateral border. Hindlimbs with striking recurved bars colored as with body stripes. Forelimbs are unicolor.
Dorsal crest scales are green and become less distinctive towards anterior.
Color in alcohol. Overall dorsal color is greatly darkened with banding less distinctive but still present.
Ventral maintains light-green to blue-grey coloration. Dewlap remains unicolor and not distinctive from ventral
coloration. Striking leg banding is obscured. Nasal scale retains light orange color within rim. Parietal eye
relatively obscure. Tympanum translucent. Tail becomes brownish and bands become less obvious towards tip.
Vari a t i on . The diagnosis above describes some of the variability in this species for the main characteristics
that distinguish it from the other living forms (Table 1). Variability in scalation and measurements for B. gau are
reported in Table 2. Overall it’s a gracile species with long legs and tail. Additional variability in this species is
described here and includes either fused or elongated subocular scales in all of the paratypes. Relatively high
femoral pore counts. Short dorsal crest height, but high number of dorsal crest scales. Males with two wide dorsal
bands, a pelvic band, and a partial nuchal band. Background color of males and females the same lime green.
Females have small blue spots anteriorly when looked at closely.
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FIGURE 5. Photos of two live female Brachylophus gau sp. nov., illustrating the overall green coloration and light spotting.
Photos by Mark Fraser and Theo Blossom.
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FISHER NEW FIJI IGUANA
FIGURE 6. Photo of preserved male paratype of Brachylophus gau sp. nov., (BMNH 55.8.16.2) collected during September
1854 by John MacGillivray on Gau Island. Photo by Sam Fisher.
FIGURE 7. Photo of preserved female paratype of Brachylophus gau sp. nov., (BMNH 55.8.16.1) collected during September
1854 by John MacGillivray on Gau Island. Photo by Sam Fisher.
Etymology. The specific epithet is for the only island where the species is known to occur, Gau.
Comparisons. Table 1 reports the most important characters that distinguish between B. gau and the other
three living Brachylophus species. One important difference is that this species has fused or elongated subocular
scales, and these scales are as long or longer as the eye, this condition is rare in Brachylophus, and absent in most
populations. Dorsal crest scales average 53.2 with a range of 45–60 (n = 50) for B. vitiensis and for B. bulabula
dorsal crest scales vary in number between 55–88 with an average of 66.2 (n = 14). B. gau has a greater count than
these two species with an average of 71.8 and a range of 67–82 (n = 7), and a lower count than B. fasciatus which
averages 74.6 with a range of 52–94 (n = 48). It also has the shortest dorsal crest scales of any species, with a
maximum height of 1.8 mm (n = 7). So comparing it to B. vitiensis, it is much shorter, with longer legs and tail,
thinner build with fewer dorsal crest spines. Its nasal is mostly the same color as the head with a slight orange tinge
on the rim. It has unicolor females, versus the banded females of B. vitiensis. When compared to B. bulabula, this
species is shorter by almost 25%, it has a uniform lime green throat, with few black spots, versus white throat in
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males that may have grey/green or black barring. It lacks a strong nuchal band in males, and has very little orange
on the nasal scale. Finally comparing B. gau to B. fasciatus, it has much wider dorsal bands in males, and a more
coppery eye, versus the gold eye of B. fasciatus. It lacks the pastel light blue that B. fasciatus tends to have on the
anterior of the face, and instead has unicolor lime green head color.
Distribution. Brachylophus gau appears restricted to the large inland forest patches on Gau Island (Figure 1).
Iguanas were reported from Gau by Watling (1986), but no specimens were collected or described. It continues to
be widespread in high quality forest habitat in the interior of Gau Island (Figure 8), with some individuals also in
degraded forest closer to the coast (Figure 9). Gau is about 136 sq. km. in size and is the fifth largest island in Fiji.
FIGURE 8. Photo showing extensive intact patch of moist forest at higher elevations on Gau Island. Photo courtesy of
NatureFiji-MareqetiViti.
Provenance of historic specimens. The two specimens at the Natural History Museum, London (BMNH
55.8.16.1-2) were obviously Brachylophus gau based on morphology, but lacked specific locality information
except that they were submitted by John MacGillivray, esq. and from H.M.S. Herald. A more complete log of the
overall expedition of the Herald within Fiji is presented by David (1995). By examining his expedition notes we
were able to determine that John MacGillivray directly discusses finding and collecting an iguana on Gau Island
(as he spells it Ngau) on page 55, on line 4 (Figure 10; MacGillivray 1855):
In the wood behind the beach I shot but lost a large parrot, and a singular chameleon—like green + variegated
lizard with extremely long tail among the foliage of an ivi (Tahitian Chestnut) tree to which it closely assimilated in
colour.
Later he summarizes his reptile samples from Gau Island collected between September 12
th
and 27
th
, 1854, and
highlights the Brachylophus, and accurately points out that as of this time there were no samples yet in the British
Museum (MacGillivray 1855). Additionally he discusses/collected Gehyra vorax, Laticauda sp. and Candoia
bibroni from Gau Island. This is from page 59 of his notes (Figure 11):
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FIGURE 9. Photo showing coastal mangroves and lowland forest patch on Gau Island, but severe habitat degradation between
these coastal habitats and the intact forest at higher elevations. Photo courtesy of NatureFiji-MareqetiViti.
FIGURE 10. Partial scan of page 55 of Volume II of John MacGillivray’s private journal from 12–27 September, 1854,
discussing field work near Sawaieke Village on Gau Island (MacGillivray 1855). This is his first time finding an iguana in the
field. Scan courtesy of National Archives, Kew.
Reptiles. Perhaps the most remarkable reptile of Ngau is a very handsome tree lizard reminding one of a
chameleon. In colour it is a yellowish green (matching the leaves of the trees among which it is found) with paler
bands of a light bluish grey tint. My largest specimen is 19.7 inches in length of which the tail forms no less than
15.4 inches. I do not find its genus described in the Brit. Mus. Catalogue, but, judging from the characters there
given it appears to be closely allied to Lophura + Physignathus especially the latter. An ugly brown gecko 10
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inches long was occasionally brought on board by the natives, and small and active lizards of one or two species
are abundant. A small banded Hydrophis—similar to one previously procured at Aneitium, was brought me the one
day. Land snakes seem to be plentiful all that I saw are of one species including two varieties. It is usually of a
greyish colour, steaked and mottled with black, and an individual of average size measures 3 ½ feet in length. It
approaches the genus Engyrus.
FIGURE 11. Partial scan of page 59 of Volume II of John MacGillivray’s private journal from 12–27 September, 1854,
discussing the reptiles collected on Gau Island during his trip there on the HMS Herald (MacGillivray 1855). He extends his
discussion of the iguana from Gau in this section. Scan courtesy of National Archives, Kew.
Discussion and Conservation implications. Our lack of knowledge of diversity within Brachylophus in the
southwest Pacific has the potential of permitting the extirpation and extinctions of other divergent populations of
iguanas (Harlow et al. 2007). The Gau iguana appears to be a single island endemic, as are several other iguana
populations that most likely warrant species status (Fisher et al. unpub.). The nearest populations of Brachylophus
to B. gau are Brachylophus cf. bulabula on Nairai Island, approximately 15 km to the northeast (Figure 12), and
Brachylophus cf. bulabula on Wakaya Island approximately 43 km to the northwest (Fisher et al. 2013). Further
analysis needs to be conducted to determine the relationships of these populations to B. gau. As with other
populations of iguanas in Fiji, this species is restricted to intact patches of forest and lives high in the canopy
(Figure 13). Currently much of the low to mid elevation forest on Gau is highly degraded and no longer serves as
iguana habitat, and iguanas are not known to cross these grassy or unsuitable plantation habitats to get to other
forest patches (Figure 12). Nothing is known of its reproduction but other species of Brachylophus lay one clutch
of a few eggs (4–6) per year or every two years. These are placed in a shallow hole in the soil under the forest
canopy.
Revisiting conservation strategies for this group of enigmatic iguanas in Fiji is a priority, as there continues to
be many threats in addition to forest loss and fragmentation, including invasive mammals such as mongoose, cats,
and rats, and these iguanas continue to be harvested for the illegal wildlife trade. Local outreach and education will
be an important tool in their long-term protection and the protection of their habitats into the future. Gau Island
had previously been identified as a site of conservation significance for Fiji due to its retention of an altitudinal
range of moist forest in a large single block that supports Fiji’s only endemic seabird (Olson et al. 2010). Although
approximately 40% of Gau contains forested habitats, many of these habitats occur as isolated patches at the lower
elevations. Although alien invasive species are widespread on the island, cats in particular have the greatest
impacts on iguanas (Gibbons 1984; Priddel et al. 2009). Thorough surveys across the island are required to
determine the actual range of the species, to identify critical hotspots of high abundance, and to identify cat control
areas. Whether this species is found in the highest uplands of the island is also unknown, as we only surveyed
coastal and at mid-elevation forests, with the highest record of an iguana being approximately 440 meters
elevation.
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As Gau Island is the home of the only endemic sea bird in Fiji (Watling 1986), it is presumed to have high
levels of endemism in other groups as exemplified by our study here and studies on groups such as invertebrates
and plants (Keppel et al. 2010; Sarnat & Moreau 2011). In a comparison of 1 ha vegetation plots across the Pacific,
Gau had the highest level of recorded tree endemism (56.5%) across islands (Keppel et al. 2010). So with Gau
Island there still is great opportunity to work with the local communities to conserve what is left of significant
habitat patches that can serve for both the conservation of the endemic iguana and the rest of the endemic
biodiversity on the island (Remling & Veitayaki 2016).
FIGURE 12. Photo from mid-elevation on Gau Island, looking across to Nairai Island, which is the next closest island that
currently is occupied by iguanas. Extreme habitat degradation is shown looking across the landscape, with invasive grasslands
and plantations separating coastal forest patches from higher elevation intact forest. Photo courtesy of NatureFiji-MareqetiViti.
Acknowledgments
Funding for this work came primarily from the International Iguana Foundation, which is greatly appreciated; with
additional funding from Mohamed bin Zayed Species Conservation Fund, IUCN- Oceania Program Office, Critical
Ecosystems Partnership Fund - Polynesia-Micronesia, Taronga Conservation Society Australia, San Diego Zoo
Global, and University of the South Pacific. Critical support has come from Mark O’Brien, Poasa Qalo, Mark
Fraser, and the Gau Petrel team. Cindy Hitchcock graciously painted Figure 1. We thank Mark Fraser for loaning
photographs, and field help. Permitting agencies include the National Trust of Fiji, Department of Environment,
Ministry of Education, Ministry of Immigration, and U.S. Fish and Wildlife Service. Joerg Kretzschmar and Theo
Blossom allowed use of their photos of other live Gau iguanas. NatureFiji-MareqetiViti supplied the habitat photos.
David Gower, Patrick Campbell, Jeff Streicher, and Sam Fisher helped with the historic assessments and
measurements at the Natural History Museum, London. The National Archives, Kew, was very responsive to my
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searches of historic documents. We thank Nunia Thomas, Tamara Osborne, Alivereti Naikatini, Jesse Grismer,
Maleli Biciloa, Pita Biciloa, Ramesh Chand, Tuverea Tuamoto, Sialesi Rasalato, and the many other people that
have helped measure iguanas over the last seven years. George Zug, Jesse Grismer, and one anonymous reviewer
provided excellent feedback on the manuscript. We thank the USGS Ecosystems Mission Area for funding and
project support. The use of trade, product or firm names in this publication does not imply endorsement by the U.S.
Government.
FIGURE 13. Photo showing typical forest composition within healthy forest patches on Gau Island. Because of cyclone
disturbance history, these patches maintain a diversity of different tree size classes (Keppel et al. 2010). Photo courtesy of
NatureFiji-MareqetiViti.
References
David, A. (1995) The Voyage of HMS Herald to Australia and the South-west Pacific 1852-1861 under the command of
Captain Henry Mangles Denham. Melbourne University Press, Carlton, Victoria, 521 pp.
Fisher, R.N., Thomson, P., Watling, D. & Harlow, P. (2013) Brachylophus bulabula (Fiji Banded Iguana). Predation.
Herpetological Review, 44, 140–141.
Gibbons, J.R.H. (1981) The biogeography of Brachylophus including the description of a new species, B. vitiensis, from Fiji.
Journal of Herpetology, 15, 255–73.
https://doi.org/10.2307/1563429
Gibbons, J.R.H. (1984) Iguanas of the south Pacific. Oryx, 18, 82–91.
https://doi.org/10.1017/S0030605300018755
Gibbons, J.R.H. (1985) The biogeography and evolution of Pacific island reptiles and amphibians. In: Grigg, G., Shine, R. &
Ehmann, H. (Eds.), The Biology of Australasian Frogs and Reptiles. Surrey Beatty and Sons, Sydney, pp.125–142.
Gibbons, J.R.H. & Watkins, I.F. (1982) Behavior, ecology, and conservation of South Pacific banded iguanas, Brachylophus,
including a newly discovered species. In: Burghart, G.M. & Rand, A.S. (Eds.), Iguanas of the World. Noyes, Park Ridge,
pp. 418–441.
Zootaxa 4273 (3) © 2017 Magnolia Press
·
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Harlow, P.S., Fisher, M., Tuiwawa, M., Biciloa, P.N., Palmeirim, J.M., Mersai, C., Naidu, S., Naikatini, A., Thaman, B.,
Niukula, J. & Strand, E. (2007) The decline of the endemic Fijian crested iguana Brachylophus vitiensis in the Yasawa and
Mamanuca archipelagos, western Fiji. Oryx, 41, 44–50.
https://doi.org/10.1017/S0030605307001639
Ineich, I. & Fisher R.N. (2016) Rediscovery of the 220-year-old holotype of the Banded Iguana, Brachylophus fasciatus
(Brongniart, 1800) in the Paris Natural History Museum. Zootaxa, 4138, 381–391.
https://doi.org/10.11646/zootaxa.4138.2.10
Keogh, J.S., D.L. Edwards, Fisher, R.N. & Harlow P.S. (2008) Molecular and morphological analysis of the critically
endangered Fijian iguanas reveals cryptic diversity and a complex biogeographic history. Philosophical Transactions of
the Royal Society B, 363, 3413–3426.
https://doi.org/10.1098/rstb.2008.0120
Keppel, G., Buckley, Y.M. & Possingham H.P. (2010) Drivers of lowland rain forest community assembly, species diversity and
forest structure on islands in the tropical South Pacific. Journal of Ecology, 98, 87—95.
https://doi.org/10.1111/j.1365-2745.2009.01595.x
MacGillivray, J. (1855) Private journal of John MacGillivray, naturalist, in HMS Herald, in a voyage to the Pacific: with charts
and views. Vol. II: with an envelope containing papers not copied into the Journal. The National Archives, Kew, ADM 7/852.
Morrison, C. (2003) A field guide to the herpetofauna of Fiji. Institute of Applied Sciences, University of the South Pacific,
Fiji, 121 pp.
Olson, D., Farley, L., Patrick, A., Watling, D., Tuiwawa, M., Masibalavu, V., Lenoa, L., Bogiva, A., Qauqau, I., Atherton, J.,
Caginitoba, A., Tokota’a, M., Prasad, S., Naisilisili, W., Raikabula, A., Mailautoka, K., Morley, C. & Allnutt T. (2010)
Priority forests for conservation in Fiji: landscapes, hotspots and ecological processes. Oryx, 44, 57–70.
https://doi.org/10.1017/S0030605309990688
Pregill, G.K. & Steadman, D.W. (2004) South Pacific iguanas: Human impacts and a new species. Journal of Herpetology, 38,
15–21.
https://doi.org/10.1670/73-03A
Pregill, G.K. & Worthy, T.H. (2003) A new iguanid lizard (Squamata, Iguanidae) from the later Quaternary of Fiji, Southwest
Pacific. Herpetologica, 59, 57–67.
https://doi.org/10.1655/0018-0831(2003)059[0057:ANILSI]2.0.CO;2
Priddel, D., Carlile, N., Moce, K. & Watling D. (2009) A review of records and recovery actions for the ‘Critically Endangered’
Fiji Petrel Pseudobulweria macgillivrayi. Bird Conservation International, 18, 381–393.
https://doi.org/10.1017/S0959270908007648
Remling, E. & Veitayaki, J. (2016) Community-based action in Fiji’s Gau Island: A model for the Pacific? International
Journal of Climate Change Strategies and Management, 8, 375–398.
https://doi.org/10.1108/IJCCSM-07-2015-0101
Sarnat, E.M. & Moreau, C.S. (2011) Biogeography and morphological evolution in a Pacific island ant radiation. Molecular
Ecology, 20, 114—130.
https://doi.org/10.1111/j.1365-294X.2010.04916.x
Townsend, T.M., Mulcahy, D.G., Noonan, B.P., Sites Jr., J.W., Kuczynski, C.A., Wiens, J.J. & Reeder, T.W. (2011) Phylogeny
of iguanian lizards inferred from 29 nuclear loci and a comparison of concatenated and species-tree approaches for an
ancient, rapid radiation. Molecular Phylogenetics and Evolution, 61, 363–380.
https://doi.org/10.1016/j.ympev.2011.07.008
Watling, D. (1986) Rediscovery of a petrel and new faunal records on Gau Island. Oryx, 20, 31–34.
https://doi.org/10.1017/S0030605300025886
Zug, G. (1991) The lizards of Fiji: Natural History and Systematics. Bishop Museum Press Honolulu, 136 pp.
APPENDIX
Description of the characters recorded for the iguana specimens. Some characters can only be recorded from living iguanas.
As the field study has progressed, additional characters have been added. Due to the technical difficulties in returning to
previously visited sites, and finding iguanas for capture and measurement, some characters are unrecorded for some islands/
populations. Additionally, some of the extreme value ranges recorded are from museum specimens with locality data that
cannot be validated. It is probable that each of the three described species include additional variation within them that
represented additional undescribed forms. Measurements are taken with dial calipers, except the snout vent length, which is
typically taken with a stiff ruler, and the tail length which is taken with a soft vinyl or fiberglass tape measure. All
measurements are recorded in millimeters. All counts and measurements are taken on the left side of the body, if possible.
Most counts are recorded from both sides and the average or left one is reported.
Measurements:
Width at widest point of body bands: This is measured horizontally at the widest point along the lateral side of the iguana. Width
is measured for each band starting with anterior, but on the left side only.
Width at dorsal crest of bands: Width is measured for each body band but at the dorsal crest. Again a measurement is taken for
each band starting with the most anterior one.
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Dorsal crest spine maximum height: This is measured at the highest spine, which is typically within the first 15 spines anteriorly.
Dorsal crest spine maximum width: This is measured at the widest spine which is typically within the first 15 spines anteriorly.
Measured where the spine hits the crest of the body.
Ear diameter: This is taken at the longest cross-section axis of the ear.
Eye diameter: This is measured across eye, at the widest point.
Forearm length: Measured from the inside of the palm to the outside of the elbow, while the arm is bent 90 degrees.
Head height: This is measured as the widest point between the parietals on top of the head to the gulars under the throat.
Head length: This is measured from the rostrum to the rear edge of the lower mandible on the lateral side of the head.
Head width: This is measured as the widest point on the sides of the head approximately at the temples. Perpendicular to the
body axis.
Internaral distance: This is the distance between the interior side of one naris to the interior side of the other naris.
Crus knee to heel: This is measured as the distance from the outside of the knee to the base of the heel.
Naris-eye length: Measured from the nearest point of the eye to the posterior edge of the nasal opening.
Width of nuchal band: If present it is measured where it hits the dorsal crest.
Width of pelvic band: If present it is the band that crosses to the dorsal crest above the hind legs.
Tail height: The highest point posterior to the hind legs. For males, two measurements are taken, one at hemipenes and another
at highest point posterior to hemipenes.
Tail length: Measured from posterior edge of cloaca to tip of tail. As the tails have no fracture planes, and a modified scale at
tip, it was easy to determine if tail was complete. Any variance from complete tail was noted.
Trunk length: Measured from the posterior insertion of the forelimbs to the anterior insertion of the hindlimbs.
Snout length: Measured as the anterior edge of eye to end of snout (rostral scale).
Snout-vent length: From the tip of rostral scale to the cloaca.
Mass for live specimens: Taken in grams with a pescola scale. Iguanas were either balanced on a tared rubber band or placed in
a tared cloth sack.
Scalation:
Femoral pores, total of left and right side, exuding only: For males, only pores with obvious dilation and exudate were counted.
Additionally, smaller pin-hole pores were counted for males and females, and for males a total of both was recorded.
Number of points/tines on third digit combs: The third toe has a modified set of discrete enlarged scales. Counted on both left
and right side; left side used for comparisons.
Toe combs with fused scales: This character is whether there is evidence of fusion between the tines on the combs of the toes if
the combs are present.
Postmentals paired: Yes if paired; no if odd number.
Total postmentals: Count of scale row directly behind mental, not counting labials.
Mental single or divided: Conditions were entire, divided, or quarters of divided (i.e. half).
Parietal eye size: Large or small were the conditions, large was the eye aperture similar in size to adjacent scales, small the
aperture was more of a pin-hole or almost absent.
Head scale size and shape: The scales in the center of the top of the head between the eyes and posterior to the parietal eye. This
is qualitative as to size and shape. Typically they were either small and knobby, or larger and polygonal shape and flat.
Labial-nasal contact: Whether the labial scales contact directly the nasal scale or not. Recorded for both sides.
Rostral-nasal contact: Whether the rostral scale contacts the nasal scales directly or not. Recorded for both sides.
Nasal color and intensity (see also Keogh et al. 2008): This is the color of the scale, and how bright that color is. Also whether
the color of the scale expands to adjacent scales or is confined to that scale. This character is maintained in preserved
specimens pretty well although the color does become somewhat muted.
Number of scales encircling nasal scales, including labials: This is the total count of scales touching, even at a corner, the nasal
scales. Recorded for both sides.
Postnasal scale enlarged and bordering entire posterior portion of nasal: This is the condition of a fused set of postnasal
scales bordering the posterior of the nasal scale, or whether they are not fused.
Internasals total: Total scales between nasal and directly posterior of rostral.
Number of scales encircling rostral, including labials: Count of scales adjacent to the rostral scale. Some might touch just at a
point or angle and these are included in the count.
Size of postnasal/preocular scales: Larger or smaller; generally if there are < 10 scales between nasal and preocular scales then
they are larger, > 10 generally smaller.
Dorsal crest spines: Count of the dorsal crest spines, from the first differentiated midline scale at posterior of head to the one
centered over the mid-point of the hind limbs.
Nostril placement: The nostril opening either centered within nostril scale, or offset. If offset then direction of offset is noted.
Nostril shape: The shape of the nostril opening is typically rounded, triangular, oval or egg shaped, or slit like.
Number combs digit three: The count of how many “toe combs” on the hindfoot of digit three if any. They could be present and
then a count is given, absent, or enlarged.
Number combs digit two: The count of “toe combs” on the hindfoot of digit two if any. They could be present and then a count
is given, absent, or enlarged.
