BIAWAK

Quarterly Journal of Varanid

Biology and Husbandry

Volume 4 Number 1
ISSN: 1936-296X

On the Cover:
Varanus caerulivirens
Varanus caerulivirens is a poorly-studied
member of the V. indicus complex, indigenous to the northern Moluccas, Indonesia.
To date, few observations of this species have
been made by scientists in the field, and documentation of its ecology have been limited
until now (Weijola, this issue).
The V. caerulivirens depicted on the cover
and inset of this issue were photographed by
Valter Weijola.The specimen to the left was
photographed basking on a fallen tree trunk
at Air Mangga, Obi Island, at ca. 600 m elev.,
on 2 January 2009 at ca. 1130 h. The specimen below was photographed around 1000 h
on 8 December 2008 near Tetawang, Halmahera.

BIAWAK

Quarterly Journal of Varanid Biology and Husbandry

Editor

Editorial Review

ROBERT W. MENDYK
Center for Science Teaching and Learning
1 Tanglewood Road
Rockville Centre, NY 11570, US
odatriad@yahoo.com

Michael J. Balsai
Department of Biology, Temple University
Philadelphia, PA 19122, US
Vze3vhpv@verizon.net

Associate Editors
DANIEL BENNETT
School of Biology, Leeds University
Leeds LS2 9JT, UK
mampam@mampam.com
Michael Cota
Thailand Natural History Museum,
National Science Museum,
Technopolis, Khlong 5, Khlong Luang,
Pathum Thani 12120, TH
herpetologe@gmail.com
Andr Koch
Zoologisches Forschungsmuseum Alexander Koenig
Section of Herpetology
Adenauerallee 160
D-53113 Bonn, DE
a.koch.zfmk@uni-bonn.de

Zoo Liaisons
GEORGE SUNTER
Herpetology Department, London Zoo
Zoological Society of London, UK
George.Sunter@zsl.org
BRANDON GREAVES
Department of Herpetology
Omahas Henry Doorly Zoo, US
brandonlgreaves@yahoo.com
MARKUS JUSCHKA
Aquazoo Dusseldorf, DE
markus.juschka@duesseldorf.de

Editorial Liaisons
JOHN ADRAGNA
Cybersalvator.com
john@cybersalvator.com
ARNAUD COLIN
Groupement dEtude des Varanids
indicus@msn.com
MATTHEW SOMMA
matt_varanid28@yahoo.com

Bernd Eidenmller
Griesheimer Ufer 53
65933 Frankfurt, DE
bernd.eidenmueller@t-online.de
Michael Fost
Department of Math and Statistics
Georgia State University
Atlanta, GA 30303, US
MFost1@student.gsu.edu
Ruston W. Hartdegen
Department of Herpetology, Dallas Zoo
650 South R.L. Thornton Freeway
Dallas, Texas 75203, US
ruston17@yahoo.com
Hans-georg horn
Monitor Lizards Research Station
Hasslinghauser Str. 51
D-45549 Sprockhvel, DE
Hans-Georg.Horn@rub.de
tim jessop
Department of Zoology
University of Melbourne
Parkville, Victoria 3010, AU
tjessop@unimelb.edu.au
Jeffrey M. Lemm
Applied Animal Ecology Division
Conservation and Research for Endangered Species (CRES)
Zoological Society of San Diego
15600 San Pasqual Valley Rd
Escondido, CA 92027, US
jlemm@sandiegozoo.org

Web Editor
RYAN OBERTO
ryan@ubbsworld.co.za

Software and Technical Support

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1 Tanglewood Road
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http://www.cstl.org

International Varanid Interest Group

www.varanidae.org
The International Varanid Interest Group is a volunteer-based organization established to advance varanid research, conservation, and husbandry, and to promote scientific literacy among varanid enthusiasts. Membership to the IVIG is free, and open to anyone with an interest
in monitor lizards and the advancement of varanid research. Membership includes subscription to Biawak, a quarterly journal of varanid
biology and husbandry, and is available online through the IVIG website.

Biawak

Quarterly Journal of Varanid Biology and Husbandry

ISSN 1936-296X

Volume 4 Number 1
March 2010

Organizational News...............................................................................................................

News Notes.............................................................................................................................

Geographic Distribution and Habitat Use of Monitor Lizards of the North Moluccas
.........................................................................................................VALTER S- WEIJOLA

Varanus spenceri: A Last Supper of Grasshoppers (Acrididae)

......................PATRICIA A. WOOLLEY, MARTIN J. STEINBAUER and GREG MIFSUD

24

Parthenogenesis in an Ornate Nile Monitor, Varanus ornatus

................................................................................................................JAMES HENNESSY

26

Book Reviews..........................................................................................................................

31

Recent Publications..................................................................................................................

42

2010 International Varanid Interest Group

Varanus albigularis. Hluhluwe Imfolozi Game Reserve, South Africa. Photograph by Jo-anne Hounsom.

ORGANIZATIONAL NEWS
New Web Editor
The International Varanid Interest Group welcomes
Ryan Oberto to the editorial board as web editor. A new
IVIG website, which will continue to host Biawak and
an assortment of educational resources, is currently in
development and should be online soon.

IVIG Logo Designs

Due to unforseen complications, decision of the winning
insignia design for the IVIG has been delayed, and will
be decided shortly, with the winning design announced
in the next issue of Biawak. .

Call for Papers on the

Herpetoculture of Varanids
Biawak is currently seeking papers describing the husbandry, breeding, and management of varanids in captivity. Submissions from private hobbyists and zoos are
both welcomed. Assistance with manuscript preparation
is available to those who inquire. For additional information, please contact submissions@varanidae.org

Varanus gouldii. Ewaninga Rock Carvings Conservation Reserve, Northern Territory, Australia. Photograph
by Ed Loveridge

Varanus salvator tracks. Teluk Kampi, Penang National Park, Malaysia Photograph by Chris Liberty chris@
chrisliberty.com

NEWS NOTES
Nile Monitor Hunt to Begin in
Florida

Indonesian Ranger Attacked by

Komodo Dragon

A hunt scheduled in the state of Florida aimed at reducing

the population of feral pythons will also include the
alien Nile monitor (Varanus niloticus). The hunt, to
take place March 8 through April 17, will occur in three
areas: Everglades and Francis S. Taylor, Holey Land
and Rotenberger Wildlife Management Areas. Anyone
with a valid hunting license and who pays an additional
$26 fee will be able to hunt the reptiles. Unlike previous
hunts, this one will be open to the public at large and not
just individuals who possess reptile-of-concern permits.
Hunters will be allowed to use all legal methods used
in taking game animals (excluding centerfire rifles), as
well as nets and snares. No live animals are allowed to
be removed.

An Indonesian park ranger was attacked and severely

wounded by a Komodo dragon on the island of Rinca.
Marselinus Sabanhadir was heading for the toilet when
a two-meter dragon which had been concealed behind a
guard post rushed towards him, grabbing his right foot
and causing deep lacerations before other rangers drove
the animal away. Sabanhadir was taken to a hospital on
Bali where he is recovering.
Source: Jakarta Globe - 23 February 2010; Mercury
News - 23 February 2010

Man Arrested with 40

Endangered Monitor Lizards

Source: Florida Fish and Wildlife Conservation

Commission 22 February 2010

A 48 year old man from West Delhi, India was arrested

and detained for 14 days of judicial custody for the

Combatting Varanus salvator macromaculatus. Lumpini Park, Bangkok, Thailand. Photograph by Tim Cox.

BIAWAK VOL. 4 NO. 1

illegal poaching of endangered monitor lizards (Varanus

bengalensis). Police seized a total of 40 monitors
which were bound within seven gunny bags when they
apprehended the subject in a park in Raghubir Nagar.
The man was planning on selling the animals for their
meat, at 2000 Rupees each (ca. $44 USD). The monitors
are planned to be released back into the wild.
Source: Indian Express.com - 13 Jan 2010

Cincinnati Zoo to Open New

Monitor Lizard Exhibit
The Cincinnati Zoo, Ohio, US, will be opening a new
Komodo dragon exhibit on 5 June 2010, after a four year
period without a dragon on display at the zoo. The new
exhibit will also feature several additional species of
monitor lizard, highlighting the largest, longest, smallest,
and some of the most colorful species in the genus.
Source: zandavistor.com - 29 January 2010

Varanus salvator salvator. Kandy, Sri Lanka.

Photograph by Lynda Hanwella.

Invitation to the Second Annual

Meeting of the AG Warane of
the DGHT
We are pleased to invite all those interested in monitor
lizards to the second annual meeting of the DGHT-AG
Warane. The meeting will take place on 24 April 2010
in Hanau near Frankfurt. The participation fee for nonmembers is 10,- , members are free of charge. The fee
will be set against the membership fee for new members
who attend the AG-Warane at the meeting. AG-members
have to be members of the DGHT (www.dght.de).
Location: Cafe-Restaurant Sandelmhle
Philipp-August-Schleissner-Weg 2a

63452 Hanau
Schedule of talks and presentations:
10:30 Begin of the meeting
11:00 Welcome: Kay Dittmar and Andr Koch,

leaders of the AG-Warane
11:15 Klaus Wesiak (Frankfurt) & Andr Koch

(ZFMK, Bonn): Successful husbandry and

first breeding of Varanus juxtindicus Bhme

et al., 2002, with remarks on the development

of juveniles of this rarely-kept endemic

Solomon monitor species (in German).

See report in Biawak 3(4): 106-121.
12:00 Lunch break and group photo shoot
13:30 General meeting (inofficial)
14:00 Thomas Hrenberg (Stuttgart): Keeping and

breeding of Macraes tree monitor (Varanus

macraei) and caesarean section in V. (Odatria)

tristis orientalis (in German).
14:45 Monika Labes (Germering): Report on the

Husbandry and Breeding of Varanus

exanthematicus (in German).
15:30 Coffee break
16:00 Andr Koch (ZFMK, Bonn): News about

Southeast Asian monitor lizards (in German).
16:30 Jochen Meyer (Stuttgart): Remarks about

keeping and breeding of Varanus rudicollis (in
German).
17:00 Farewell and social gathering
Further information can be found on the AG Warane homepage
at www.ag-warane.de, or contact Kay Uwe Dittmar (working
group leader) at dittmar@ag-warane.de or Andr Koch
(scientific leader) at a.koch.zfmk@uni-bonn.de.

ARTICLES
Biawak, 4(1), pp. 7-23
2010 by International Varanid Interest Group

Geographical Distribution and Habitat Use of

Monitor Lizards of the North Moluccas
VALTER S- WEIJOLA
Departement of Biosciences
bo Akademi University
FI-20520 Turku, Finland.
Email: vweijola@abo.fi
Abstract: Observations on the ecology and distribution of monitor lizards were made on three different
field visits to Halmahera, Morotai, Bacan, Gebe and Obi islands. Data was collected on four species:
Varanus caerulivirens, V. rainerguentheri, V. cf. salvator and V. yuwonoi. No specimens of V. zugorum
were observed during the course of the study, and this species also appears to be virtually unknown
even among local hunters and animal collectors. Varanus caerulivirens, V. rainerguentheri and V. cf.
salvator are reported from Obi for the first time. Additionally, V. caerulivirens and V. rainerguentheri
were recorded from Morotai and Bacan. The different species appear to avoid competition by
differences in habitat use, body size, and foraging mode. Varanus rainerguentheri is largely restricted
to the coasts, while V. caerulivirens and V. yuwonoi primarily occur in inland forest habitats.
Introduction
Currently, the V. indicus species group is, despite
the efforts of several workers, quite possibly the least
resolved group of varanoid lizards in the world. They
occupy a vast and difficult to access geographical region
from the Solomon Islands in the east, across New Guinea
with its shelf islands and parts of northern Australia,
through the Moluccas as far as Talaud in the northwest,
as well as some remote Pacific Island groups. This is
a region of unparalleled complex geological history
consisting of thousands of variously sized islands where
isolation and other evolutionary forces have driven
populations to diverge and differentiate, resulting in a
phylogenetic and ecological diversity that is still poorly
understood.
The Moluccan islands (Maluku) of east Indonesia
forms part of the biogeographical region known as
Wallacea. Traditionally considered a transition zone
between the Australian and Asian faunas, many of the
islands also hold a considerable number of endemic
species and have at least for some animal groups played
an important role in speciation processes (How and

Kitchener, 1997, Ziegler et al., 2007b). The number of

identified species in the Varanus indicus species group
in the Moluccas has risen from one to eight during the
last decade (of which seven are endemic to this region),
and more will inevitably be described (e.g., Weijola
and Sweet, in press). Sweet and Pianka (2007) review
the reasons for the high diversity of small monitors
(less than 130 cm total length) east of Wallaces Line,
concluding that the historical absence of predatory
placental mammals has played a fundamental role in the
radiation of lineages such as Euprepiosaurus.
The numerous islands with their different
communities provide the ideal natural experiment to
study resource partitioning and ecological release in
monitors. Since studies on varanid communities have
historically been largely restricted to dry and seasonally
wet Australian environments, it seemed relevant to
compare these dry communities with ones in the
wet tropics, such as in the Moluccas. It is particularly
interesting since most of the species within Moluccan
communities are very similar in size and more closely

BIAWAK VOL. 4 NO. 1

related than within most Australian communities (where

assemblies include species of several subgenera).
Despite the recent discovery boom of new species,
there has been very little fieldwork involved, and all
the descriptions, except for V. lirungensis Koch et al.
(2009), have been based solely on museum or animal
trade specimens. The first ecological observations of
many of these species were made during this study
on three separate field trips in 2008 and 2009. This
paper concerns varanids of the larger islands of the
biogeographical northern Moluccas: Halmahera,
Morotai, Bacan, Kasiruta, Gebe and Obi. The Sula
Islands are also included in the administrative unit,
but group biogeographically with Sulawesi and are not
considered further here.
Methods and Study Area

Field work was conducted during March 2008,
December-February 2008-2009, mid April- mid June,
and late October-mid November 2009. Investigations
were made in coastal habitats (Fig.4), lowland forests

Fig. 1. Map of the Moluccas with the study area darkened.

(Fig.3), swamps (Fig.5), and hill forests (Fig.2) on

Halmahera, Bacan, Morotai, Gebe and Obi (Fig.1). These
are all climatologically rather similar and are, or have
historically been, covered primarily by tropical rain- and
evergreen forest. At present, much of the lowland forests
are degraded, converted to plantations or even mined
for minerals. Annual rainfall varies according to land
topography, but averages between 1500 and 2000 mm.
(Monk et al., 1997; Bacan Agricultural University, pers.
comm. 2009), and lowland temperatures are 25-30 oC
year-round. December through March tends to receive
the heaviest rainfall, but most of the Moluccas rarely
experience prolonged dry periods.
Many of the V. indicus group monitors can be
observed by searching on foot in suitable habitats.
Alternatively, some of the shyer species can be attracted
to bait (fish/meat). Varanus yuwonoi is particularly
difficult to observe, and the author was forced to follow
a professional animal collector at work in order to see
this species at all. Locating monitor lizards in tropical
wet forests presents particular difficulties since it
excludes the possibility of using tracks as aid and the

WEIJOLA - MONITOR LIZARDS OF THE NORTHERN MOLUCCAS

dense vegetation provides ample cover and places to

hide. Recurrent periods of rain and clouds often decrease
activity levels of the animals, making observations
difficult for days or weeks on end. For each observation,
a set of data (most importantly: habitat use, location and
activity etc.) were recorded. In a few cases, dead animals
were encountered and stomach content and reproductive
condition was analyzed.
Abbreviations for museum collections mentioned in
this article are: BYU: Brigham Young University, Utah;
RMNH: Naturalis, National Museum of Natural History,
Leiden; MZB: Museum Zoologicum Bogoriense, Java
Halmahera
Halmahera is the largest and most geologically
complex island in the Moluccas (Hall, 1998). It is of
composite origin and consists of a younger, volcanic
western part and an older eastern part. These collided
somewhere between one and three million years ago
(Hall, 1999), after having moved westwards for 15-20
million years. Their composite nature and historical
proximity to New Guinea probably contributes to the
high species diversity of this island. Four species have

Fig.2. Mangrove forest, Tetewang, Halmahera

been confirmed (V. caerulivirens, V. rainerguentheri, V.

yuwonoi and V. zugorum), which is more than on any
other Moluccan island.
Observations were made around a number of sites
on the northwestern, northeastern and eastern peninsulas
in a variety of habitats from coastal mangrove swamps,
Nypa swamps, coastal forests, sago swamps, plantations,
secondary forest, disturbed forest, and primary forest, at
elevations from 0 to 700 m.
Morotai
Morotai is a part of the East Halmaheran Crustal
Fragment (Hall, 1999) and is at present separated from
Halmahera by about 10 km of open sea. According to
Voris (2000), Morotai would have been connected to
Halmahera by land bridges repeatedly during Pleistocene
glacial periods. Only one field site was visited on
Morotai: the surroundings of Pilowo village on the
southern end of the island. There are vast mangrove
forests in this area but unfortunately the lowland forests
are disturbed or secondary. Most observations were
made in the mangroves.

BIAWAK VOL. 4 NO. 1

Fig. 3. Nypa-palm swamp, Kasiruta.

Fig. 4. Freshwater stream and lowland forest, Sumahode, Halmahera.

10

11

WEIJOLA - MONITOR LIZARDS OF THE NORTHERN MOLUCCAS

Bacan Islands

Gebe

The Bacan islands are geologically a part of western

Halmahera (Hall, 1999) and were also periodically
connected with southern Halmahera during the
Pleistocene (Voris, 2000). Two field sites were based on
the island of Kasiruta, one at Dobo (north east coast)
and the other at Kasiruta Dalam (southern interior). The
site at Dobo was poor since most of the surroundings
were heavily converted to nutmeg and clove plantations.
Kasiruta Dalam was reached by ascending the Kasiruta
River by a small motorized canoe. It is a small village
surrounded by sago swamps and disturbed lowland
forest.
On the main island of Bacan, observations were
made outside of Labuha and Wayamiga. Wayamiga lies
at the foothills of the mountain Gunung Sibela, from
where a day long excursion into the virgin hill forests
was made.

Gebe is situated ca. 40 km southeast of the

easternmost tip of Halmahera and 70 km west of Waigeo.
The geology is mostly Tertiary limestone (raised reef)
and ophiolitic red soils. The western part has been
heavily mined for nickel and large areas have been
destroyed during the past decades. Thus observations
were concentrated in the eastern part where pockets
of primary lowland forest, drier limestone forests and
mangroves remain.

Fig. 5. Hill forest on Gunung Sibela, Bacan; habitat of

V. caerulivirens.

Fig. 6. Limestone forest, Air Mangga, Obi.

Obi
Large portions of Obi are composed of raised reef
limestone (Monk et al., 1997). Field work was conducted
at two sites in the northeastern part of the island. The
coasts were heavily forested by mangroves, coconut
groves and coastal swamps, with inland areas including
cocoa plantations, disturbed hill forest, and limestone
forests.

BIAWAK VOL. 4 NO. 1

Results
Species accounts
Up to nine species have been reported to occur on these
islands collectively; V. caerulivirens, V. cerambonensis,
V. doreanus, V. indicus, V. melinus, V. rainerguentheri, V.
salvator, V. yuwonoi and V. zugorum. Of these, Varanus
melinus was erroneously reported from Obi (Bhme and
Ziegler, 1997), but later omitted by Ziegler and Bhme
(1999); the absence of V. melinus from Obi is supported
by fieldwork (this study). Examination of photographs
of voucher material (RMNH 3184) and fieldwork
reveals that the record for V. cerambonensis from Obi in
Philipp et al. (1999) is based on a misidentification of V.
rainerguentheri, and thus Obi should be omitted from the
range of this species. Based on four 19th century voucher
specimens, Koch et al. (2007) included both Seram and
Halmahera in the range for V. salvator; however, these
records are not supported by fieldwork (this study;
Edgar and Lilley, 1993; Setiadi and Hamidy, 2006) or
interviews with locals and local dealers (T. Baadilla and
B. Baadilla, pers. comm. 2009) and should be omitted
until more substantial evidence is presented. Varanus
cf. salvator is however here documented to be native to
Obi. Yuwono (1998) mentioned a record of V. doreanus
from Halmahera, and genetic studies by Ziegler et al.
(2007b) include a specimen allegedly from that island,
but fieldwork does not support such an occurrence. Thus
five species are confirmed from within this region: V.
caerulivirens, V. rainerguentheri, V. cf. salvator, V.
yuwonoi and V. zugorum.

on that island. No observations were made on Kasiruta,

a large island just west of Bacan, although considering
the close proximity between these islands it is likely that
they do occur there.
This species was also recorded in the field on Obi
for the first time, an island separated from Halmahera by
almost 60 km of open sea. On Obi, they are sympatric
with V. rainerguentheri and V. cf. salvator. Investigations
on Gebe strongly suggest that V. caerulivirens does not
reach that island, most likely making east Halmahera
and Morotai the eastern limit for its range.
Ecology: A total of 59 observations of V. caerulivirens
were made during almost five months of work throughout

Varanus caerulivirens
Despite having been described so recently (Ziegler
et al., 1999), the turquoise monitor (Figs. 7 and 8) is
widely distributed in the northern Moluccas and is
actually often quite common in suitable habitats.
Distribution: The type locality for V. caerulivirens is the
northern Moluccan island of Halmahera, and the only
precise locality previously published is Patani at the far
eastern tip of that island (Ziegler et al., 2004). A specimen
collected on Morotai in 1944 (BYU 7477) confirms its
presence on that island. Investigations on Halmahera
found the species to be widely distributed throughout the
lowlands up to about 700 m elevation in all but the most
disturbed forested habitats. Several sightings were also
made in primary forests in Gunung Sibela on the island
of Bacan, and this species is expected to be widespread

12

Fig. 7. Varanus caerulivirens. Tetewang, Halmahera.

13

WEIJOLA - MONITOR LIZARDS OF THE NORTHERN MOLUCCAS

Fig. 8. Varanus caeurulivirens. Air Mangga, Obi.

BIAWAK VOL. 4 NO. 1

its range. This species is a forest generalist and occurs

in most non-salt water influenced forest types: from
coastal forest and beach vegetation, plantations, primary
and secondary lowland forests, limestone-, alluvial, hill- and mountain forests up to at least 600-650 m
elevation. They are often found basking around small
clearings. Occasionally, specimens are encountered
while patrolling freshwater streams, and on one occasion,
two specimens were observed near a lake at over 600 m
elevation (Tables 1 and 2).
This species is most often encountered while actively
foraging through the leaf litter layer of the forest floor,
and stomach content analysis reveals that they consume
burrowing animals such as earthworms, mole crickets,
etc. They almost invariably seek refuge in trees when
frightened by or pursued by humans and use tree cavities
as night refugees, and probably for nesting as well. The
claws and scales of the feet are well adapted for climbing
and they make much use of tree trunks and canopies for
basking, sometimes even jumping from one tree to the
other.
No significant difference in habitat utilization could
be observed between juveniles and adults - both were
equally terrestrial and arboreal, though admittedly far
fewer sightings were made of small vs. large individuals.

Fig. 9. Varanus rainerguentheri. Ibu, Halmahera.

14

It is possible that younger individuals make more

extensive use of tree hollows.
Dissection of sexually mature specimens revealed
males to have enlarged testes throughout the year,
suggesting that reproduction takes place year-round. The
largest adult specimen measured 110 cm in total length,
which is probably near maximum size.
Varanus rainerguentheri
Varanus rainerguentheri (Figs. 9-14) was described
as a cryptic species within the V. indicus complex based
on molecular and morphological evidence (Ziegler et
al., 2007a). Unfortunately, the genetic comparison of the
new species with V. indicus in that paper was restricted to
a single sequence of mtDNA from a V. indicus specimen
without specific locality data. Additionally, the type
locality of V. rainerguentheri (Jailolo, Halmahera)
is disputed by the original collector (L. Wagner, pers.
comm. November 2008), making the genetic and
morphological argumentation a comparison between
two unknown localities.
Two of the characteristic morphological features
of the holotype were the blunt snout and occurrence
of a light postocular stripe. The blunt snout was never

15

WEIJOLA - MONITOR LIZARDS OF THE NORTHERN MOLUCCAS

Fig. 10. Head and tongue of V. rainerguentheri. Tetewang, Halmahera.

Fig. 11. Varanus rainerguentheri. Pilowo, Morotai.

BIAWAK VOL. 4 NO. 1

Fig. 12. Varanus rainerguentheri. Air Mangga, Obi.

Fig. 13. Adult V. rainerguentheri. Gebe Island.

16

17

WEIJOLA - MONITOR LIZARDS OF THE NORTHERN MOLUCCAS

Fig. 14. Juvenile V. rainerguentheri. Gebe island.

observed in the field and appears to be unique to that
particular specimen. Field investigations of larger
sample sizes also show that the postocular stripe is
variable among Halmaheran and north Moluccan
populations of V. rainerguentheri, and usually fades with
age. For example, the two specimens depicted in Setiadi
and Hamidy (2006) as V. indicus and V. rainerguentheri
appear to only show intraspecific variation, and cannot
be allocated to different species based solely on the
occurrence of a temporal stripe. The temporal stripe is
usually less pronounced on specimens from Obi and
Gebe compared to Halmahera and Morotai, though
based on external features, the populations on these
islands are very difficult to distinguish. Specimens from
Obi do appear to have dark pigmentation further back
on the tongue.
Despite the potential weaknesses in the original
description, I have opted to use the name V.
rainerguentheri for populations of the V. indicus-type
monitors in the northern Moluccas included in this
paper. A more detailed taxonomic investigation of
animals from verifiable localities and with larger sample
sizes is much needed to confirm the taxonomy and

specific characteristics of this species, particularly since

the description of V. rainerguentheri failed to make a
comparison with the very similar animals of nearby
Waigeo, which were described as V. chlorostigma
by Gray (1831). This name was synonymized with V.
indicus by Bhme et al. (1994) and earlier authors,
but the subsequent redefinition of V. indicus (Philipp
et al. 1999) excludes animals from Waigeo on several
characteristics (such as throat markings, tongue color
and scalation). The redefinition of V. indicus invalidates
this earlier synonymization, and makes V. chlorostigma
a potentially available name for V. rainerguentheri.
Distribution: Varanus rainerguentheri is widely
distributed, particularly around the coastlines, on
Halmahera, Ternate (RMNH voucher), Tidore (MZB
voucher), Morotai, Bacan, Kasiruta, Gebe and Obi (and
probably on many of the smaller islands of this region).
Ecology: One hundred and twelve (112) observations
were made during the course of fieldwork. On all
islands except Gebe, they are mostly restricted to
coastal areas and mangrove swamps (Tables 1 and 2).

BIAWAK VOL. 4 NO. 1

18

Table 1. Habitat use of monitors on Halmahera, Morotai and Bacan.

Habitat
Mangrove
Beach, littoral vegetation/swamp
Nypa swamp
Sago swamp
Brackish water river delta
Freshwater stream and river
Inland lake
Secondary lowland forest 2-50 m.a.s.
Primary lowland forest 2-50 m.a.s
Hill forest 50-200 m.a.s
Hill forest over 200 m.a.s
Plantation
Total # of observations

V. caerulivirens
1 (1.8%)
5 (8.9%)
2 (3.6%)
20 (35.7%)
11 (19.6%)
15 (26.8%)
2 (3.6%)
56

V. rainerguentheri
13 (38.2%)
6 (17.7%)
3 (8.8%)
6 (17.7%)
3 (8.8%)
3 (8.8%)
34

V. yuwonoi
2 (66.7%)
1 (33.3%)
3

Table 2. Habitat use of monitors on Obi.

Habitat
Mangrove
Beach, littoral vegetation/swamp
Nypa swamp
Sago swamp
Brackish water river delta
Freshwater stream and river
Inland lake
Secondary lowland forest 2-50 m.a.s.
Primary lowland forest 2-50 m.a.s
Hill forest 50-200 m.a.s
Hill forest over 200 m.a.s
Plantation
Total # of observations

V. caerulivirens
1 (33.3%)
1 (33.3%)
1 (33.3%)
3

High densities are also reached in brackish water river

deltas and Nypa swamps. Occasionally, individuals are
found further inland along freshwater streams, lakes and
smaller swamps where they occur syntopically with V.
caerulivirens. One individual on Bacan was encountered
at an elevation of more than 200 m in a small mountain
swamp. This species was only once (on Gebe)
encountered in sago swamps, despite several searches
in that habitat. They heavily utilize trees for basking and
nighttime refuges. Diet consists of crabs (Tanner, 1950)
as well as other suitable aquatic and terrestrial prey and
carrion. One individual was seen chasing aquatic prey in

V. rainerguentheri
17 (73.9%)
3 (13.0%)
1 (4.4%)
1 (4.4%)
1 (4.4%)
23

V. cf. salvator
3 (60%)
2 (40%)
5

a small stream, while another individual was observed

digging for food in a steep riverbank. On Obi and Gebe,
they were also frequently seen foraging around human
waste heaps at the margins of villages. The ecology of
V. rainerguentheri appears to be more generalized on
Gebe, where it occurs alone, from that of Halmahera
and Obi where they face competition from other monitor
species, probably indicating niche release (Weijola, in
prep.). The largest specimen measured was 133 cm in
total length, however, animals estimated to be nearer to
150 cm were observed on Obi.

19

WEIJOLA - MONITOR LIZARDS OF THE NORTHERN MOLUCCAS

Varanus yuwonoi
Distribution: The black-backed mangrove monitor was
only observed around the villages of Akesahu and Kao,
both in the western part of Teluk Kao (Kao Bay) on
Halmahera. The type series was collected near Jailolo
(Harvey and Barker, 1998). Interviews with locals and
animal dealers suggest that this species is widespread
throughout the island. Eyewitness accounts of V.
yuwonoi-like animals from Morotai, Bacan, and even
Obi exist, but still need confirmation.
Ecology: Varanus yuwonoi (Figs. 15 and 16) is a difficult
species to study. Despite frequent observations by local
hunters, specimens are very rarely encountered out in
the open. Considering the difference in size, which is
presumably also reflected in trophic level, this species
probably does not reach population densities as high as
V. caerulivirens. A less active mode of hunting (sit and
wait/ambush) may also result in the fewer encounters
with V. yuwonoi.
Traps used by locals specifically for this species are
always set up around megapode bird (scrubfowl) nests in

Fig. 15. Large V. yuwonoi. Akesahu, Halmahera.

inland forests. The unusual pigmentation of the tail, body

and neck breaks up the silhouette of the animal very well
as they lie on the forest floor. Considering that the smaller
and more predator-vulnerable species in the Moluccas
have not evolved such an elaborate camouflage, it may
have evolved rather as a means to escape detection from
potential prey. The head is comparatively powerfully
built and the teeth proportionately longer than any of
its close relatives (Fig.16), even in comparison to most
other more distantly related varanids. Considering
these facts, the author hypothesizes that V. yuwonoi is
an ambush predator, favoring sites in the forest such
as megapode nests, that are frequented by a variety
of potential prey items. It is uncertain whether large
individuals can kill adult scrubfowl, but the large nest
mounds attract a number of smaller birds such as pigeons,
and also lizards, snakes, and invertebrates, as well as
the frequently hatching megapode chicks. According
to local people, this species is also occasionally seen
digging into these nests in search of eggs. Additionally,
observations of intraspecific aggression within the V.
yuwonoi and V. doreanus clade (Ast, 2001) in captivity
would support a hypothesis that V. yuwonoi could be an

BIAWAK VOL. 4 NO. 1

20

Fig. 16. Head and teeth of V. yuwonoi at a trader in Dumdum, Halmahera.

ambush predator. With few exceptions (Sweet, 1999,
2007), widely foraging monitor (and other scleroglossan
lizards) species are not territorial; however, the two
species (V. scalaris and V. glebopalma) known to be
territorial are both ambush hunters. For a sit and wait
predator favoring prey hotspots, a certain degree of
territoriality could be expected to evolve.
The largest individual measured 146 cm in total
length; this specimen however did not appear to be fully
grown, and according to locals they occasionally grow
much larger.
Varanus cf. salvator
The distribution of V. salvator (Fig. 17) in the Moluccas
has been uncertain for a long time, probably starting
with what appears to be an erroneous record for
Halmahera in de Rooij (1915), widely cited since (for
example Bennett, 1995, Harvey and Barker, 1998). Obi,
Seram and Bacan were mentioned by Mertens (1930,
1942, 1959), and Buru by Bleeker (1857). Since recent
fieldwork, including this study, has not found support for
many of these localities, it is suggested that all except
Obi be omitted from the range of V. salvator.

Since the animals on Obi do not completely

correspond with any of the described V. salvator group
taxa, it is here employed with a cf. (meaning compare
with).
Distribution (in the north Moluccas): Obi Island,
probably also on nearby Bisa, but not confirmed.
Ecology: Five observations were made of this species
during two weeks on Obi. The sympatry between
members of Soterosaurus and Euprepiosauris is
interesting, since some authors have presumed that
they might be ecologically exclusive (Mertens, 1942,
Sprackland, 2004, Ziegler et al., 2007b). This proves
to be incorrect since this study confirms sympatry of V.
cf. salvator with V. rainerguentheri and V. caerulivirens
on Obi, as well as two other species in the Sula islands
(Weijola and Sweet, in press).
All observations of V. cf. salvator on Obi occurred in
coastal mangrove forest and periodically flooded littoral
vegetation (Table 2), where it is widely syntopic with V.
rainerguentheri. They most likely also occur in inland
forests and around freshwater streams side by side with
V. caerulivirens. On Obi, they appear less numerous than

21

WEIJOLA - MONITOR LIZARDS OF THE NORTHERN MOLUCCAS

Fig. 17. Varanus cf. salvator. Air Mangga, Obi.

V. rainerguentheri (5 vs. 22 observations), and similarly
on Sanana, many fewer V. cf. salvator were observed
compared to a sympatric member of the V. indicus
complex during the course of fieldwork.
Varanus zugorum
This is the only species which was not observed
during fieldwork. It appears to be almost completely
unknown by locals, and only two reasonably reliable
local eyewitness accounts were collected, in Akesahu
(lower western side of Kao Bay) and Labi Labi (northeast
Halmahera). Searches and interviews around the type
locality Pasir Putih were unproductive. The holotype
was collected in December 1980 by a villager, so further
details from the initial discovery remain unknown
(Adam Messer, pers. comm. 2008).
Discussion
Monitor communities and niche segregation by
sympatric species have been studied by several workers,
including Pianka (1994), Shine (1986), Sweet (1999,
2007) and Philipp (1999). Communities range from
simple one species systems up to as many as 11 species

in certain areas of northern Australia (Sweet, 2007).

These studies have shown that interspecific competition
is typically minimized by differing use of habitat,
different body sizes, and foraging mode. Additionally,
phylogenetic distance may aid in reducing niche overlap
in the most diverse communities of Australia and New
Guinea.
A community comparable to the one on Halmahera,
though slightly less diverse, was studied by Philipp
(1999) in West Papua. He investigated the habitat use
of three closely related species of the indicus-group (V.
doreanus, V. indicus and V. jobiensis), and concluded
that competition and interaction is largely avoided by
the use of different habitats and/or microhabitats.
This study found that on Halmahera and Obi,
V. rainerguentheri is similar in habitat use to that of
V. indicus on New Guinea (Philipp, 1999), while V.
caerulivirens appears to fill a similar niche to that of V.
jobiensis. Varanus yuwonoi overlaps widely in habitat
use with V. caerulivirens but grows significantly larger,
may use a different hunting strategy, and concentrate
on larger prey items. Thus there seem to be clear niche
separations in communities of V. indicus group animals
in Moluccan multi-species communities as well. The
ecological separation where members of Euprepiosaurus

BIAWAK VOL. 4 NO. 1

and V. salvator overlap is not equally obvious.

Acknowledgements- I especially wish to thank Dan,
Baba, Mayur, Mastur and Nelson for help in the field.
Sam Sweet, Daniel Bennett and two anonymous
reviewers provided valuable comments on an initial
draft. Travel support was provided by the National
Geographic Society, Svenska studiefonden, Nordenskild
Samfundet, Svensk-sterbottniska Samfundet and
bo Akademi University. I am also grateful to Mikael
von Numers and Erik Bonsdorff at bo Akademi, and
my parents, grandparents and friends for support and
encouragement.
References
Ast, J.C. 2001. Mitochondrial DNA evidence and
evolution in Varanoidea (Squamata). Cladistics 17:
211-226.
Bennett, D. 1995. A little book of monitor lizards. Viper
Press, Aberdeen. 208 pp.
Bleeker, P. 1857. Berigt omtrent eenige Reptilien von
Sumatra, Borneo, Batjan en Boero. Nat. Tijdschr.
Ned. Ind. 13: 470-475.
Bhme, W., H.-G. Horn and T. Ziegler. 1994. Zur
Taxonomie der Pazifikwarane (Varanus indicusKomplex): Revalidierung von Varanus doreanus
(A.B. Meyer, 1874) mit Beschriebung einer neuen
Unterart. Salamandra 30: 119-142.
Bhme, W. and T. Ziegler. 1997. Varanus melinus sp.
n., ein neuer Waran aus der V. indicus-Gruppe
von den Molukken, Indonesien. Herpetofauna 19:
26-34.
Edgar, P. and R. Lilley. 1993. Herpetofauna survey of
Manusela National Park. Chapter 8. Pp. 131- 141.
In: Edwards, I.D., A.A. McDonald and J. Proctor
(eds.), Natural History of Seram. Intercept Ltd,
Hampshire, England.
Hall, R. 1998. The plate tectonics of cenozoic SE Asia
and the distribution of land and sea. Pp. 99-131. In:
Hall R. and J.D. Holloway (eds.), Biogeography
and Geological Evolution of SE Asia. Backhuys
Publishers, Leiden.
Hall, R. 1999. Neogene history of collision in the
Halmahera region, Indonesia. Proceedings,
Indonesian Petroleum Association, 27th Annual
Convention & Exhibition, October 1999: 1-7.
Harvey, M.B. and D.G. Barker. 1998. A new species
of blue tailed monitor lizard (genus Varanus) from
Halmahera Island, Indonesia. Herpetologica 54(1):
34-44.

22

How, R.A. and D.J. Kitchener. 1997. Biogeography of

Indonesian snakes. Journal of Biogeography 4:
725-735.
Koch, A., M. Auliya, A. Schmitz, U. Kuch and W. Bhme.
2007. Morphological studies on the systematics of
South East Asian water monitor (Varanus salvator
complex): Nominotypic populations and taxonomic
overview. Pp. 109-180. In: Horn, H.-G., W. Bhme
and U. Krebs (eds.), Advances in Monitor Research
III. Mertensiella 16, Rheinbach,.
Koch, A., E. Arida, A. Schmitz, W. Bhme and T. Ziegler.
2009. Refining the polytypic species concept of
mangrove monitors (Squamata: Varanus indicus
group): a new cryptic species from the Talaud Islands,
Indonesia, reveals the underestimated diversity of
Indo-Australian monitor lizards. Australian Journal
of Zoology 57(1): 29-40.
Mertens, R. 1930. Die Amphibien und Reptilien der
Inseln Bali, Lombok, Sumbawa und Flores. Abh.
senckb. naturf. Ges. 42(3): 115-344.
___. 1942a-c Die Familie der Warane (Varanidae). Abh.
senckenb. naturf. Ges. 462, 465, 466: 1-391.
___. 1959. Liste der Warane Asiens und der
Indoaustralischen Inselwelt, mit systematischen
Bemerkungen. Senck. Biol. 40 (5/6): 221-240.
Monk, K.A., Y. de Fretes and G. Reksodiharjo-Lilley.
1997. The Ecology of Nusa Tenggara and Maluku.
Periplus Editions Ltd., Singapore. 966 pp.
Pianka, E.R., 1994. Comparative ecology of Varanus
in the Great Victoria desert. Australian Journal of
Ecology 19: 395-408.
Philipp, K. M. 1999. Niche partitioning of Varanus
doreanus, V. indicus and V. jobiensis in Irian Jaya:
preliminary results. In: Horn, H. G. and W. Bhme
(eds.): Advances in monitor research II. Mertensiella
11: 307-316.
Philipp, K. M., W. Bhme and T. Ziegler. 1999. The identity
of Varanus indicus: redefinition and description of a
sibling species coexisting at the type locality (Sauria:
Varanidae, Varanus indicus group). Spixiana 22(3):
273-287.
Setiadi, M.I. and A. Hamidy. 2006. Jenis-jenis
Herpetofauna di Halmahera.. Report for LIPI. Bogor,
Indonesia: Museum Zoologicum Bogoriense, Puslit
Biologi Lembaga Ilmu Pengetahuan Indonesia
(LIPI). 41 pp.
Shine, R. 1986. Food habits, habitats and reproductive
biology of four sympatric species of Varanid lizards
in tropical Australia. Herpetologica 42(3): 346-360.

23

WEIJOLA - MONITOR LIZARDS OF THE NORTHERN MOLUCCAS

Sweet, S.S. 1999. Spatial ecology of Varanus glauerti and

V. glebopalma in northern Australia. Pp. 317-366.
In: Horn, H.-G. and W. Bhme (eds.), Advances in
Monitor Research II. Mertensiella 11, Rheinbach.
Sweet, S.S. 2007. Comparative spatial ecology of two
small arboreal monitors in northern Australia. Pp.
378-402. In: Horn, H-G., W. Bhme and U. Krebs
(eds.), Advances in monitor research III. Mertensiella
16, Rheinbach.
Sweet, S.S. and E.R. Pianka. 2007. Monitors, mammals
and Wallaces Line. Pp. 79-99. In: Horn, H.-G., W.
Bhme and U. Krebs (eds.), Advances in Monitor
Research III. Mertensiella 16, Rheinbach.
Tanner, V.M. 1950. Pacific islands herpetology, No. 3.
Morotai island. The Great Basin Naturalist (Brigham
Young Univ.) 10: 1-30.
Voris, H.K. 2000. Maps of Pleistocene sea levels in
Southeast Asia: shorelines, river systems and time
durations. Journal of Biogeography 27: 1153-1167.
Yuwono, F.B. 1998. The trade of live reptiles in Indonesia.
Pp. 9-15 In: W. Erdelen (ed.). Conservation, Trade
and Sustainable Use of Lizards and Snakes in
Indonesia. Mertensiella 9, Rheinbach
Ziegler, T. and W. Bhme. 1999. Genital morphology
and systematics of two recently described monitor
lizards of the Varanus (Euprepiosauris) indicus
group. Pp. 121-128. In: Horn, H.-G. and W.
Bhme (eds.), Advances in Monitor Research II.
Mertensiella 11, Rheinbach.

Received: 20 February 2010; Accepted: 16 March 2010

Ziegler, T., W. Bhme and K. M. Philipp. 1999. Varanus

caerulivirens sp. n., a new monitor lizard of the V.
indicus group from Halmahera, Moluccas, Indonesia
(Squamata: Sauria: Varanidae). Herpetozoa 12(1/2):
45-56.
Ziegler, T., W. Bhme and K.M. Philipp. 2004. Varanus
caerulivirens. Pp. 161-164. In: Pianka, E.R. and
D. R. King (eds.), Varanoid Lizards of the World.
Indiana University Press, Indianapolis.
Ziegler, T., W. Bhme and A. Schmitz. 2007. A new
species of the Varanus indicus group (Squamata,
Varanidae) from Halmahera Island, Moluccas:
morphological and molecular evidence. Mitteilungen
des Museum fr Naturkunde Berlin. Zoologische
Reihe 83 (Supplement): 109-119.
Ziegler, T., A. Schmitz, A. Koch and W. Bhme. 2007. A
review of the subgenus Euprepiosaurus of Varanus
(Squamata: Varanidae): morphological and molecular
phylogeny, distribution and zoogeography, with an
identification key for the members of the V. indicus
and the V. prasinus species groups. Zootaxa 1472:
1-28.

Biawak, 4(1), pp. 24-25

2010 by International Varanid Interest Group

Varanus spenceri: A Last Supper of Grasshoppers (Acrididae)

PATRICIA A. WOOLLEY, MARTIN J. STEINBAUER and GREG MIFSUD1
Department of Zoology, La Trobe University
Bundoora, Vic. 3086
p.woolley@latrobe.edu.au
m.steinbauer@latrobe.edu.au
Greg.Mifsud@deedi.qld.gov.au
1

Present address: Department of Primary Industries and Fisheries, Toowoomba, Queensland 4350

Abstract: The stomach of a roadkilled Varanus spenceri contained many largely-intact grasshoppers,
together with the well-digested remains of a lizard limb, and nematodes, Abbreviata hastaspicula.
Spencers monitor, Varanus spenceri, is one of
Australias larger (ca. 1.2 m in total length), grounddwelling varanid lizards that is found on the Mitchell
grass plains of north-western Queensland and the Barkly
Tableland in the Northern Territory (Wilson and Swan,
2008). The diets of Australian varanids are not well
known, but they tend to swallow their prey whole or, in
larger species, in pieces that can be readily identified,
and they are capable of catching fast moving prey (Green
and King, 1993). Based on an analysis of the stomach
contents of preserved museum specimens of nine
species (not including V. spenceri), James et al. (1992)
found that most had generalized diets with lizards and
orthopterans being the most common prey items, but a
variety of other invertebrates were also taken.
Only three reports were found on prey taken by V.
spenceri and it appears that this species feeds on a broad
range of prey items including invertebrates, reptiles, and
mammals. Pengilley (1981) found isopods, grasshoppers,
reptiles (including single records of an unidentified
agamid and an elapid snake, Pseudonaja ingrami) and
unidentified mammals (probably Rattus villosissimus) in
the stomachs of eight specimens collected on the Barkly
Tableland from September to October, 1976. Valentic
and Turner (1997) found three lizards (two Ctenotus
joanae and one Tympanocryptis tetraporophora), and a
large roach (blattodea, approx. 30 mm long) too large

to have been secondarily ingested in the stomach of

a V. spenceri found injured on the Barkly Highway in
the Northern Territory, 32 km west of the border with
Queensland, in October 1996. Jackson and Lemm
(2009) identified the tail of an agamid lizard, Pogona
henrylawsoni, in the stomach of a road-killed V. spenceri
found on the Landsborough Highway, 110 km south of
Winton, Queensland in October 2008.
A fresh road-killed V. spenceri was collected near
Waterloo Plains (2057S; 14143E), approximately
33 km south of the town of Julia Creek, Queensland on
22 April 1995. The stomach contained a large number
of mostly intact, undigested grasshoppers (Acrididae)
as well as the well-digested remains of part of a limb
of a small lizard and numerous large nematodes. The
appearance of the acridids suggested that they were
the last food eaten before the death of the monitor. The
acridids, 47 in all, comprised one male Spur-throated
locust (Austracris guttulosa [Walker]), four male and
11 female Leaping Lagoonia (Lagoonia sp. 1) and 13
male and 18 female Speckled Yrrhapta (Yrrhapta striata
Sjstdet). Austracris guttulosa and Y. striata are fullywinged species whereas Lagoonia sp. 1 is brachypterous,
but known to be a powerful jumper. Rentz et al. (2003)
give details of size and seasonal occurrence for these
grasshoppers.
About 106 nematodes, all Abbreviata hastaspicula,

25

WOOLLEY ET AL. - VARANUS SPENCERI: A LAST SUPPER OF GRASSHOPPERS

were extracted from the stomach. This is the most

common species of Abbreviata in the drier inland of
Australia, principally in larger species of Varanus such
as V. gouldii, V. panoptes and V. spenceri (H. Jones, pers.
comm.).
These observations and those made by others on
the stomach contents of V. spenceri indicate that this
species, like the nine other varanids studied by James et
al. (1992), has a generalized diet.
Acknowledgements- We thank David Rentz for
confirming the identities of Lagoonia sp. 1 and
Yrrhapta striata, and H. Jones for identification of the
nematodes.
References
Jackson, R., and J. M. Lemm. 2009. Stomach content
note for a road-killed Varanus spenceri. Biawak
3(1): 18-20.
James, C.D., J. B.Losos and D. R. King. 1992.
Reproductive biology and diets of goannas
(Reptilia: Varanidae) from Australia. Journal of
Herpetology 26(2): 128-136.

Received: 28 January 2010; Accepted: 23 February 2010

King, D. and B. Green. 1993. Goanna: The Biology of

Varanid Lizards. New South Wales University
Press, Kensington. 102 pp.
Pengilley, R. 1981. Notes on the biology of Varanus
spenceri and V. gouldii, Barkly Tablelands,
Northern Territory. Australian Journal of
Herpetology 1(1): 23-26.
Rentz, D. C. F., R. C. Lewis, Y. N. Su and M. S. Upton.
2003. A Guide to Australian Grasshoppers and
Locusts. Natural History Publications (Borneo),
Kota Kinabalu. 419 pp.
Valentic, R.A. and G. Turner. 1997. Diet and
reproductive status of a road-kill Spencers
monitor Varanus spenceri. Herpetofauna 27(2):
43-45.
Wilson, S. and G. Swan. 2008. A Complete Guide to
Reptiles of Australia, 2nd ed. New Holland
Publishers Pty Ltd., Australia. 512 pp.

Biawak, 4(1), pp. 26-30

2010 by International Varanid Interest Group

Parthenogenesis in an Ornate Nile Monitor,

Varanus ornatus
JAMES HENNESSY
Reptile Village Zoo
Demesne Road, Gowran, Co.
Kilkenny, Ireland
E-mail: james@reptilevillage.net
Abstract: Parthenogenesis is documented in Varanus ornatus for the first time. A ten year old captive
female V. ornatus laid a clutch of 21 eggs in July 2008 without ever coming into contact with a male.
Two of the 21 eggs contained embryos. Information on the history and husbandry of the adult female
is given as well as details of the reproductive event.
Introduction
Parthenogenesis was recently described in two
varanid lizards, Varanus panoptes horni (Lenk et al.,
2005) and V. komodoensis (Watts et al., 2006). Here,
parthenogenesis is described for the first time in a V.
ornatus, maintained in captivity at the Reptile Village
Zoo in Kilkenny, Ireland.
Acquisition and Husbandry
A female V. ornatus (Figs. 1 & 2) was obtained from
a dealer in 1998 as an unsexed, wild-caught juvenile (11
cm snout to vent length [SVL]). The exact country of
origin is unknown.

Since January 2006, the female V. ornatus in question

has been housed in a glass-fronted enclosure of timber
construction measuring 150 x 120 x 140 cm (l x w x h).
The female was housed alone her entire life except for
three instances when attempts were made to house her
with another adult female V. ornatus in the collection. All
attempts at cohabitation were unsuccessful, with fighting
occurring within seconds of introduction. The enclosure
is furnished with an artificial rock background with
several basking ledges and large climbing branches. A 5
cm deep layer of bark mulch of is used for substrate and a
large water tub allows for bathing. A large cork bark tube
is provided for refuge. A 300 watt UVB basking lamp

Fig. 1. Female Varanus ornatus which reproduced by

parthenogenesis.

27

HENNESSY - PARTHENOGENESIS IN VARANUS ORNATUS

until test digging was first observed on 9 July, eight days

prior to oviposition. Test digging occurred in the rear
of the enclosure, away from the public-viewing side of
the enclosure. During gravidity, the females feeding
patterns did not change, and she consumed two large fish
three days prior to oviposition.
Incubation and Parthenogenetic Embryos

Oviposition

Previous V. ornatus clutches at the zoo were

discarded once retrieved. However, due to the recent
reports by Lenk et al. (2005) and Watts et al. (2006)
documenting parthenogenesis in Varanus, and the onsite availability of a large incubator, it was decided
to artificially incubate the eggs on this occasion. The
clutch was split up and placed in three plastic containers
each measuring 25 x 18 x 8 cm filled with dampened
vermiculite at a water:vermiculite ratio of 1:2 by weight.
The containers were then placed inside an incubator
converted from a refrigerator, heated by infrared lamps
on the bottom connected to a digital thermostat. A
small fan circulated air inside the incubator. The eggs
were incubated at 27 C and were inspected every other
day by opening each container and visually inspecting
each egg. Within a week, some of the eggs had begun
to deteriorate. Eggs which collapsed and had begun to
grow mold were removed and discarded. On 6 November
2008, after 113 days of incubation, seven eggs remained,
of which four appeared healthy. Assuming that all eggs
were non-viable after losing the majority of the clutch, it
was decided to discard all remaining eggs.
Beginning with the three deteriorating eggs, each was
dissected to check its contents. The three deteriorating
eggs contained solidified yellow material and showed
no signs of vascularization. Upon dissection of the first
healthy-looking egg, a partially-developed embryo
was discovered (Fig. 3). Coloration and patterning had

On 4 June 2008, the V. ornatus was given a routine

general health assessment, was weighed, and measured.
At this time, the female measured 52 cm SVL and 131
cm in total length (TL), and weighed 4.48 kg.
On 16 July 2008, the female deposited 21 eggs in
the corner of the enclosure, with oviposition lasting ca.
30 minutes. Eggs were retrieved once the female had
finished covering them with substrate. Defensive nest
guarding behavior was observed, however the female
appeared noticeably fatigued from oviposition.
Up until a few days prior to oviposition, the females
behavior did not change, exhibiting normal basking
behavior and activity levels. Gravidity was not suspected

Fig. 3. Parthenogenetic embryo, removed from egg.

Fig. 2. Profile of female V. ornatus.

fixed to the ceiling of the enclosure provides basking
temperatures of up to 45 C. An ambient temperature of
25 C (in winter) to 32 C (in summer) is maintained by
a panel heater controlled by a thermostat. As an adult,
the female has been fed weekly on a diet of weaned rats,
fish, chicks and eggs.
Both V. ornatus were proven to be females, evidenced
by egg production. The female in question laid 15 eggs
on 12 March 2007; the second female laid 12 eggs on
20 April 2007 and 15 eggs on 11 June 2008. Aside
from the brief, unsuccessful introductions to each other,
both females never came into contact with any other
individual. As a result, when eggs were laid by either V.
ornatus, they were assumed to be non-viable and were
immediately discarded.

BIAWAK VOL. 4 NO. 1

Fig. 4. Parthenogenetic embryo, dorsal view.

Fig. 5. Parthenogenetic embryo, ventral view.

28

29

HENNESSY - PARTHENOGENESIS IN VARANUS ORNATUS

begun to develop, and distinct banded markings on the

tail were visible (Figs. 4 & 5). All limbs and digits,
including the claws were formed. The embryo had
a SVL of ca. 5.3 cm and a TL of ca. 13.0 cm. Given
this discovery, the remaining three eggs were returned
to the incubator, where they were checked on a daily
basis for an additional 88 days. During this time another
egg began to deteriorate and was removed. Dissection
confirmed it was infertile.
On 2 February 2009, after 201 days of incubation,
the remaining two eggs started to deteriorate. One of the
eggs showed mold growth and was found to be infertile
upon dissection. The remaining egg, while slightly

beginning to smell of decomposition, appeared full and

turgid. Upon dissection, a dead, fully-developed embryo
was discovered (Figs. 6, 7 & 8). It had noticeably been
dead for some time and decomposition had begun to
set in. The yolk sac was fully absorbed and the embryo
appeared fully formed, apart from developing only three
digits on the front left foot. The embryo measured 9.3
cm SVL and 22.7 cm TL.
The same female laid another clutch on 2 August
2009. Eleven eggs were retrieved for incubation, though
at least three were eaten by the female before they could
be collected. All eggs began to decompose within the
first week of incubation.

Fig. 6. Full-term parthenogenetic V. ornatus embryo. Note the expended

yolk sack.

Fig. 7. Parthenogenetic
embryo, dorsal view

BIAWAK VOL. 4 NO.1

30

Fig. 7. Parthenogenetic
embryo, ventral view
Outlook
Since V. ornatus is now the third documented
species of Varanus to undergo parthenogenesis, with
previous reports documenting surviving parthenogenetic
offspring (Lenk et al., 2005; Watts et al., 2006), it seems
as though this reproductive mode may be more common
in captive varanids than realized. Further research is
needed to determine how widespread parthenogenesis is
in captive varanids, and whether or not it occurs in wild
populations. The two parthenogen have been preserved
and are currently held at the Reptile Village Zoo awaiting
future DNA analysis.

Received: 9 November 2009; Accepted: 15 March 2010

Acknowledgements- I would like to thank Phill Watts

for providing information for further research and Todd
Lewis for his support and guidance. I would also like to
thank Robert Mendyk for his guidance and suggestions
and two anonymous reviewers who helped improve this
manuscript.
References
Lenk, P., B. Eidenmueller, H. Staudter, R. Wicker and

M. Wink. 2005. A parthenogenetic Varanus.
Amphibia-Reptilia 26: 507-514.
Watts, P.C., K.R. Buley, S. Sanderson, W. Boardman,
C. Ciofi and R. Gibson. 2006. Parthenogenesis in
Komodo dragons. Nature 444: 1021-1022.

BOOK REVIEWS
A 21st Century Book Written for 20th Century Herpetoculture

Giant Lizards: The Definitive Guide to the Natural

History, Care, and Breeding of Monitors, Iguanas
and Other Large Lizards
ROBERT G. SPRACKLAND
TFH Publications, Inc. Neptune, New Jersey, USA. 2009. 335pp.
Hardcover. ISBN: 9780793805815

Reviewed by

ROBERT W. MENDYK
E-mail: odatriad@yahoo.com
One of my earliest introductions to herpetological
literature as a teenage reptile enthusiast was Robert G.
Spracklands (1991a) book entitled Giant Lizards, which
I came across one day in a local pet store. Back then, as
a budding hobbyist who knew absolutely nothing about
herpetological natural history, taxonomy, or husbandry,
I considered the book to be an excellent resource for
educating myself about lizards. But as time went on and
my interest in reptiles and knowledge of their biology and
husbandry grew, I began to notice a number of mistakes
in the book, whether they were outdated and potentially
hazardous husbandry advice, mislabeled photographs,
or photographs of taxidermied museum displays being
used to depict living specimens (see Fig. 1). Today, the
book has lost its value, but serves as a slight reminder of
my early years as a young reptile enthusiast.
Despite the poor composition of Giant Lizards, and
my disappointment with the authors (2001) book entitled
Savannah and Grassland Monitors (very critically
reviewed by Bennett, 2002), I was excited to learn
that TFH Publications would be publishing a new and
completely revised edition of Spracklands Giant Lizards
in 2009, this time incorporating written contributions
from several successful and esteemed lizard breeders.
Would this second edition be as disappointing as the
original, or would the author constructively learn from

and build upon its mistakes and errors, and criticisms

from others? Would the new book benefit from the input
of these invited authors? Subtitled as The Definitive
Guide to the Natural History, Care, and Breeding of
Monitors, Iguanas and Other Large Lizards, the revised
edition of Giant Lizards would surely have its work cut
out for it just to afford such a designation.
Reading this book has left me with a dissatisfying
sense of dj vu. Despite seeking to present new material
on the larger lizard species of the world, this revised
edition suffers from many of the same types of mistakes
seen in the original 1991 version, but also includes many
new mistakes and questionable material; all of which I
will discuss in detail below.
The Second Edition

31

Printed in a smaller size than the original version,

the second edition of Giant Lizards resembles a coffee
table book on account of its square-ish shape. Judging
the book by its hard cover, which depicts a beautiful
Varanus melinus, it would seem as though the publisher
invested significant time and resources into producing
this publication. Inside, the books pages are glossy
and of high quality, and printed in full color. The book
features more than 260 color photographs and hand-

BIAWAK VOL. 4 NO. 1

32

Fig. 1. A taxidermied display of Varanus komodoensis at the American Museum of Natural History, New York. A
photograph of this display is used on p. 52 in the original 1991 editon of Spracklands Giant Lizards to depict a live
dragon eating a wild boar. The same photograph also appears on p. 26 in Coborn (1997).
drawn illustrations, and 13 tables.
The book begins with a brief preface and introduction
which explain the motivation for the new edition, offer
acknowledgements, and a brief overview of the history
of herpetoculture. The book is then divided into two main
sections before concluding with a glossary, references,
resources section, and index. The first major section is
comprised of individual chapters dealing with a number
of topics relevant to the biology and husbandry of lizards
including anatomy, physiology, taxonomy, acquisition
and care, reproduction, veterinary care, laws, and
herpetoculture. The following section is comprised of
individual species accounts, divided into four chapters
highlighting giant members (species reaching over .9
m in total length; although some species below the .9 m
mark are also included [e.g., V. prasinus, V. keithhornei, V.
tristis] and some over the mark arent [e.g., V. reisingeri,
V. spinulosus]) of the Gekkota (1 species), Iguania (33

species), Scincomorpha (9 species), and Anguimorpha

(43 species). Each species account covers etymology,
distribution, maximum size, captive suitability, natural
history, captive care and breeding, and includes a list of
relevant publications. Included within some sections are
brief accounts on the captive husbandry and reproduction
of certain species written by invited authors. Bert
Langerwerf (Physignathus lesueurii; Ophisaurus
apodus), Gunther Kohler, A.J. Gutman and John Binns
(Ctenosaura), Robyn Markland and Chad Brown (V.
albigularis), Paul Rodriguez (V. salvator), and Ben
Aller and Michaela Manago (V. dumerilii) contribute
information based on their personal experiences with
certain species. The glossary defines a total of 90 terms
which also appear bold-faced within the text, and the
references section provides a bibliography of published
works cited throughout the book as well as sources for
further reading. The resources section lists a number of

33

MENDYK - A CRITICAL BOOK REVIEW OF GIANT LIZARDS

herpetological societies and informational websites, and

is followed by an index.
Comments
Knowing the Audience
Before I express my thoughts on this book, I feel it is
necessary to point out that since TFH Publications, Inc.
identifies itself as the source for authoritative books on
your companion animals as per the companys website
(http://tfhpublications.com), the target audience of this
book is most likely beginner and marginally-experienced
pet reptile-keeping hobbyists, probably those belonging
to younger age groups. These assumptions are further
evidenced by the colorful and playful tribal-like
borders, fonts, and artistic designs used throughout the
book, which I feel would deter more serious enthusiasts
and herpetological professionals from thumbing through
its pages or purchasing.
What immediately stands out when reading the first
chapter on lizard anatomy, physiology, and taxonomy, is
the authors textbook-like treatment of the material.
While this information may be useful and relatable to a
more informed audience, I feel that much of it is beyond
the comprehension level of the books major audience
despite the laymans definitions accompanying technical
terminology throughout the chapter. On page 29 the
author provides a list of bones found in the lizard skull.
While potentially useful to someone with an educational
backing in anatomy, I am doubtful that the average pet
lizard keeper will retain knowledge that the basioccipital,
epipterygoid and septomaxilla bones are part of the
upper skull, whereas the prearticular and surangular
bones are part of the lower jaw. Another example can be
found on the following page, where several bones of the
skull are individually sketched. Without a diagram in the
book referencing all of the bones found in an articulated
lizard skull, an illustration of a lone premaxillary is of
no use to an uninformed reader since it can be extremely
difficult to visualize where the bone is actually located
in the skull. Semi-technical discussions on circulatory
physiology, lung function, taxonomy and the rules
of zoological nomenclature, and chromosomal sex
determination systems (as well as CITES, and other
wildlife trade laws presented in a later chapter) are also
likely to escape the majority of the books readership.
Similar to my thoughts on certain sections of Bayless
(2006) book on V. exanthematicus (Mendyk, 2008), I get
the impression that the author attempted to create a semitechnical work for the wrong audience. The following

chapters read more in line with what would be expected

for a beginning keeper audience; for instance, on page
245, V. yuwonoi is said to resemble the Velociraptors of
Jurassic Park in temperament and behavior.
In addition to material that is likely beyond the
comprehension level of most readers, the book also
presents material which I find irrelevant to the subject
matter. The most obvious example is a full-page
table presenting the Greek alphabet system with a
pronunciation key (p. 303). Greek terminology, using
Greek letters, also appears in some species accounts
when considered relevant to a particular taxons
etymology. While the author states on pages 116-117
that he has received feedback from others interested in
herpetological etymology, I wonder how useful could
the Greek alphabet be to the average reptile keeping
hobbyist, and what proportion of readers will actually
take interest in such a resource?

Photographic Quality
One of the more upsetting yet perplexing aspects of
this book is its shortage of crisp and brilliant photographs.
I would estimate that more than 1/3 of the photos in the
book appear to be at least 15 to 20 years old, evidenced
by their dull, grainy and faded appearances, with many
of them terribly out of focus (e.g., p. 62, 173, 184, 188).
Additionally, I recognize several photos from previously
published pet hobbyist books and popular pet magazine
articles of the 1990s. The book does contain some nice
photography (mostly appearing on the title pages of
some chapters); however, the abundance of dreadful
photos detracts from any real enjoyment or appreciation
of the photography in this book.
I consider the overall photographic quality to be
highly unacceptable for any contemporary book of this
nature, and cannot believe that a publisher would print
seemingly decades-old, recycled, faded, and out of focus
material given the advent and widespread usage of digital
photography. Quality photographs of most lizard species
are no longer difficult to come by, thanks to online search
engines and photo hosting websites. Today, anyone
with a computer and working internet connection can
easily access high-quality digital images of reptiles from
around the world, and I am confident that most amateur
photographers would be delighted to contribute their
digital images to a book of this nature, gratis. While I
doubt these issues are the fault of the author, since the
publisher does have a well-known history of reusing
photographs in many of its reptile and amphibian titles, I
question whether those responsible for the photographic

BIAWAK VOL. 4 NO. 1

content of this book have ever heard of the internet, or if

they were just lazy?

Editing

34

Just like the original edition of Giant Lizards, this

version has its share of poor editing. In one example
in a sidebar discussion on page 25, 1980s Scotsman
should read 1780s Scotsman, otherwise the entire
section is historically incorrect and fails to make
any sense. On the title page of chapter two and in all
subsequent page headings in the chapter, acquisition
is misspelled as aquisition. Similarly, in all page
headings throughout chapter five, herpetoculture
is misspelled as heptoculture. Redundancy is also
an issue. For example, the change in coloration of V.
beccarii from hatchling to adult is mentioned twice in
the same species account (p. 231-232); similarly, the
long-distance eyesight of V. giganteus is also discussed
twice in the same account, in successive paragraphs (p.
271).
Editorial mistakes are not limited to proof reading
and fact-checking errors; poor layout and design are also
common throughout the book. The placement of sidebar
discussions, tables, and photographs in a chapter often
has nothing to do with their actual content. For example,
a sidebar discussion on page 32 discussing a veterinary
procedure for extracting blood appears within a section
on teeth, limbs, and tails. An illustration depicting head
pattern diversity in several varanid taxa appears in a
section dedicated to parietal eye and behavioral control
(p. 40). A sidebar discussion highlighting the homology
of the human hand, bat wing, and whale fin mysteriously
appears in a section on skin and scales (p. 45). While these
are just a few examples, several additional instances of
poorly-placed objects occur throughout the book which
might confuse, or be missed by readers.

according to its original description, which remains

the only published account on its occurrence to date,
V. yuwonoi is known only from inland forested areas of
Halmahera (Harvey and Barker, 1998). The author also
lists several preferred prey items for V. yuwonoi, yet no
dietary studies of wild or museum specimens have been
conducted on the species to date.
A statement which immediately caught my attention
is a claim on page 295 that V. salvadorii is often referred
to as death from above by local people because of the
lizards tendency to drop from a tree and kill hunting
dogs and to sometimes attack people. I am unaware of
any authenticated reports documenting such a behavior,
and while this claim is undoubtedly a case of folklore,
the way in which it is presented and implied in the text
will cause nave readers to believe that this is indeed a
natural habit of V. salvadorii.
In the natural history section of the V. salvator
species account on page 254, the author states that a
well-fed V. salvator may weigh 68 kg (150 lbs). This
is almost three times the maximum weight reported for
the species (25 kg/ 55 lbs) by Gaulke and Horn (2004).
Although specimens of V. salvator in captivity frequently
become obese, I am unaware of any authenticated report
or record which documents a captive specimen weighing
anywhere in the vicinity of 68 kg. A photograph in the
first edition of Giant Lizards (Sprackland, 1991a; p 127;
but also included again in the second [2009] edition, p.
104) purportedly depicts a 59 kg (130 lbs) V. salvator,
however judging by the size of the animal, this is most
likely a gross embellishment. Regarding size records,
the author also states that V. komodoensis holds the
maximum length record for any living lizard. While V.
komodoensis certainly attains longer average lengths
than any other species, the current authenticated record
for the longest lizard is held by a Sri Lankan V. salvator
which measured 3.21 m in total length (Randow, 1932).

Taxonomy and Natural History

Taxonomic Conventions

Natural History Information

Although this book is intended for pet reptile

hobbyists and is by no means a taxonomic treatise, there
are a number of taxonomic conventions used by the
author which are incorrect or invalid. I fear that these
misnomers will cause readers to accept invalid taxa and
confuse those who have read other accounts which use
different, or conflicting taxonomic conventions.
On page 231, the author refers to the taxonomy of
V. caerulivirens, V. cerambonensis, and V. juxtindicus
as controversial and suggests that they may represent
examples of normal variation within V. indicus; however,

There are a number of claims made throughout

the book pertaining to varanid natural history which
are dubious and lack literature citations to verify their
validity. I will focus on just a few that I feel are likely to
misinform or mislead readers.
On page 245, the author states that V. yuwonoi is
known to forage on the beach and swim in the ocean.
There are no published reports documenting a coastal
occurrence or sea-going behavior in this species, and

35

MENDYK - A CRITICAL BOOK REVIEW OF GIANT LIZARDS

no explanations or supportive evidence are given.

Several additional statements about the taxonomy of
the V. indicus complex are equally as dubious. Varanus
cerambonensis is also incorrectly identified throughout
the book as ceramboensis.
I am bothered by the authors use of Teris monitor
as one of the vernacular names of V. keithhornei. The
name Teris monitor originates from Spracklands
(1991b) description of V. teriae (named after his wife,
Teri), a species which had already been described six
years earlier as V. keithhornei by Wells and Wellington
(1985). Since V. keithhornei had taxonomic priority over
V. teriae, the latter was dropped and deemed invalid;
thus, so should its associated vernacular name.
The author fails to acknowledge V. kordensis as a
distinct taxon from V. prasinus, despite its elevation to
species status in 2002 (Jacobs, 2002) and widespread
recognition and usage of V. kordensis in current literature
(Jacobs, 2004; Eidenmller, 2007; Eidenmller and
Philippen, 2007; Bhme, 2003; Ziegler et al., 2007).
The author also rejects the validity of subspecies within
V. albigularis, citing a taxonomic revision of his own
published in a non-peer reviewed popular pet hobbyist
magazine (Bayless and Sprackland, 2000a,b).
Varanus ocellatus is surprisingly treated as a
valid taxon despite its long synonymization with V.
exanthematicus (Mertens, 1942a,b,c), also citing the
same hobbyist magazine revision as before (Bayless
and Sprackland, 2000a,b). Modern authorities follow
Mertens (1942a,b,c) treatment and fail to recognize V.
ocellatus as a valid species (Bhme, 2003; Pianka and
King, 2004).
The most problematic taxonomic treatment in the
book is the recognition of V. rubidus as a valid species.
Cited by the author in the rubidus species account, Storr
(1980) described rubidus as a subspecies of V. panoptes,
not a distinct species. Unless a mistake, the authors
treatment of rubidus most likely follows Wells and
Wellingtons (1985) controversial taxonomic revision of
Australian reptiles which raised rubidus to specific rank
without providing any supportive evidence or explanation
why. Due to a number of problems associated with Wells
and Wellingtons (1985) report, many of its proposed
taxonomic changes were never adopted, including
the elevation of rubidus. Interestingly, the author has
heavily criticized the Wells and Wellington (1995) report
in the past for its taxonomic questionability and lack of
adequate research when arguing for the conservation of
his V. teriae and the suppression of their earlier-described
V. keithhornei (Sprackland et al., 1997).

Photo Identifications
Much like the original version of Giant Lizards,
there are several misidentified species in the books
photographs. On page 14, a V. salvator is incorrectly
identified as V. niloticus. On page 241, a photograph
depicting a green and patternless specimen of a presently
undescribed species belonging to the V. indicus complex
(Bayless, unpub. ms.), is labeled as V. juxtindicus. On
page 252, a sulphur water monitor (Varanus salvator)
is identified as V. cumingi. On pages 254 and 255, three
of the four photographs are mislabeled or incorrectly
identified. Instead of reading V. s. marmoratus, V. s.
komaini, V. s. togianus, and V. s. salvator, the photographic
captions should read V. cumingi, V. togianus, V. nuchalis,
and V. s. salvator.
Captive Husbandry
I am very disappointed with the captive husbandry
and breeding information presented in this book.
Although the written contributions from the invited
authors are a welcoming addition since they present
useful and progressive information based solely on firsthand knowledge and experience, they quickly become
overshadowed by the authors own recommendations. In
addition to pushing many of the same outdated standards
and practices from some of his earlier herpetocultural
publications, there is also an abundance of contradictions,
inconsistencies and questionable material presented
throughout the text which will undoubtedly misinform,
mislead, and confuse readers, and in turn adversely
affect captive specimens.
Captive Suitability
For determining the captive suitability of a particular
species, the author has devised a numerical rating system
purportedly based on the disposition, hardiness, and
breeding potential of a species. A suitability rating of 1
represents a species considered to be extremely difficult
to keep, potentially aggressive, and unsuitable for
captivity, whereas a rating of 5 represents a beginnerslevel species easy to care for and not difficult to breed.
Suitability ratings for all varanid species presented in
Giant Lizards are given in Table 1.
I find most of the suitability ratings assigned to
varanids in this book to be preposterous, and question
the authors familiarity with the species in captivity.
Many of the ratings appear to have been randomly

BIAWAK VOL. 4 NO. 1

generated and are not at all representative of species

dispositions, hardiness, or ease of breeding in captivity.
For example, V. exanthematicus is given the highest
possible rating of 5, deeming it a worthy beginner-level
species. Since few V. exanthematicus live long lives
in captivity, with more specimens dying at the hands
of inexperienced beginner keepers each year than any
other varanid species, it cannot be considered a hardy
captive. Captive reproduction of V. exanthematicus has
also been extremely rare with respect to the sizeable
numbers of specimens being kept in captivity. Only a
few published reports on its successful breeding exist,
most of which report on single hatching events rather
than consistent and repeatable success; an indication
that V. exanthematicus is indeed difficult for most people
to breed in captivity. Given this overall lack of success
with V. exanthematicus, I fail to see how anyone can
consider it a species suitable for beginner keepers.
Despite clearly stating that V. niloticus is unsuitable
for private collections on page 226, the author assigns
it a suitability rating of 4 later on in the book. Of the
dozens of varanid species currently available in the pet
trade, I cannot think of many other species which are
less suitable for captivity than V. niloticus. Having only
been bred a few times in private collections, it is not
an easy species to maintain or reproduce in captivity.
Furthermore, V. niloticus attain very large adult sizes
(up to ca. 2 m) and can be extremely defensive, posing
serious safety risks to the keeper. Their inexpensiveness
and diminutive size as juveniles have led to countless
impulse purchases by inexperienced keepers over the
past several decades. As a result, most captive V. niloticus
do not survive to adulthood, and those that do are often
dumped off on rescue groups or sometimes released into
the wild when they become too large or unmanageable (V.
niloticus is now established in parts of southern Florida,
US [Enge, et al., 2004]). Suggesting that V. niloticus is a
semi-acceptable species for beginners by ranking it a 4
is ridiculous and also highly irresponsible.
Another rating which disagrees with the species
history in captivity is that given to V. nebulosus (V.
bengalensis nebulosus). Despite the author claiming it
to be a beginner-level species (5), captive breeding of
V. bengalensis nebulosus has been extremely rare with
few, if any records documenting successful reproduction
or long-term keeping of this species in existence. Also
attaining a large adult size (ca. 1.7 m), it is well-known
for its defensive nature in captivity (M. Cota, pers.
comm; M. Bayless, pers. comm.), and can pose serious
safety risks to keepers.
Most shocking of the rankings is that assigned to

36

Table 1. Suitability ratings of varanid species from

Spracklands (2009) Giant Lizards. 1 = least suitable for
captivity; 5 = most suitable for captivity
Taxon
albigularis
exanthematicus
ocellatus
niloticus
ornatus
griseus
yemenensis
beccarii
boehmei
keithhornei
macraei
prasinus
doreanus
finschi
indicus
jobiensis
melinus
yuwonoi
caerulivirens
cerambonensis
bengalensis
nebulosus
flavescens
cumingi
salvator
dumerilii
rudicollis
mabitang
olivaceus
giganteus
gouldii
panoptes
rosenbergi
rubidus
spenceri
glebopalma
tristis
mertensi
varius
komodoensis
salvadorii

Suitability Rating
3
5
4
4
3
4
3
3
3
3
2
2
2
3, 2
3
4
4
4
3
3
4
5
3
3
3
5
4,3
1
4
3
4
5
4
4
4
4
4
3
4
2
1

37

MENDYK - A CRITICAL BOOK REVIEW OF GIANT LIZARDS

V. olivaceus (4). As one of only two fruit-eating dietary

specialists within the genus, it has proven to be very
difficult to maintain and breed in captivity (Card, 1995a),
with just two live offspring produced in captivity to
date (Card, 1995b; M. Yuyek, pers. comm.). To assign
V. olivaceus, a large (ca. 2 m) and highly endangered
species notoriously difficult to keep and breed, the
same suitability rating as V. tristis, a small (to 80 cm),
easily-manageable species which has been successfully
kept in captivity for decades and bred to multiple filial
generations (Eidenmller, 2007; Husband and Bonnett,
2009), is ridiculous.
Inconsistencies in suitability ratings occur between
closely related taxa with identical or near-identical
husbandry requirements. For example, despite being
nearly identical in size, morphology, habit, diet,
disposition, husbandry requirements, and breeding
biology, members of the V. prasinus complex are given
different suitability ratings; V. macraei and V. prasinus
are given lower ratings than V. boehmei, V. beccarii,
and V. keithhornei. For reasons unknown, V. prasinus
is ranked as one of the lowest in the group in terms of
captive suitability, yet has been kept longer and bred
more frequently and consistently than any other member
of the complex, to multiple captive generations. Within
the V. indicus complex, V. doreanus is ranked two grades
below V. yuwonoi, yet both species attain similar sizes,
have similar morphologies and dispositions, require
similar husbandry parameters, and all known captive
specimens originate from the wild. Other members of the
V. indicus group, whose size, husbandry, and disposition
do not differ substantially from one another (e.g., V.
indicus, V. finschi, V. cerambonensis, V. caerulivirens,
V. jobiensis, V. melinus), are given a range of scores
which are not at all representative of their hardiness,
disposition, or history of reproduction in captivity.
While I do not think any large lizard species is suitable
for beginner keepers, which certainly undermines the
concept of this book, I do agree with the author in that V.
panoptes, V. gouldii, and V. tristis make hardy captives.
Since specimens of all three species maintained in
captivity outside of Australia are captive bred, they tend
to be hardier, have generally calmer dispositions, and
are more likely to reproduce than wild-caught species. I
also agree that V. salvadorii makes a terrible captive, but
since the authors rating for this species is based on its
potential dangerousness, why not extend the same rating
to V. komodoensis and other truly giant species such as V.
salvator, V. niloticus, or V. giganteus which are equally
as formidable?
These rankings are also disappointing because they

promote and endorse the wild-caught reptile trade. Since

successful captive breeding has been absent or extremely
limited in V. exanthematicus, V. niloticus, V. jobiensis,
V. melinus, V. yuwonoi, V. bengalensis, V. dumerilii,
V. rudicollis, and V. olivaceus (level 4 and 5 species
according to the author), specimens in the pet trade are
almost entirely of wild-caught origin. Promoting these
species as easy or relatively-easy to keep will encourage
inexperienced, beginner hobbyists to purchase wildcaught specimens, further sustaining the wild-caught
trade. I cannot see how anyone can look at the numbers
of wild-caught (or captive-hatched, which is even more
ecologically-destructive) V. exanthematicus imported
into the United States alone each year (ca. 25,000 in
2007, ca. 23,000 in 2008; CITES trade database) and
fail to see a problem with recommending the species to
inexperienced keepers.
If these wild-caught species were identified and
promoted as poor, unsuitable captives, the demand for
specimens would decrease, and fewer animals would
have to be removed from the wild each year to supply
the pet trade. Beginner keepers should be introduced
to captive-bred species which are hardier, and where
deaths caused by keeper inexperience will not affect
wild populations. Wild-caught species should be left
to experienced, dedicated keepers and breeders who
are more likely to succeed in establishing captive-bred
lineages.
Husbandry Recommendations
Prior to reading this book, I assumed that 21st
century herpetoculture had advanced well beyond the
primitiveness of feeding processed meats and dog food
to captive reptiles, and that those days were now a
thing of the past. I was wrong. On numerous occasions
throughout the book, the author condones feeding beef
(p. 54, 187, 241), cooked turkey sausage (p. 133, 139.
201, 211, 219, 234, 241, 246, 260, 270, 274), and dog
food (p. 133, 187, 196, 201, 208, 238, 246) to lizards.
Also advocated is the feeding of bananas to V. prasinus,
V. macraei, and V. boehmei. Improper and lazy dietary
items such as these can easily become adopted by
beginner hobbyists who dont know any better, and can
adversely affect the health of captives. I am not aware
of a single report of long-term keeping or reproductive
success which documents or advocates the feeding
of beef or dog food to varanids. These are potentially
hazardous, unnatural dietary items which mainstream
herpetoculture has done away with many years ago, for
good reasons.

BIAWAK VOL. 4 NO. 1

The authors support for a number of unnatural

substrates in this book is equally as appalling. Although
he does recommend deep soils for V. albigularis, V.
exanthematicus and V. griseus, which would seem to be
a progressive departure from some of the advice given
in his earlier works, these suggestions are negated by
his endorsements of bare wood, glass and concrete
floors, wood shavings, shredded newspaper, rabbit food
pellets (alfalfa), paper towels, and brown butcher paper
as acceptable substrates elsewhere in the book (p. 61).
These suggestions conflict with advice given by highly
successful long-term varanid keepers and breeders such
as Vincent and Wilson (1999), Eidenmller (2007),
and Husband and Bonnett (2009), who understand
that terrarium substrates serve more purposes than just
collecting fecal material. Unnatural substrates serve
absolutely no benefit to the physiological or behavioral
requirements of lizards and should have no place in
herpetoculture.
Another major problem with the books captive
husbandry information is its lack of consistency between
species accounts. For some accounts, recommendations
for enclosure sizes are outlined, but for others they
arent. Suggested dietary items are given for some taxa,
but not for others. Incubation temperatures, humidity
levels and durations are given for some, but not for
others. What I especially find strange is why many
closely related taxa, such as members of the V. prasinus
complex, have different husbandry guidelines. Why is
the recommended diet for V. macraei different than V.
prasinus, considering their near-identical morphological
and ecological similarities? Another example is the
difference in feeding frequencies recommended for V.
cumingi and V. salvator, two very similar species which
were once considered the same species. Why is V.
cumingi to be fed daily, but V. salvator just twice weekly?
Inconsistencies and contradictions like these occur for
many other species in the book (not just varanids) and
give the impression that these sections were hastily
assembled in a random, unorganized manner. It also
begs the question of where is the information coming
from?
Many of the species accounts fail to cite or
incorporate important information from published
husbandry and breeding reports, and instead present
questionable material from outdated or unknown
sources. For example, despite a number of detailed and
informative publications existing (e.g., Biebl, 1993;
Dedlmar, 1994; Bosch, 1999; Polleck, 2004; Baldwin,
2006), the species account for V. prasinus does not cite
a single report of successful long-term keeping and

38

breeding of the species. Instead, the author refers readers

to dated articles of his own on the keeping of V. prasinus
in captivity (Sprackland, 1989, 1991c). Even when
successful published accounts are cited for a species, the
information is sometimes mixed together with and skewed
by questionable material from other, unknown sources.
An example of this can be found in the V. boehmei species
account (p. 232), which cites the only published report
on the successful husbandry and breeding of the species
(Reisinger and Reisinger-Raweyi, 2007). Nowhere in
Reisinger and Reisinger-Raweyi (2007) do the authors
report or advocate the feeding of bananas, yet they are
included by the author as a suggested dietary item in the
books species account. Another example demonstrating
a poor grasp of herpetocultural literature can be seen in
the V. macraei species account, which states that there
are no published reports on captive bred offspring of V.
macraei, yet two such reports (Jacobs, 2002b; Dedlmar,
2007) are cited in the accounts references section.
Occasionally, the author compares the husbandry
requirements of one species to a non-related species
originating from an entirely different environment. For
example, little information is given on the husbandry
of V. varius (p. 287); instead, the author suggests
that its husbandry should mimic that of V. salvator.
Varanus varius and V. salvator inhabit vastly different
environments, have different habits (e.g., V. salvator is
semi-aquatic; V. varius is semi-arboreal), and differ in
their reproductive biology (e.g., V. salvator often nests
in the ground; V. varius in termite nests). In another
example, the author recommends that the husbandry of
V. nebulosus (V. bengalensis nebulosus), an inhabitant
of open forests and disturbed environments of southeast
Asia, should be the same as that for the African grasslanddwelling V. albigularis.
On more than one occasion, the author recommends
dangerously high ambient temperatures which can be
deadly to any reptile maintained under such conditions.
For example, on page 271, he claims that captive V.
giganteus do best if kept at temperatures above 43.3 C
and given several options to escape the heat. In virtually
every conceivable captive situation, particularly indoor
enclosures, cool areas to escape extreme heat would be
extremely difficult if not impossible to provide if the
enclosures ambient temperature was being maintained
near or above 43.3 C. Unless the author is wrongly
referring to basking temperatures of 43.3 C, extended
exposure to these excessive ambient temperatures
will quickly kill a reptile. Similarly, on page 249 the
author recommends seasonal increases in daytime
temperature to 39.7-45 C for V. bengalensis. Like

39

MENDYK - A CRITICAL BOOK REVIEW OF GIANT LIZARDS

the excessive ambient temperatures recommended for

V. gigatneus, these markedly high temperatures will
quickly kill captive reptiles when cool temperatures
are unable to be provided. Beginner keepers can easily
misunderstand the concept or importance of cool refuge
sites when providing hot ambient temperatures, and will
quickly overheat and kill their captives if they follow
these suggestions.
Many of the books photographs depict outdated
husbandry practices and inadequate enclosure designs.
To use the expression, a picture is worth a thousand
words, I fear that beginner hobbyists flipping through
this books pages will see some of the pictured lizard
enclosures, substrates, hide spots, and furnishings, and
adopt them for their own captives. There are several
photos which depict lizards housed in screen cages
or enclosures with large wire ventilation areas (p. 59,
62,108, 239, 247). Since most keepers do not live in
tropical regions with consistently high humidity levels,
these types of enclosures promote low humidity levels
and can cause chronic dehydration, related illnesses, and
eventually death. Although the author briefly discusses
the importance of proper humidity levels and hydration
in the text, he does not address the humidity issues
wire screening will cause. Moreover, the photos of
inadequate enclosures are more likely to be noticed by
readers, and can be devastating to the health of captives.
On page 61, a young V. komodoensis is pictured in its
enclosure. The images caption discusses the importance
of shelter in reducing stress levels in varanids; however,
the half log-type hide depicted in the photo is too large
and vacuous to provide any real sense of security for the
animal. Inappropriately-sized hide areas occur in several
photographs (p. 60, 65) and may give readers the wrong
impression about what suitable refugia should look like
and provide.
Non-Varanid Material
Though I have clearly focused my attention on the
varanid-related content of the book since this review is
appearing in a varanid-specific publication and because
its varanid content makes up almost half of the books
species account section, I did find some errors in nonvaranid related sections as well. One of the more
noticeable mistakes occurs in the chapter on agamids
where the author repeatedly refers to Hydrosaurus
pustulatus incorrectly as pustulosus (p. 135-137).
Hydrosaurus pustulosus is also claimed to be the most
heavily-exported member of the genus based on United
States CITES records (p. 136); however, no members of

Hydrosaurus are listed in any of the CITES appendices,

therefore, the source for this statement is incorrect and
the claim dubious. Many of my concerns regarding
the husbandry advice given for varanids also apply to
other taxonomic groups discussed in this book. Given
the poor quality and accuracy of information in sections
pertaining to Varanus, I have no reason to believe that
other areas of the book do not suffer in similar ways.
Conclusions
To be blunt, I feel that Giant Lizards 2nd ed. is
one of the worst books of the last two decades to
cover the natural history and captive husbandry of
varanid lizards. My criticisms outlined in this review,
although numerous, do not represent frivolous or trivial
mistakes, but serious flaws which affect the accuracy,
credibility and educational value of the book, and are
indicative of its overall quality. Its poor composition,
careless mistakes, informational inconsistencies and
contradictions, unsupported statements, questionable
taxonomic conventions, dreadful photography, and
appalling captive husbandry advice render this book
the antithesis of anything worthy of being considered a
definitive guide.

Of my many criticisms, I consider the biggest
disappointment to be the information and advice given
on captive husbandry and breeding. From the information
presented in this book, I am convinced that the author
has very little, if any practical experience with varanids
in captivity, much less the 41 species he has provided
husbandry and breeding advice on. Admittedly having
no experience or interest in breeding lizards (p. 7), the
author clearly lacks the qualifications necessary to be
offering recommendations on their long-term care and
reproduction in captivity.
Although lacking the product placements which
were rampant throughout the first edition that
undoubtedly helped boost reptile product sales in pet
shops worldwide, the second edition of Giant Lizards
is destined to become a new personal favorite of pet
shop owners if their customers can afford its $79.95
USD suggested retail price (I have been told this price
has recently been lowered to around $50 USD). Not
only does it encourage readers to purchase wild caught
species, which have a much higher retail markup value
than captive-bred species, but inexperienced keepers
are also likely to return to the pet shop to purchase a
replacement animal once his or her current specimen
dies as a result of some of the poor husbandry advice
outlined in this book. Lastly, one mustnt forget the

BIAWAK VOL. 4 NO. 1

books dietary recommendations of canned dog food for

lizards, which will also require returning trips to the pet
shop.
Sadly, instead of improving herpetoculture, as any
new book on the subject should, Giant Lizards sets varanid
keeping back more than a decade with its outdated and
potentially harmful husbandry recommendations, some
of which will kill captives if adopted. The author ignores
many of the significant advancements and breakthroughs
made in varanid herpetoculture over the past decade
which have enabled specimens to live longer, healthier
lives and possibly reproduce in captivity. This is now
the authors third poorly-written book reporting on the
captive husbandry of varanid lizards, which I consider to
be a great disservice to the progression and advancement
of herpetoculture.
Due to its poor educational value and overall
quality, I cannot in good conscience recommend this
book to anyone, not even for its photographs, as I
have done for others in the past (Mendyk, 2008) since
a sizeable percentage of them are poor in quality. For
current and progressive information on the keeping and
breeding of varanids in captivity, I recommend Monitor
Lizards: Natural History, Captive Care and Breeding
by Eidenmller (2007), The Savannah Monitor Lizard:
The Truth About Varanus exanthematicus by Bennett
and Thakoordyal (2003), and Keeping and Breeding
Australian Lizards, edited by Swan (2007). For
detailed information on the biology and natural history
of varanids, I recommend Pianka and Kings (2004)
Varanoid Lizards of the World.
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Publications, Neptune, New Jersey. 128p.
Bayless, M.K. and R.G. Sprackland. 2000. The
taxonomy of Africas savannah and cape monitor
lizards, part 1. Reptiles 8(6): 76-85.
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cape monitor lizards, part 2. Reptiles 8(7): 40-47.
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RECENT PUBLICATIONS
2009
Bowker, R.G., C.L. Wright and G.E. Bowker. 2009.
Patterns of body temperatures: is lizard
thermoregulation chaotic? Journal of Thermal
Biology 35(1): 1-5.
Das, A., U. Saikia, B.H.C.K. Murthy, S. Dey and
S.K. Dutta. 2009. A herpetofaunal inventory of
Barail Wildlife Sanctuary and adjacent regions,
Assam, north-eastern India. Hamadryad 34(1):
117-134.

Doody, J.S., J. Roe, P. Mayes and L. Ishiyama. 2009.

Telemetry tagging methods for some freshwater
reptiles. Marine and Freshwater Research 60(4):
293-298.
Flesch, J.S., M.G. Duncan, J.H. Pascoe and R.C.
Mulley. 2009. A simple method of attaching GPS
tracking devices to free-ranging lace monitors
(Varanus varius). Herpetological Conservation and
Biology 4(3):411-414.
2010

Christy, B. 2010. The kingpin: an expose of the worlds

most notorious wildlife dealer, his special
government friend, and his ambitious new plan.
National Geographic 217(1): 78-107.
Douglas, M.E., M.R. Douglas, G.W. Schuett,
D.D. Beck and B.K. Sullivan. 2010. Conservation
phylogenetics of helodermatid lizards using
multiple molecular markers and a supertree
approach. Molecular Phylogenetics and Evolution
55: 153-167.
Ficetola, G.F., A. Crottini, M. Casiraghi and E. Padoa-
Schioppa. 2010. New data on amphibians and
reptiles of the northern areas of Pakistan:
distribution, genetic variability and conservation
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Janzen, P. 2010. Herping the Pearl. Reptiles 18(4):
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Sprackland, R.G. 2010. Savannah savvy. Reptiles

18(2): 32-41.
Okafor, A.I. 2010. The influence of body temperature
on sprint speed and anti-predatory defensive
responses of the North African monitor lizard,
Varanus griseus. African Journal of Biotechnology
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Pare, J. and A. Lentini. 2010. Reptile geriatrics.
Veterinary Clinics of North America: Exotic
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Rothschild, B. 2010. Macroscopic recognition of
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Smith, J.G., A.D. Griffiths and B.W. Brook. 2010.
Survival estimation in a long-lived monitor lizard:
radio-tracking of Varanus mertensi. Population
Ecology 52(1): 243-247.

42

Varanus albigularis. Tarangire

National Park, Tanzania. Photograph by Daniel Virella.