NOTES ON TIIE BIOLOGY OF VARANUS GRISEUS KONIECZNYI ...

NOTES ON TIIE BIOLOGY OF VARANUS GRISEUS KONIECZNYI /MERTENS
SAURIA: VAP*ANIDAEI
wALTm. AuFFsNBERc2, Harrpzun REHMAN, FErnrcna IFFAT Ar\tl zatgnepm.vnnt3
QVit h four text-fi g ures)
The preserrt surdy documents several aspects of thebioiogy of.Varanus griseu.s konieczzyiin Pakistan, Adequate
material now available makes possible a more detailed descriprion of the molphological features and rhe geographic
disrribution of this species than was prcviously possible. The annual reproducrive and "Mo*it ul fur "y"t"r "]e oirtit"a
and the common prey organisms idenrified.
hr"rR,oouc'[oN
This srudy was undertaken as pafi of an extensive
research programme involving the varanid
lizards of India and Pakistan. It is the second of a
series of publications intended to make theresults of
this work available to biologists. With the exceplion
ofa few casual nores (Corkhilt 1928), literature contains
noihing regarding the bioiogy of this subspecies
(a significant unpublished thesis research
study on food and burrow use was conducted by
Dave (1961) at rhe University of Rajasthan (see
below). Severalof &e earlierpublicarions {i.e. Smith
i935) that deal with tlle eastern populations of
Varanus griseus (now considered to represent the
distinctive V. g. koniecznyi) include behavioural and
other data rclating to more western populations now
considered as distinct geographic races, so that it is
not clear what information pertains !o the eastern
race and what does not.
Varanus griseas (desert monitor lizard) is disu'ibuted
from northern Africa to north--central India-
Within this large area three geographic races are
recognized V. g. griseusfiom Africa trc approximately
eastern kan, Z g. caspius from t}te westem shores
of the Caspian Sea, Turkman and thelranianPlateau,
easfward through Afghanistan and Baluchistan to
western Sinkiang Province, China; and V. g.
koniecznyi from cenral Pakistan to north-+entral
India.
Varanus g. kaniecznyi was described by Mertens
in 1954; the rlpe localiry is Khorangi, near
Karachi, Pakistan. ia addirion ro rhose morphologi-
lAccepred Seprember I 988
{he Florida Srate Museum, University of Florida, Cainesville,
Florida 32611, U.S.A.
'A1l remaining authors Zooiogical Suwey Depr, Block 61,
Pakistan Secretariat, Karachi, Pakisran.
cal features he iisted as distinguishing it from irs
closest geographic conspecific V. g. caspius,heincluded
a few other localities. To these he added
several more in 1969. Minron (1966) published a
few natural history notes, added still more localiries
in southern Pakistan, and funher characterized the
subspecies on the basis of a few additionai morphological
features. Our studies were primarily conducted
to befter understand the biology of this poorly
known subspecies. The study was intended to
suppiy baseline information on distribution, morphology,
reprod.ucfion and nuririon as an aid to
developilg a conservalion programme for this (and
orher) monitor species in southern Asia.
The following data were obtained as a result of
field wort conducted from 1984 through most of
1987 in both countries. Addirionally, importanr
museum specimens were examined in both these
countries and in Europe and the United States. Total
field time was approximately 22 months (India 10,
Pakistan L2). Data were obtained from 150
specimens (52 in museums, 98 in rhe field). Al1
measurements of rotal length (TOL), tail length (tL)
and snout-vent length (SW) were made to the
closest mm; all internal measurements (testes, ova,
etc.) weremade to the closest 0.1 mm; all weights to
*re closest 0.1 g. l
REsuLTs
Distribution: Fig. 1 shows the published literature
records as well as those Pakistan localities
demonstrated as possessing Vararuts giseus during
the present sfudy. These data make it quite clearthat
the subspecies V. g. kaniecznyi is restricted to what
is best described as the historic and the presenr Indus
River Valley (seeFig-g,However, within this large
area, it is more or less restricted to sandy tracts. This
habital preference' has already been rnentioned by

BIOLOGY OF VARANUS GRISEUS KONMCffYI
Fig. 1. I-ocality records for Varanus griseus koniecznyi; literature tecords hollow dots, examined speci-ens solid dots. Shaded area
repres€nts what we believe to be the entire range of the subspecies. The zone of intergradation with Y g. caspir.rs along the western
border is nol yet clear (see texr). Only one specimen is considered an intergrade (half solid dor). The tningles represen! the closesl
localities known at present for Varanus g. caspius.
Minton (1966); it'is rarely found in clay deserts.
Common plants noted in areas where tr4 g. koniecznyi
were seen during this study we Acacia jacquemontii,
Prosopis cineraria, Salvadora oleoides, Capporis
aphylla, Leptadenia pyrotechnica, ffid Colligonum
polygonoides.
Localiries from which this subspecies are now
known and the data source follow:
iliDiA. Uttar Pradesh Prov., Sikaiidra (this
study), nearAgra (Carlieyle 1869); Dat'albagh (this
sludy), Agra (Boulenger 1885, basedon Carlleyle?),
Delhi (Jerdon i870), Oudh (Murray iB84);
Maharashtra Prov., Narsingarh (Smith 1932, see
text, locality questioned). Gujarat Prov., Surat,
(Gleadow 1905), Deesa (ZSI and Smith 1932);
Kutch (Stoliczka 1872). Haryana hov., Ambala;
Rajasthan hov., Bikaner, Barmer and Pirgal (Dave
1961), Pokaran (this study), Jodhpur (ZSU, Smith
1932 utd this study), Jaipur (rhis study).
PAKISTAN. Baluchistan Rov., Gwador DisE.:
Ormara (this study); Quetra Distr., Quetta (ZSD);
Las Bela Prov., Bela (this study), Goth Mauladad
(Mertens 1969), Uthal (ZSM), Panjgur Disrr.,
Panjgur; Punjabkov., Jhellum Distr.: Goolpour, 9.6
km Pindo Dadau Khan (this study); Jhang Disn'.:
Jhang (UMMZ), Rabwah (this study), Bhravi Desert
(il{inton 1966); Sargodha Distr.: Khepra, nr" Jhellum
River (this study); Lahore Distr.: Lahore (Jerdon
1870): 32krn SE I-ahore; Kolakhatai; Narang
Mandi; Sri Rampura; Changa Manga (all this study),
Mianwali Disr.: Salt Range (Iheobald 1868). Sind
Prov., Badin Distr.: Badin (I4CZ and Mertens 1969),
Thar Parkar Distr.: (Smith 1932), Islamkot @SD),
Nagar Parkar (this study), Umarkot (ZSD and this
study); Karachi Distr.: Karachi (Minton 1966 and
many museum collections), Dabiji (Mertens 1969),
Clifton (lt4inton 19ffi), 6.4 km E landhi ([4inton
1966), Khff Centre nr. Hab Dam (this study), Sanghar
Distr.: Sher Khadra (this study); Thatta Distr.:
(Mertens 1969, Jhermck (Mertens L969),
'Jati
Jungshahi (Mertens 1969 and this study);4.8 km NE
Gharo @linton 1966); llyderabad Disf.: nr. Bhalori
([4inton 1966), MirpmKhas (ZSD); Dadu Distr-: nr.
-
Thana Bulla Khan (ZSD, Minton 1966 and this
study).
The locality Narsingarh (Smith 1932) requires
verificalion, foi'it is out of the expected mnge on
zoogeographical grounds. In the most eastsrn parts
of this subspecies range Nikolsky (1907) reports
Varanw griseru (subsp.?) from SE Iran; but the subspecies
represented is probably V. g. caspius on the
basis of Mertens's record from kanshali-(1956). The

.)e
western limits of I4g. koniecznyi is apparently at the
edg0'of the Iranian Plateau (i.e., the Sulaiman,
Bruhai and Kirthar Mountain Ranges in cenfrai
Pakistan. At the southeastern limit the subspecies
ranges into, but not beyond, the GujaratPlains (Suw.
Indja Maps, pl. 41), and eastward to, but apparently
not byond the Ganga-Jamuna Doab. I have seen a
specimen (not captured) from the sandy river bed
near Firozabad that I believe represents this species.
If corre.ctly identified, it is the most eastern locaiity
lanown.
Systematics: At an early date the disrinctive coloration
and pattern of the Yaranus griseus populations
from India had already been reported in the literature
(I{ardwicke and Gray 1827, Anderson 1898). In
1942, lv{ertens suspected that the Indian and Pakistani
specimens represented an undescribed race on
the basis of its colour pattem, but had too few
specimens for study to be certain. Additional Pakistan
material was made available through colleclions
made by Mr Konieczny. Mertens described the subspecies
V. g. koniecznyl in the collector's honour in
1954. Unforunately, Mertens was unable to define
the geographical limis at the western edge of the
range, for at that time there were no Baluchi
specimens with precise locality data.
Today, the situation is hardly better, though
Minnn (1966) reported a specimen referred to V. g.
caspins from Chagai District, 17.6 km NW Nushki.
It is also presumed that the specimen reported by
Wall (i912) from Chitral Distr., N.W.F.P., was also a
member of this race. Thus systematists believe the
populations along the Afghani-Baluchi border are
referable n V. g. caspius.It is presumed that those
populations between these and those examined from
the most northwestern localities in the Indus Valley
trrg. 1) are intermediate, though this has not been
proven with the material at hand. Even less is lnown
about the subspecfi c allocation of populations along
the Irani-Baluchi border. Specimens from the middle
and eastem part of the Mekkran Coast (Ormara,
Uthal, and Beia) are clearly assignable to V.g.
koniecznyi, and show no evidence of intergradation.
However, a single adult from Panjgur, Baluchistan
(ZSM 161i1985) is intermediate in body and tail
colour pattern between V.g. knniecznyi and ltg.
caspius. The closest undoubted V. g. griseus records
are in central Iran (fuck 1971).
l
JAURNAL, BOMB,4Y NATURAL HIST. SOCIETY, Vol.87
A good black-and-white photograph of
Varanus griseus kaniecznyi is provided by Minton
(1966) and a coloured one by Auffenberg (1986).
The photograph by Pilleri (1970) is not of this
specie:s as stated, but of 14 bengalensis bengalensis,
as frst noted by Mertens (1971). Detailed descnptions
of material then available are provided by Mertens
(1954) and Minton (1966). The following short
description is based on a much wider seieclion of
specimens available to us in this study:
Tongue long, slender, bifid and renacnble into
a sheath, Head moderately broad, with a pointed
snout, External nares, iocated near the eye (2840 Vo
of the distance from the nostril to tip of the snout).
Tail rounded in cross s€ction throughout most of its
length,longer than snout-vent, distance {IW-L28 Eo
of SW). Head scales small, polygonal; mid-dorsal
body scales small (108-139 scale rows around
body), granular above, squarish ventrally.
Dorsai head colour dark grey to black, lighter
in the young. Body duli yellow to light grey,
speckled with dark grey to black; 3-4 dark grey,
greenish grey (to dark brown or black in young)
cross bands on trunh bordered by and sometimes
enclosing white to yellow spots. This pattern is
somelirnes obscured by a long triangular dark grey
to black patch that extends from the top of the head
over the dorsal part of the neck onto the shoulders,
somelimes farther posteriorly. This "cape" is best
developed in older individuals, especiaily from the
Indian side of the Thar Desgrt (photograph in Auffenberg
198Q. There is always a dark brown to black
stripe extending from the canthus through the ear
opening onto the neck and a shorter, narrower one
from behind the eye. The base and proximal two
thirds of the tail are often crossed with from &-12
greyish bands; distal third to one haif black with a
white rip. The limbs are greyish with yellow spots
and the venfal surfaces are white, often with dark
speckling on the throal
Corkhill (1%8) reports that during early summer
the bellies of males have a pinkish cast Though
this is probably cdnect, we were not able to conf,rm
a seasonal colour change in either sex.
The few Pakistan specimens of. Varanus g.
caspius that are available are similar, but have a
ionger tntl (l48Vo SVL), a higher number of middorsal
scales (143) and more body (6-8) and

tailgraph (15-17) bands. The body bands are not
bordered by lighter spots (photograph in Minton
1966). The distal part of the tail is whitish. Additionally,
they reach larger size.
Size and Mass: In the sample of adult Z g.
koniecznyimeasured and weighed in the field during
this study (N = 94), female SVL varied from 19G-
335 mm (N = 39); males i83-365 mm (I'{ = 55).
Males were siightly, but significantly larger than
females [K male SVL 286.7 t 34.6 mm, X fema].e
S\lL 268.0 x 33.22; t test = 6.5; p < 0.001). The
proportional difference in SVL between the sexes is
less than in all other monitors studied so far. Total
length in maies varies fromS'4&to 835 mrn, and in
females a6m:4ffiq.?52 mm. The uil is longer than
the body, as in most monitor species. There is no significant
sexual dilference in the proportional length
of the tail, the mean SVLi Tail L 6 1.23 t 1.0 in
males, 1.20t 0.1 in females). Its tip is missing in i1
Vo,of the males and 12.8 Vo of the females, with no
significant sexuai difference betlveen them in this
regard.
Total weight varied from 62 to 580 g in females
CK 250.9 t i33.1 g) and'85 to 520 g in males (X
296.2x 106.0 g).
Among sexually mature adults there is no significant
relation betqeen SVL and Wt (best fit with
exponentirl curve, R2 = A.29), suggesting that total
weights vary greatly intra-individually and
seasonally. This high variance is probably due to
variation in local insect abundance.
Sex ratio: A total of 96 sperimens were dug from
their burrows in Sind Province, Pakistan. Of these,
the sex of twelve was indeterminant (gonadal immaturity).
The sex ratio of adults (84) was 58 males
to,26 females (2.23:1), which is s^ignificantly different@

1 000
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< 600
o
- 4oo
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fr zoo
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JFMAMJJASOND
Clutch size of oviducal eggs ftuges from 6-11;
luteal bodies remain in the ovary for at least one year,
and range in number from 2-15. Thus our'evidence
from Pakistan suggests that mature females lay from
2 tn 15 eggs, with the rnean for this area calculated
as 8.7 t 4.3.
Females appear !o produce only one clutch per
breeding season. All females throughout the subspecies
range are synchronous in respect to gonadal
acdviry. The ovaries of individuals u.ith ovi$ucal
eggs generally conBined both corpora lutea and a
number of degenerating ova. We found no evidence
thateggspass through theoviducts of this subspecies
as rapidly as suggested by Jacob and Ramswami
(1976) for 11 bengalenis, who reporred the
presence of mafure follicles, and both shelled and
unshelled eggs in a single individual. Ondarion occurs
mainly in August and September, with continuing
slow decrease in ovary volume through January
Fig. 2), after which there is a radical reducrion in
ovary size March through June- Ovulation in this
subspecies is later than that in the sympatric Z
tlav escens (Auffenberg e t al. in press).
Coqpora lutea vary frlcm 2 to 15. There is no
significant relationship (R'= 0.0i) berween female
size (SW) and number of ovulated eggs (number of
corpora lufea).
Testes volume of all males with SVL >250 mm
shows a significant seasonal change 1Fig. 2). Testes
volume is highest in Aprii, gradually becoming more
JOURNAL, BOMBAY NATURAL HIST. S0CIEW, Vol.87
Fig. 2. Seasonal change irr ovary and testes volume and the diameter of the largest follicle in Va ranus griseus knniecznyi.
or less smaller in the following mon&s through Juiy.
In August there is a dramatic decrease in testes
volume, after which it slowly increases through
December, followed by a more rapid rise to April.
Varanus g. koniecznyi exhibits a composite
reproduclive cycle. Malesproduce sperm during the
Wring, after they emerge from a period of dormancy
(or at least, reduced activity) from November
through March. Thus testicular recrudescence takes
place when ambient temperalures are increasing in
spring. Combaf takes place May, June and early July
(ust before tha monsoon, which in Pakisun and
western India usually begins between mid-July and
mid-August). This is suggested both by the presence
of combat scars on the shoulders of adult males (see
Auffenberg 1988) and combatantpairs during these
monihs. Ovarian activity (as well as courShip and
breeding) occurs during August. Mosteggs are laid
in September, a few in October.
The timing of reproductive activiry in this subspecies
of monitor is of considerable interest. We
have demonstrated that the onset of testicular ac.
tivity is correiated with increasing ambient. temperafure;rln
contrast, ovarian activity is not corelaBsd
with temperature, but is greatest during the monsoon
season. Because ambient temperafure and precipitation
are not correlated, the gonadal activity of males
and females are aslnchronous.
Reproductive activity is controlled by complex
environmental cues, including temperature,

phomperiod an{ precipitarion (see Duvall et a/'
iggz'for reuie1t).-Lichi (f9?1) and Marion (1982)
have demonstrated thai lizard tcsticular activity is
stimulatedby increasing temperature. Thus the male
desert monitor annual reproductive cycle with spermatogenesis
in spring is not surprising. However,
the fact that female gonadal activity does not begin
until much later in the year is unusual among
monitors. Guillette and Sullivan (1985) have
demonsfaied a similar asynchronous reproduclive
cycle in male and female iguanid lizards (S celoportts
formosus).
BIOL,OGY OF VARANUS GRISEUS KONiECZ}I).]
However, in that case females are preg-
nant iluring spring. They point out that. pregnancy
hormones would block any stimulatory environmental
cues (through action of the anti-gonadal hormone
progesterone). However, there is no evidence that
female V. g. koniecznyl are pregnant during spring,
so that the mechanism inhibiting ovarian activity in
females during the earlier parr of the year remains
unknown.
Unfortunately, we are not cefiain when the
eggs hatch. Minton (i966) collected what he
beiieved n be newly hatched individuals in August
and September (SVL 94 mm). Three small
specimens (K.SVL i02 mm); in collections of the
Caiifornia Academy of Science and M.S. Khan
(Rabwah, Pakistan) were collected together during
.Iu.ly fr.on the roof of a thatched hut, suggesring they
had hatched several weeks previousiy (sibling hatchling
monitors sometimes remain together for some
time (Auffenberg 1 98 1, i 988). Thus the curent data
suggest that 14 g. koniecznyi harch from July through
September (monsoon), firrther suggesting that incubation
takes about 10 months.
This premise requires confirmation from
Pakistani and lndian biologists. In Tunisia, Thilenius
(l897freports hatchlings 3 to 4 months after;helled
eggs are found in the oviducts. Hoiveveq his total
length measuremenb of about 30 cm are much loo
great for young of the year, su ggesting that his "hatshlings"
were already one season old, Thus there is
no information on how long incubation lasts in
Tunisia.
The long incubation period suggested for V g.
koniecznyi is not unique among monitors (though
exceptionally long when compared to lizard species
in other families), f.or Varanus niloticrts requires a
similar incubation period in some parts of its range
(Corvles 1930). Other workers have demonstraed
that the incubaticn of the eggs of other monitor
species 4nay take 4 to 6'months (see Auffenberg i988
for review). In every case wheie comparisons are
possible, the hatchlng of monitors seems correlated
wltn tle onset of the raily season, which evidence
suggests is also true of V. g. ianiecznyl' This is, of
course, *re time when insectprey usuaily reach their
hrghest annual peaks.
Thilenius (1897) suggests the possibiliry that
in Tunisia, female desert monitors may reLrm to the
nest near the time of hatching. Auffenberg (198i)
repeats a loca1 rural story that the females of 14
knmodoensis do likewise, though he saw no
evidence for such behaviour. Additiorral informarion
on such possible maternal behaviour on the part of
aduit females wouid be very important, if ccnfirmed-
Ndwly hatched monitors of many species are
rarel| seen in the wild even by professional monitor
hunters. We presume this is usually due to their often
arboreal habits when young. However, hatchlings of
Varanw griseus are also rarely caught when compared
to aduits. The paucity of trees in their usual
habitat suggests they are spending their time else-
whe.re. This is another area in which local biologists
can shed much light on - where very young monilors
spend most of their time. In conE ast, ovarian activity
is not. correlated with temperanue, but is greatest
during. the monsoon season. Because ambient
temperattue and precipitalion are nol correlated, the
gonadal activity of maies and females are
asynchronous.
On the basis oi limited growth data on captive
Vqranus grisew caspius (Segeev 1939), V. g.
knniecznyi may become sexually marurs at the end
of the second year of life, certainly during the third-
Thilenius (1897) believes that the larger nominal
subspecies may become rnatue as early as two
years, but for most individuals he believes it is 4 or
5 years. They have been loown to live in captivity
at least ten years @ower 1925).
Abdominal Fat Bodies. Male and female fatbodies
exhibitidentical annual cycles (Fig. 3), as they do in
tlre tropical evergreen species Varanus olivaceus
(Auffenberg 1988). Inrbo& sexes of V. griseus the
fat, body weight. begms !o accumulate during the
mon$oon season, reaching a peak ia September and
Octffi*IT'his is' directiy associated with increased
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Fig. 3
JOURNAL, BCMIIAY NATURAI, HIS|. SOCIET|. Vct. t7
TyTrtM.JJASO
AMominal fat body rycles of male @iI) and female (l) Varanus grisere koniecmyt
FMAMJJASO
Fig. 4. Annual rycle of proportional Jiver weighr (SV!4|_iver Wr g x 100)
abundance of insect prey (Auffenberg, MS). Ab- dulng vitellogenesis in varanid lizards. Auffenberg
dominal far in both sexes begins tolecrease in *a f"rienle?i (in press) have suggesred rhar the
November' correlated with the ueginningof annual g.uygul tur .ay"u" the source of some (or ali) of the
dormancy and following egg aeposiriin during lipidsusediniilttogenaslsinphilippinescincidsas
August and Septembet. ftriJfit UoAy reOuciion is well.
later than viteilogenesis,__suggesting that the fat In both sexes abdominai fat weight is least
bodies in v' giseus' as in 14 oTiuorrul, are not used. during April, May and June. while ir is rue that this
for this purpose' The senior author has suggested. inverJe retationship agrees with spermatogenesis in
elsewhere (Auffenberg 1988) that considerffrle fat m*e aeserimonlmrs lmost *arkedin April), it does
is also stored in the ail of varanids and that this notugr."*i*ruienogenesisin *ierem'ate (mainly
varies.seasonally- It is possible that caudal fat, rather eugu"st).
than abdominal fat is a more important lipid source Thus the weight of abdominal fat in both sexes
N
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M.
D

TAslar
SiIf VACH CONTE|ITS OF Va rants g r is e us ko n i e c z ny i L\
RAJASTI{AN. BASM ON DATA PROVTDED l\t DAVE (1%1)
.Food
lnsects
Noninsect invertebrales
Amphibians
Lizards
Snakes
Birds
Eggs (various)
category
BIOI.OGY OF VARANUS GRISEUS KONIECZ{I'I
Percent ofrotal
48.0
5.0
2"9
19. 1
10.0
4.2
10.8
'$'-WgrEz
sToMAcrI coNTENTs orlaiazzs griseus koniecznyi FR0M
SNIT:SATTSTAN BASED ON DATA OBTAINED DURING THIS
. STIJDY
Food category
Centipedes
Scolapendridae
Cockroaches
Blanidae (Ar e r,a r i rts)
Beetles
Scarabidae
Tenebriqridae
Toads
Bufcrridae (B ufo s t oma t b us)
Reptiles
Agamrirlae (Uro rnastiz hardw icki i,
Calotes versicolor, Agann sP.)
l-acerridae (Aca ilhocephalus canlor;s)
Reptile eggs
RodcnLs
Mnridae (Meriones cf. hurriants)
Percent of total
1.3
5.3
21.1
56.2
1.3
7.8
1.5
4.0
l?
taken by Varanus giseus koniecznyi' Corkhill
(1928) reporred that they fed chiefly on small rodenis,'lizards,
snakes and crickets. Minton (i966)
rcports that his captives fed on mice, rats, ftsh, meat
-a "ggs, and small toads- The inference from these
repo{Jis thatV. g. koniecznyi feeds-largely upon
sina[ vertebrates. IIowever, Dave (1961) provided
a long list of prey taken from the stomachs of individuais
captured in Rajasthan,India (no indication
of number examined). More importantly, his data
show that insects comprise the major food category
(Iable 1).
' Food remains were recovered from the
stomachs of 75 specimens during this-study' All
those: individuals captured do-.ng December
through,Mafclr- hadpdpiy stomachs and intestines'
Or:r'6talshow that during these three months 14 g'
koniec:znyi is usually inactive. Insects are the most
common food category t*en (t2.6%o); vertebrates
(including their eggs) comprise 1 sm{l gerleltaSe
nf tle totat stomach contena (15.97o; Table2)' Thus
in both of tiose studies in which food habia are
based on wild caught specimens, insects
predominate; vertebrates are not the most common
food of this subsPecies.
Almost all the prey in Pakistan specimens are
partly fossorial. The gerbill' desert cochoaches'
ienripedes, tenebionid beedes, toads, and reptile
esss are all most commonly encountered under
sio"nes and surface detritus, or in rheir bunows' It is
aiso highly likely that at least the juvenile Uromas-
exhibit cyclic changes, with ttre greatest weight
achieveil during and immediately after the monsoon
tix hardwickil were also taken from their burrows'
season (Fig.3).
Acantho dac ry lus cantori s regularly hides in burrows
Liver:Thg pattem of seasonal changes in propor-
and could have been retrieved from hoies. However,
tional liver weight has been investigated inVaranw
Calotes versicolor was almost certainly taken while
oliv aceus (Auffenberg 1988) and Z/Tavescens (Auf-
on the ground (nonnally an arboreal form). Scarib
fenberget al.in press). Fig' 4 demonstrates seasonal
beetles are commonly found either on the ground at
differJnces in the weight of this organ in I4g'
cow or camel pats, or within pats or the soil beneath
konkcznyi, The'annual peak in both males and
them. When'the entire sp€ctrum of prey taken is
females is during August, when, in the females,
reviewed it becomes obvious that almost all the prey
vitellogenesis occurs. However, the fact that liver
--are
fbund- below the surf;ace or exricated from
weighi is also highest in males during this same
beneath debris on the desert floor.
monttr suggests that greater liver weight is related to
The lar-ggst prey taken are Uroma'stix
increased insect prey abundance during the same^
lnrdwickii-bffwere juveniles with a mean SVL of
month (Auffenberg, unpubl. data). Lowest. liver
77 inm. One adult deseitmoriitor contained 5 hatc-
weights occur during May and June, when insect
hiing U. hmdwickii,tvto of which apipeared t9 have
prey are the least abundant.
Ueen,eaten the day before, andthree'the day it was
hood and its seasonal utilization- Several previous
captured, Another adult monitor contained remains
workers have reported in a general way on the prey

JOUP!^'|AL, BOMBAY NATARAL HIST. SOCIEW, Vol.BZ
SEASONAL PREY urlI-zATIoN Lr{ uoron* gri""* k:rl:"?n"ri o,orr ro*, nuxtsTAN Gi-qo oF Mot-tTrly TorA$)
Prey Type Jan. M"y Jun
Cenripedes
Bmtles
Blaurds
Toads
Lizalds
Reptile eggs
Rodmrs
1.4 0.5
x.0 '70.6 93.3
17.6
50.0
.4J.6
of two ha tc hlings. Most beetle prey were represen ted
by a number of individuals in each sromaah (mean
10.2 per stomach). The mean cockroach length is
10.0 mm; mean beetle length 11.9 mm. Thus the vast
majoriiy of the prey eaten are small in proportion to
predator lengrh. In spire of prey lisrs in the literatue
that tend to emphasize the vertebrate prey of
V-aranus griseus koniecznyi, most. prey are small
beetles they dig from beneath detritus or from shallow
burrows, usually unCer desert slrrubs.
Dave (1961) demonstrates considerable
seasonal variation in the prey taken by the desert
monitor in Rajasthan, India. He concludes thar insects
comprise the major prey during July and
August; reptiles are eaten primarily in Maich
tlrcugh April, and eggs (onjer not derennined) in
IV{ay and June (but no quantitalive seasonai data is
providei).
Table 3 provides data obtained during the current
study on seasonal prey utilizatron in Sind
kovince, Pakistan. No foocl was found in any desert
monitor stomachs from December through March.
Beedes were taken every rnonth that the monitors
w3re active, varying from abou t 4 n 9 6 Vo o f the O fal
monthly items. Lizards were taken only during the
monsoon season; reptile eggs mainly before the
monsoon. The remainder of the prey categories
reflect no obvious pattem. Thus the seasonal utilization
of prey in Pakistan is somewhat different from
what Dave descrioed in western India.
Utilization. In connection with study of the behavioural
ecology of monitor lizards in pakistan and
India during five yeais by the senior author, there is
no clear evidence thar this monitor lizard is regular-
l;r (if ar all) hunted for iE leather. This is fqking in
rFQf.p.Itltte tacr that$.appropdate habirats ir is rather
common. Not once have we seen a leather product
(handbag, waller; etc.) made of ix skin in either of
Jul, Aug S"p. Oct. Nov. Dec.
98.9
11.8 L2 f .i
t.J
79.3
8.8
78.8
10,1
5.9
18.0 2.3 2.4
1.3
96.5
these counlries. Nor have we noted it among *re
thousands of confiscated skins ready for proceising
in tanning establishments. Such skins are always oi
Varanus bengalensis utd,V. flavescens (in pakrsun
and western India). The reason seems to be that individuals
of both the last two species tend to be concentrated,
whereas individuais of the desert monitor,
while cleariy more abundant on the basis of comparative
sizes of the areas intrabited by each, tend {.o
be more widely scattered over the vast sandy arid
ftacts in which they are found. We found no evidence
that the desert monitor ltzard is avoided by hunters
(though many uninformed rural people believe if is
venomous, the professionai animal catchers do not).
In general, through both India and paHstan
much of the original preferred habitat of this subspecies
remains intact. This is due marnly to its
atidiry. Some schemes have been proposed in both
countries to convert some of these desert lands
(where they lie near a major water source) to agricultural
land. This will undoubtedly result in some
habitat desruction of importance to the desert
monitor. However, viewed in broad perspective,
such schemes will have tiftle significanr effecr on rhe
totalify of the geographic area in which it eKisrs.
lCost of this iand cannot be irrigated and thus the
total range will probably not experience as much significant
future modification as the habitats of rhe
other indo-Pakistani monitor species. In fact, the
continued salination (and thus abandonment for
agricultural purposes) of marginal desert habitat
through hydro-agricutfure may, in the long run,
match and even exceed the effects of habitatdestruction
through urbanization and highway construction.
However, proof of this remains for the future. These
observafions suggest that of the three species tbund
in southern arid Asia, V. g. koniecznyl is the least
ftreatened.
3.5

AcxNowI-EDcENcNTS
Thanks are extended to the following individuals
for allowing the study of prepared
materials in their care (museum abbreviations in
parentheses where appropriate): Dr Farooq M.
Ahmed, Direc t or, Zoolo gical S urvey Dept", Karachi
(ZSD); Dr N. Arnold, British Museum (Narural l{istcry),
I-ondon (BMNII); Mr K.J. Baig, Pakistan
Museum Natural History, Islamabad; Mr J.C.
Daniel, Bombay Nanrai History Society GNHS);
Dr W. Bohme, Alexander Koenig Museum Natural
History Bann; Dr'J.Eiseli, Vienna Museum Natural
History; Dr O. Gruber, Zoologisches Staatmuseum
Munchen (ZSlv!; Dr K. Klemmer, Senckenberg
Museurn Natural History, Frankfurt (Sil{F); lt{r H.
Marx, Chicago Museum Natural History, Chicago.
Mr S"lvt. Nair, Director, lr{useum Natural History,
New Delhi; Mr J-P. Rosada, Museum Compararive
Zoolog.v, I{arvard University, Cambridge @{ass.);
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