Asiatic

December 1993

Herpetological ResearchVol. 5, pp. 85-95 1

Notes on a Collection of Squamate Reptiles from Eastern Mindanao,
Philippine Islands Part 1: Lacertilia

Brian E. Smith
Museum of Natural History. Louisiana Stale University, Baton Rouge, Louisiana 70803-3216. Present Address:
Department of Biology. University of Texas at Arlington Box 19498 Arlington, Texas 76019-0498
Abstract. -In 1982, I spent six months collecting the herpetofauna of several areas in eastern Mindanao,
conclusions
Philippine Islands. I present species accounts for all lizards collected at these sites, and draw

about the biodiversity of second-growth and primary forest habitats.

I

found second-growth habitats to beSpecies which may depend on primary forest habitat andalso species apparently restricted to such habitats are detailed. The importance of primary forest reserves,
selective logging, and a mosaic of successional habitats within primary forest reserves is discussed.

depauparate compared to primary forest habitats.

Keywords:

Reptilia,

Squamata,

Lacertilia,

Introduction
Little

is

known

of

the

ecology,

Taylor (1922a, b, c; 1923; 1925)
thesystematics andinvestigatedofthePhilippine
zoogeography
herpetofauna and provided some ecological
Brown and Alcala (1961,
information.
1964, 1978, 1980) undertook ecologicaland systematic studies on some islands,
Alcala (1986) recently reviewed theherpetofauna, and Auffenberg (1988) has
detailed the ecology of the Gray's monitor,
However, many
Varanus olivaceus.
islands remain poorly known. This and asubsequent paper provide information on
lizards.

the natural history of squamate reptiles
collected in eastern Mindanao, Philippine
Islands, during a collecting trip to the area
in

1982.

I

also

biodiversity

selectively logged one or two years
previously. It has abundant ground coverof rattan, herbaceous vines, shrubs, andA few small (<2 m) trees werelianas.
present, and the ground was littered withfallen logs.

should primary forest continue to be lost to
logging and agricultural practices.

—

The
Vegetation.
vegetation of the Philippines was described
Description

by Brown (1919) and Dickerson (1928)
and a detailed study of southeast Asian rain
forests was published by Richards (1952).
Trees of the family Dipterocarpaceae
dominate the rain forests of the Philippines.
For the purposes of my study, primary
dipterocarp forest is defined as dipterocarp
forest that has apparently never been

© 1993 by Asiatic Herpetological Research

Selective logging was practiced

study sites, and some small
dipterocarps were left standing, though notenough to cast appreciable shade. Lateatmysecond-growth dipterocarp forest is defined
as forest selectively logged three or more
years ago. It usually had a closed canopy
composed of young fast-growing trees(mostly Trema species) about 10-15 m tall.
Small dipterocarps were present, and theherbaceous understory was extremely
dense. The vegetation of my study areas is
discussed in more detail in Smith (1985).

comment onsquamate assemblages occurring in primaryand second-growth dipterocarp forest in aneffort to pinpoint species of special concern

Site

ecology,logged. Early second-growth dipterocarp
forest is defined as dipterocarp forest

distribution, and life history of Philippine

madetaxonomy,

Philippines,

Site

1.

—This

site

was located in the

coastal mountains (Diuata Range) of eastcentral Mindanao, 55 km south, 20 km

west of Bislig Bay and 10 km southeast of
Mt. Agtuuganon (Fig. 1). Areas from 500800m elevation were sampled. Vegetationof this area was early second-growth andprimary dipterocarp forest. Slopes weregenerally very steep (Fig. 2, 3, 4). This
site was sampled from April 6 to August
15, 1982.

Site 2.

—This

Range, 33

site was also in the Diuata

km south and 7 km west of

Bislig Bay (Fig. 1). Vegetation at this site
consisted entirely of late second-growthAsiatic Herpetological ResearchVol. 5 p. 86

December 1993

occurs from November to March, with theheaviest rains in December and January(Census Office of the Philippine Islands

1920; Dickerson, 1928; Willmott et al,
Annual rainfall in Surigao (the1981).
1 and 2) is
with
2360
mm,
falling fromof thisNovember to March, and 1 191
amount in December and January alone(Willmott et al., 1981). Surigao is in thelowlands, and rainfall at my collecting sites
may have been higher. Site 3 also does nothave a marked wet or dry season (Census
Office of the Philippine Islands, 1920;
Annual rainfall in
Dickerson, 1928).

nearest weather station to sites

mm

3647

mm

M-f

12^°

125°

Davao (at the base of the Mount Apo range)

2#1942 mm (Willmott et al., 1981). This
figure undoubtably increases withisEast Longitude

FIGURE 1. Map of Mindanao, Philippine Islands,
showing locations of collecting sites

1

and 2

(triangle) and site 3 (circle).

dipterocarp forest.

Slopes were moderately

The site was 400 m inwasandelevation,
sampled from August
inclined to steep.

21 to September 4, 1982.

—

I also visited Mount Talomo, aSite 3.
2693 m peak located 30 km west of Davao
City in the Mount Apo Range of southern
Mindanao (Fig. 1). I sampled areas aroundthe Philippine Eagle Captive Breeding
Project (PECBP) field station (about 1000m elevation) from September 8-13, 1982.
Workers at the field station made incidental
collections at this site from April to

According to residents,
September.
inthisarea was discontinued
loggingsometime in the mid- to late-1960's. The
area was selectively logged and appears to
be more similar to typical pristine forest

many other logged areas I visited.
Human disturbance is considerable on theslopes of Mount Talomo. Farms extend up

elevation. During some times of the year,
the PECBP station may be shrouded inclouds for weeks at a time. Temperature

was relatively constant at all sites. At site 1
it usually ranged from 20-25 C under the
canopy, and from 20-30 C in the open.
The minimum temperature recorded was 18
C, the maximum was 34 C. Temperaturesat site 2 were similar.
Temperature
readings were not available from site 3, but
it was slightly cooler because of the higher
elevation. Humidity at all sites varied from79-100%. Occasionally typhoons strike
Mindanao, but they generally lack theseverity of those hitting more northerly
islands (Census Office of the Philippine
In
Islands, 1920, Dickerson, 1928).

March of 1982 a typhoon struck Bislig Baynear sites 1 and 2. The only effect at the
study sites was moderate, steady rain forseveral days.

Methods

thanthe slopes from Davao Gulf to about 900m

Coconuts, bananas, pineapples,
coffee, and various fruits and vegetables
are grown. Areas sampled range from 600-

elevation.

1050 m.

Slopes are gently to steeply

inclined.

Specimens were collected using drift
fences (Gibbons and Semlitsch, 1981) andhigh andby hand. Drift fences were 0.5inas
detailed
constructed
and
18
long
Gibbons and Semlitsch (1981). Pit-cansmm

and funnel traps were placed at either endof the fence.

capturing—

Climate.
Rainfall was heavy at allsites. At sites 1 and 2, there is no marked
dry season. The wetter season usuallyall

Pit-cans seemed effective atterrestrial lizards except

Varanus spp. Funnel traps were primarily
useful in capturing snakes. Drift fences
appeared to adequately sample reptiles
moving on the soil surface. To sample

December 1993

Asiatic Herpetological ResearchVol. 5 p. 87

FIGURE 2. Primary forest at the edge of a road cut at site 1. The ridge top is about 900m elevation.
arboreal fauna, I examined epiphytic ferns
and trees felled during logging. These

were largely unsuccessful andarboreal species are under-represented in
Data on macrohabitat
the collection.
(primary, early, or late second-growth
dipterocarp forest), microhabitat (fossorial,
terrestrial, or arboreal), elevation, date, andtime of day were recorded for eachpreviously published data was noted, andscale count data are not reported herein.

efforts

Species Accounts
Family Gekkonidae—

Specimens ofCyrtodactylus agusanensis:
this species were collected in all habitats atand 2. This species' morphology
suggests nocturnal habits (verticallyStandard
collected.
measurements including snout-vent (SVL),
tail (TL), and total length (TTL) were takenin the field on freshly killed specimens.
General weather conditions were noted
daily.
Specimens were dissected in thelaboratory to determine sex, stomach
contents, and reproductive condition of
females. Food contents were identified

sites

speciesisusually to family for the arthropod prey ofrivers.

My specimens were all taken far

specimenthese lizards.
also taken,

Standard scale counts were
but no deviation from1

elliptical pupils), but half the specimens
were captured in drift fences during theSpecimens captured at night weretaken on bushes and logs, 1-3 m above
ground. My observations do not agreewith Alcala (1986), who states that this
day.

found

in

swamps and alongfrom such habitats. I also did not find thisspecies to be particularly rare, as did Alcala

Asiatic Herpetological ResearchVol. 5 p.

(1986). Females taken June 14 and 23 had
one large egg (15.7-18.2 mm long) in each
oviduct. Small males were captured April 2
and 9 and a small female July 31. One
juvenile was captured at site 2 on August26.

December 1993

reported from Catagan and Malindang

Mountainin

Mindanaonorthwestern(Taylor, 1922a) and the Diuata Mountains
in the province of Davao del Norte, eastcentral Mindanao (this study).

Stomach contents included insects ofseveral families and a shed skin, prohahlyof C. agusanensis.

Specimens examined:

Specimens examined:
41680.

LSUMZ 41601-

Draco

volans.

LSUMZ 41678-—This species

is

very

common in early second-growth forest and41609, 41640.

probably the most conspicuous lizard
species in this or any other habitat.
Individuals are commonly seen runningalong branches, displaying, and gliding.
is

Family Agamidae

From

their

morphology,

allagamids

collected on the study sites appear to bearboreal, although most specimens, except
Draco species, were caught on or near the

ground.

20.

—

A female wasCalotes cristatellus:
collected at about 550 m in early secondgrowth forest at site 1. A male specimen
lacks additional data. This species is rare,transient in the habitats sampled, or mostly

arboreal and hence under-represented in theIt has morphology indicative ofa highly arboreal lifestyle. The female,collected on July 1, had one large egg (38.4

collection.

and 35.2 mm) in each oviduct. Stomach
contents included lepidopteran larvae,
unidentified insects, and a snail.

Specimens examined:
41738.

LSUMZ 41737,—

base of Malindang Mountain, northwestern

Mindanao.

Inger (1983) examinedtaxonomic characters in nine specimens but
gave no ecological data for them. My
specimens were taken at 650 m at site 1 in
The species may be
primary forest.
confined to primary forest.
Draco
mindanensis is diurnal and arboreal. A
female contained two oviducal eggs 17.5
and 18.4
long. The date of capture ofmm

specimenisunknown.

Taylor (1922a) stated, and my data

confirm, that this species feeds exclusively

on ants.

Specimens examined:
41748.

LSUMZ 41741-

Gonyocephalus semperi.

— Althougharboreal by morphology, five specimens
were captured on the ground in drift fences.

One was caught by hand 1 m above theground on a large tree. This species isdiurnal, as is its' congener G. godeffroyi in
the Solomon Islands (McCoy 1980). Fourwere captured in primary forest, two in late

Due to the species'
arboreal habits, it is highly likely that G.
semperi does not occur in highly disturbed
areas largely lacking trees. McCoy (1980)
found that G. godeffroyi also avoids open
areas in the Solomons. One G. semperi
exhibited aggression and grunted whenhandled. The single female captured June
16 contained three developing eggs (5.8,
in length).
Stomachs
6.6, and 7.3
examined contained the remains of insects
second-growth forest.

Draco mindanensis: Taylor (1922a)collected only two specimens of thisspecies. They were taken at 1 100 ft. at thethisThey are diurnal and exclusively arboreal.
This species was never seen in primary
forest. A female containing two eggs (14.7
and 13.9 mm long) was collected on July

Stomachcontents consisted of several families of
insects. This species is apparently not an
ant-feeding specialist like its congener, D.
volans (see below). D. mindanensis ismmof the families Chilopoda, Coleoptera,Hymenoptera, Orthoptera, and larval
Lepidoptera.

Specimens examined:
41735.

LSUMZ 41730-

Hydrosaurus pustulosus.

—This

is

a

juvenile specimen collected by a native onAsiatic Herpetological Research

December 1993

Vol. 5 p. 89

Figure 3. Early second-growth forest at site 1. This site had been selectively logged two or three years
prior to this photograph. Trees in the middle backgound were deliberately left standing during the selective
logging procedure.

June

19.

Its'

stomach was empty.

It is

Family Scincidae

said by Alcala (1986) to be omnivorous,

which generally agrees with observationsof captive specimens at the Dallas Zoo.
Auffenberg (1988) states that adult H.
pustulosus are entirely folivorous in thewild. Captive specimens usually lay 6-8
in length
eggs measuring roughly 50
about once a year (Mitchell 1985). This
species is said to be common in thePhilippines near unpolluted mountainstreams (Alcala 1986). It has also been
observed around coastal fishing villages,

mmutilizing as vertical perches the stilts orpiers that support homes over water (L. A.

Mitchell, personal observation).

In the Philippines, skinks far exceed theother lizard families in number of species,
abundance, and probably in the variety ofniches they occupy. They are often themost abundant lizards

in

all

habitats

sampled, with the exception of earlysecond-growth forest, where Draco volans
The leaf litterismore abundant.
herpetofauna is dominated by species of thefamily Scincidae.

Brachymeles

gracilishilong:—Specimens were captured only in latesecond-growth forest (site 2, 400 m) and at

Mount Talomo (site 3). Brown and Alcala
Specimen examined: LSUMZ 41739.

(1980) stated that this fossorial species isfound under leaves, duff, rotting logs, andAsiatic Herpetological ResearchVol. 5 p. 90

Figure 4.

Small permanent stream in primary forest at site

December 1993

Small streams such as this one were1.

common in areas of primary forest at all sites.
in loose soil, usually only in primary forestfrom 50-1000 m elevation. This species isapparently rare or absent in early secondgrowth forest.

Stomach contents indicatethat this species is a generalized insectivore,

however, part of a skink tail (Mabuya or
Sphenomorphus species) was found in one
stomach.

Specimens examined:

LSUMZ 41719-mm

in her oviduct. Brown andAlcala (1980) stated that this subspecies isovoviviparous and usually has 2 or 3
young. This species is also a generalized
In addition, a lizard tailinsectivore.

and 15.0(probably Brachymeles species) was foundin one specimen's stomach.

Specimens examined:
41729.

41725.

—

Brachymeles schadenbergi oriental is.
This fossorial species was often caught infences after long, steady rains. It isfound under logs and in leaf litter and loose
soil in primary and second-growth forests
at elevations from 50- 10(H) m (Brown andAlcala, 1980). A female collected July 5
had three developing embryos 18.1, 15.9,

drift

LSUMZ 41726-—

Lamprolepis smaragdina philippinica.
This was a very common arboreal species
in cultivated areas and villages, and a single
specimen was taken in a coconut plantation
on Mount Talomo between 700 and 800 m
elevation. In contrast to Brown and Alcala(1980) and Alcala (1986), I never observed
this species in primary or second-growthforests.
study sites did not encompass

MyAsiatic Herpetological Research

December 1993

myagricultural areas or villages.

sites

in

individuals.

Specimen examined:

LSUMZ 4 648.

—

stumps and hollow

Alcalaandrare.
arborealthat
itisstates(1986)
in
from
an
was
taken
daylight
My specimen
in
4
m
above
fern
primary
groundepiphytic
forest at 800

trees.

m elevation.

Specimen examined:—

This is a very active and abundant skink
that favors open areas. It is diurnal andAll but two specimens wereterrestrial.

caught in early second-growth forest. One
specimen was taken in late second-growth
An additional specimen wasforest.
observed in primary forest, but this waswithin 100 m of a logging road and it may
have been transient. This species seems to
be a lizard of open areas, and it may haveoriginally occupied natural treefall gaps in
the forest. With extensive clearing of areas

logging and cultivation, M. m.
multicarinata has probably increased
Of elevensubstantially in numbers.
females collected throughout this study,
only two failed to contain well-developed
eggs. The rest had 2 (6 specimens) or 3 (3forspecimens) large oviducal eggs 11.0-15.5
SVL werelong. Juveniles 29-41
24
andAugust 1,4, 12,captured on Juneand 26. Another juvenile was taken fromthe stomach of a snake (Cyclocorus

mm

mm

nuchalis taylori) that was collected July 1.
Stomachs examined contained insects ofmany taxa, and I consider this species ageneralized insectivore.

Specimens examined:
41639.

species are mostly diurnal

Sphenomorphus acutus:— Brown andAlcala (1980) state that this species isstrictly arboreal, which may account for thepaucity of specimens collected. However,three specimens were collected in drift
fences during daylight, so they are at leastoccasionally found on the ground. Thisagrees with Alcala's (1986) observations.
Stomach contents included arachnids and

LSUMZ 41740.

Mabuya multicarinata multicarinata.

number of species andIts'and terrestrial.

1

Taylor (1922a) noted
Lipinia semperi.
that this species is commonly found in old
treeVol. 5 p. 91

LSUMZ 41610-

orthopterans.

Specimens examined:
41715.

Sphenomorphus coxi coxi.

—This wasthe most common ground-dwelling lizard inprimary forest and is the mostconspicuous member of the leaf-littertheherpetofauna in this habitat. It is fairly
common in second-growth forest as well,
but it is not seen in the open as often as
Mabuya multicarinata multicarinata.

Although unquantified, I believe that theseobservations represent a real difference,and that M. m. multicarinata replaces 5. c.
coxi as the dominant skink in second-

growthhabitats.

Gravid females weremmcollected April 27 (1 egg, 5.2long),
July 3 (2 eggs, 8.5 and 9.2 mm), andAugust 5 (2 eggs, 14.7 and 15.2 mm).
One juvenile was collected May 2 1 and two
on May 24. Four others were collected

August 9, 13 (2), and 28. These juveniles
measured 34-45 mm SVL. One small
specimen identified as a male measured 45
mm SVL. These data suggest two hatchingseasons during the six months of thisstudy, one in May and the other in August.

Apparently, reproductive maturity is
SVL.
reached at approximately 45
Stomachs examined contained insects ofmany taxa. I consider this species to be ammSphenomorphus Species

generalized insectivore.

Lizards of the genus Sphenomorphus
dominate the leaf litter herpetofauna of theSpecimens examined:
41677.

primary forest in the areas I investigated,
and they are sometimes conspicuous inwell.
as
growthsecondaryallothergenera atSphenomorphus exceeds

LSUMZ 41713-

LSUMZ 41649-

Sphenomorphus decipiens.

— Although

Alcala (1986) states that this species is rare,
I found it to be a fairly common diurnal

Asiatic Herpetological ResearchVol. 5 p. 92

member of the leaf litter herpetofauna inforest.

It

was never foundprimary
second-growth forest.

in

A juvenile (25 mm
Stomachs

SVL) was collected June 23.

contained larval lepidopterans and otherinsects. Specimens examined:
41704-41711.

Sphenomorphus fasciatus.

LSUMZ— Brown andAlcala (1980) noted that this species was acommon terrestrial skink.

Duringmystudy, I collected only one specimen at site
1.
Since my capture techniques seemed
especially efficient in sampling theterrestrial herpetofauna, I conclude that S.
fasciatus was rare at my collecting sites.
The stomach of this specimen contained
insects of several taxa.

Specimen examined:

were either captured in drift fence traps or
found under cover. At night, individuals
were found actively foraging. This speciesisSphenomorphus steerei.

—This species

commonforest,in

insects of several different taxa.

Specimens examined:
41703,41712.

LSUMZ 41697-

apparently secretive during daylight.

Specimens frequently grunted duringAll
capture and attempted to bite.
specimens were caught in primary forest
except one caught in late second-growthforest. This species is apparently absent
from highly disturbed areas.
Gravidfemales were caught March 16 (5 eggs,
3.5-6.3 mm) and June 14 (5 eggs, 12.313.9 mm).

Specimens examined:
41647.

The continuing destruction of tropicalrain forest worldwide makes it imperativethat species of special concern are identified
in these habitats.

It was never foundsecond-growth forest, but two
specimens were taken in late secondgrowth forest. S. variegatus appeared to becompletely absent from highly disturbed
areas. Females taken April 1 1 and May 24
each contained two eggs (11.0 and 10.3mm; and 6.5 and 6.1 mm, respectively).
Juveniles (SVL 24-39 mm) were captured
May 23, June 29, August 9, and August
26. Stomachs contained insects of manyIn the section below, I

will attempt to pinpoint species which could

be adversely

affectedbycontinuing

deforestation, where my data are adequate
to do so. I make special reference to thelizards of the forest floor, since I feel thatthese lizards were the most accurately
sampled species with the techniques that I
used.
These species are all skinks:

MabuyaSphenomorphus variegatus. — This
species is a common diurnal leaf-litter lizard

LSUMZ 41641-

Discussion

LSUMZ 41716.

and isprimaryinfoundsecond-growthoccasionallyforest. In my study, it was more common
above 650 m elevation, although Brown
and Alcala (1980) stated that it occurs fromsea level to 2000 m. Stomachs containedisDecember 1993

multicarinata

multicarinata,

Sphenomorphus coxi coxi, S. decipiens, S.
steerei, S. variegatus, and Tropidophorus
partelloi. Sphenomorphus fasciatus is notof the primary forest.

considered, since only one specimen of thisin early

species was collected.

different taxa.

Specimens examined:
41696.

LSUMZ 41681-—
Tropidophorus partelloi. This species
was found to be active during the day andnight. Specimens collected during the day

Primary Forest
This was the richest habitat sampled,
containing all six of the skinks mentioned
above.
Of these six, five seem to be
inhabitantsof the primary forest. In
regular
there
were
two common arboreal
addition,

The gecko Cyrtodactylus
agusanensis and the agamid
The most
Gonyocephalus semperi.
lizards
listedterrestrial
collected
commonly
lizards:

indecreasing order of abundance were:

Sphenomorphus coxi coxi, S. variegatus,
S. steerei, S.

decipiens, and Tropidophorus

Species which were sparsely
collected in primary forest include thepartelloi.

Asiatic Herpetological Research

December 1993

Vol. 5 p. 93

arboreal lizards Calotes cristate/Ins, Draco

whereas Draco volans, common in earlysemperi, andandthe terrestrial
Sphenomorphus acutus,

second-growth forest, was absent from latelizard Mabuya multicarinata multicarinata.

skinksmindanensis,

LipiniaI

do not consider M. m. multicarinata to be a
regular inhabitant of primary forest.

Early Second-Growth Forest

second-growth

extremely abundant Mabuya multicarinata
multicarinata. This habitat contained only
three of the six terrestrial skinks of interest.

The skink Sphenomorphus steerei was only
represented by two individuals collected inthis habitat, and I consider it to be rare in

M. m.
second-growth habitats.
multicarinata was virtually absent inprimary forest, but it was found to be twice
as common in second-growth habitats as
the next most common lizard, S. c. coxi. I
spent approximately two man-months
apiece working in primary forest and earlysecond-growth forest at site 1. I haveassumed that the time spent in these two
habitat types was equal and have calculated
early

the Brillouin diversity index (Krebs 1989)
for each habitat using the absolute numberof specimens collected in each habitat typeof the six terrestrial skinks of interest noted
above. Indices of 0.610 and 0.309 are
calculated for primary forest and earlysecond-growth forest habitat types,
Although not an exact
respectively.
thismeasurement,rough estimate ofoutthemajor difference indiversity points
these two habitat
betweenbiodiversity

The

terrestrial

Sphenomorphus variegatus andTropidophorus partelloi, both absent fromearly second-growth forest and common inprimary forest, were collected in latesecond-growth forest.

This was the most depauperate habitat
Two arboreal lizards were
sampled.

common: Cyrtodactylus agusanensis andDraco volans, the latter species being themost common lizard observed in any
The common terrestrial lizards
habitat.
were: Sphenomorphus coxi coxi and theforest.

Species of Special Concern
It is clear thatless complex habitats will

support a less complex community ofplants and animals, by definition.
Although my data are sparse, it is obviousthat the biodiversity of terrestrial skinks(and other lizards) is far different after
logging operations alter the forest structure.
The clearing of primary forest creates
habitat for open habitat specialists such as
Mabuya multicarinata multicarinata andDraco volans, while decreasing habitat forthe species which seem to be confined tothe primary forest. Especially notable is dielack of Sphenomorphus variegatus, which
common in primary forest but absent inisearly second-growth forest.

Other species

which may be adversely affected bylogging activities include the skinks
Tropidophorus partelloi, Sphenomorphus
steerei, and 5. decipiens; and the agamids
Draco mindanensis and Gonyocephalus
semperi. Except for D. mindanensis, allthese species were collected in late secondD.
growth and primary forest.
mindanensis was only found in primary
forest.

Sphenomorphus coxi coxi was the

only lizard commonly found in all habitats,although it was less frequently seen in the
open where it was syntopic with Mabuyamulticarinata multicarinata.

Conclusions

types as regards terrestrial skinks.

The collection of primary forest species
in areas of late second-growth forest points

Late Second-Growth Forest
spent very little time in this habitat
type, yet the species collected here provide
valuable insights into possible lizard
The
population successional patterns.
arboreal agamid Gonyocephalus semperi,absent in early second-growth forest, was
I

found

in

late

second-growthforest.

towards the possibility of a sustained
harvest of the primary forest for lumber.
However, there is absolutely no data on the
periodicity of such a harvest, nor any
precise information on changes inbiodiversity through a successional series.
It is imperative that primary forest be

conserved as a refuge for certain species

Asiatic Herpetological ResearchVol. 5 p. 94

which may only occur there, such as Draco
mindanensis.
Possibly, a system ofreserves with rotating areas of selective
logging may help to conserve Philippine
lizard species.
However, at the present
time research towards such an end isentirely lacking, and the conversion of large
a muchparts of the Philippines towards

impoverishedlizardfauna

continues

unabated.

December 1993

Ministry of Natural Resources and University of
the Philippines, Manila. 195pp.

AUFFENBERG, W.

1988. Gray's Monitor Lizard.
Press, Gainesville,

University of Florida
Florida. 419pp.

BROWN, W. C., AND A. C. ALCALA.

1961.

Populations of amphibians and reptiles in thesubmontane and montane forests of Cuernos de
Negros, Philippine Islands. Ecology 42:628636.

Acknowledgments
BROWN, W. C., AND A. C. ALCALA.
This paper is part of a thesis submitted to
the Louisiana State University Graduate
School in partial fulfillment of therequirements for the degree of Master of
Science. Dr. D. A. Rossman supported
and guided all phases of this work. Dr. J.

V. Remsen, Dr. J. W. Fleeger, and Dr. W.
J.

Harman also assisted in innumerable

ways. Dr. R. S. Kennedy of the Philippine
Eagle Conservation Project initiallysuggested this study and was instrumental
in obtaining funding as were Dr. J. P.
CTNeill and Dr. D. A. Rossman. The
Philippine Paper Company provided the

study

site,

transportation.

andA collecting permit wasstoragespace,

graciously provided by the government of
Mario Caleda,
the Philippine Islands.

Hector Miranda, Kevin Moore,
Wischusen, Mark Witmer, and especially
Susing Babao, all of the Philippine Eagle

1964.

Relationship of the herpetofauna of the nondipterocarp communities to that of thedipterocarp forest of southern Negros Island,
Philippines. Senckenbergiana Biologia 45:591611.

BROWN, W. C., AND A. C. ALCALA.

1978.

Philippine Lizards of the Family Gekkonidae.

Silliman

Sciences

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University

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146pp.

BROWN, W. C, AND A. C. ALCALA.

1980.

Philippine Lizards of the Family Scincidae.

SillimanNatural

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264pp.

Bill

Conservation Project, assisted in obtaining

Robin Lawson, Bill
Sanderson, and Randy Vaeth assisted in

specimens.

identificationof specimens

and food

remains. Jim Murphy and staff at the
Dallas Zoo Herpetology Department
provided an intellectually stimulating
environment in which to finish final editingof this paper. Ardell Mitchell shared his
observations on both captive and wild-caught Hydrosaurus

pustulosus.

Jim

Murphy, Dr. E. D. Brodie, Jr., and Dr. J.
A. Campbell provided helpful comments on
the final draft.

1919. Vegetation of Philippine
Mountains. Manila Bureau of Printing, Manila.

434pp.

CENSUS OFFICE OF THE PHILIPPINE ISLANDS.
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Climatology.
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I,

Manila. 630pp.

DICKERSON, R. E., ED.

1928.

Distribution ofLife in the Philippines.
Monograph of the
Bureau of Science, Manila, Number 21. 322pp.

GIBBONS, J. W., AND R. D. SEMLITSCH.

1981.

An
Terrestrial drift fences with pitfall traps:
effective technique for quantitative sampling ofanimal populations. Brimleyana7:l-16.

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