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Organization of Contiguous Communities of Amphibians and Reptiles in

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Article in Ecological Monographs · July 1977

DOI: 10.2307/1942516

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Organization of Contiguous Communities of Amphibians and Reptiles in
Thailand STOR
Robert F. Inger, Robert K. Colwell

Ecological Monographs, Volume 47, Issue 3 (Summer, 1977),229-253.

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Ecological Monographs (1977) 47: pp.229-253

ORGANIZATION OF CONTIGUOUS COMMUNITIES OF

AMPHIBIANS AND REPTILES IN THAILAND1

ROBERT F. INGER
Field Museum of Natural History, Chicago, Illinois 60605 USA
AND

ROBERT K. COLWELL
Department of Zoology, Ullil'ersity of Cali/omia, Berkeley, California 94720 USA

A bstract. In adjacent areas of broad leaf evergreen forest, deciduous dipterocarp forest,
and agricullural land in northeastern Thailand. >4,000 individuals of !OS species of reptiles
and amphibians were systematically collected. The position of capture of each individual was
recorded in terms of a complex microhabitat code. The 3 environments differ substantially
in pattern of human disturbance, in vegetation structure, and in range and predictability of
temperature and total evaporation, the evergreen forest having the most buffered climate near
the ground.
Of the 3 environments studied, the evergreen forest has the most diverse and distinct
herpetofauna, and the largest percentage of autochthonous species. The evergreen forest
community has more southern or tropical affinities, while the deciduous forest community has
more northern affinities.
Analysis of the use of microhabitat categories by coexisting species reveals significant dif-
ferences, even among the most similar sets of species. Differences in species richness among
the three communities are not explained by differences in microhabitat diversity, mean niche
breadth, or mean niche overlap. However, an ith nearest neighbor analysis of the dispersion of
niche centers in resource space reveals closer packing and larger and more distinct guilds in the
evergreen forest: the environment richest in species and the most predictable in climate. Both
a graphical treatment and an analysis of sets of species grouped a priori according to their
general natural history confirm the existence of larger and significantly tighter guilds in the
evergreen forest.
It is suggested that unpredictable environments tend to prevent the formation of distinct
guilds. while the greater species richness of more predictable environments may be a function
of guild formation.
Key words: Amphibia; ('olll/lwllity e\'olution; CO/ll/I1Ullity strllcture; faunal silllilarity; for-
eSI; guild; microhabitat; nearest neighbor; niche; predictability; Reptilia; species dil'IT,lity;
Thailal1ll; tropical ecology.

INTRODUCTION sites of three vegetation types, we first discuss appor-

General
Biological commullitles, however defined, are
vaguely bounded in space and time. Nonetheless,
the basic heterogeneity of natural environments con-
tinually impels ecologists to categorize habitats and
delimit assemblages of species for the sake of com-
parative studies, and we arc no exceptions. In this
paper we examine, at several levels of resolution, the
ecological organization of the herpetofauna of a
heterogeneous tropical area.
The vegetation of northeast Thailand is a mosaic
of forest types and agricultural lands. In addition to
major differences in vegetation, we show that the en-
vironments also differ considerably in the pre-
dictability in time of critical microclimatic parame-
ters. Using quantitative samples from adjacent study
1 Manuscript received 20 April 1976; accepted 16
February 1977.
tionment among them of the species of amphibians
and reptiles comprising the regional fauna. Looking
then at habitat groups (arboreal, fossorial, terrestrial,
and aquatic) within each of the three communities,
we examine hahitat partitioning on a gross level. At
the next level of resolution we use detailed informa-
tion on microhahitat occupancy for each species in
each community to analyze and contrast the ecologi-
cal organization of the three communities. We are
ahle to account for differences in species diversity
among the communities by means of niche and guild
analysis, in conjunction with differences in micro-
climatic predictahility.
In the sense of Whittaker et al. (1973), the am-
phibians and reptiles of each environment (or habi-
tat) comprise a community, and the partitioning of
available microhabitats is thus one aspect of the
niche, structure of these communities. We llse the
terms "niche breadth" and "niche overlap" in this
paper with reference only to patterns of microhabi-

229
230 ROBERT F. INGER AND ROBERT K. COLWELL
FIG. 1. View from observation tower, looking down
on canopy of dry evergreen forest at Sakaerat, Thailand
during dry season (March).
tat utilization, recogmzmg the potential importance
of trophic and perhaps other kinds of resource par-
titioning in a complete analysis of niche structure.
However, two considerations lead us to believe that
a parallel study of trophic differences among species
of reptiles and amphibians in these communities
would add relatively little to the picture.
First, many of the species involved (particularly
the insectivores) are trophic generalists, eating what-
ever is available in the microhabitats they occupy.
Thus, to the degree that prey categories are asso-
ciated with microhabitats (e.g., arboreal vs. forest-
floor arthropods), niche metrics based on trophic
categories would be highly correlated with those we
compute. Secondly, cases of trophic differenc~s
among species of the herpetofauna which overlap
greatly in microhabitat are often quite obvious, and
are frequently associated with gross taxonomic dif-
ferences.
We have not attempted to separate resting, forag-
ing, and breeding sites. Besides the inherent difficul-
ties of such a task, there is theoretical justification
for eschewing it, as all uses of a microhabitat po-
tentially interact. Exclusion from consideration of
Ecological Monographs
Vol. 47, No.3
all but feeding sites, for example, would fail to reveal
the true extent of both similarities and differences
among species. We will return to these issues in the
Discussion.
Study site
The Sakaerat station is situated =250 km from
Bangkok at 14°30'N, 101°55'E in Province Nakhorn
Ratchasima near the western edge of the Korat
Plateau (=200 m) in an area having a strongly
marked dry season (November to April or May).
Total annual precipitation is =1,500 mm. Although
the station occupies =80 km2, our field activities
were confined to about one fourth of the area.
Three major environmental types exist at and
around the Sakaerat station: dry evergreen forest,
deciduous dipterocarp forest, and agricultural areas.
Both forests are dominated by trees of the family
Dipterocarpaceae. Detailed information is available
on vegetation and climate of the two forest types,
which for simplicity we will refer to as "evergreen"
and "deciduous," but relatively little is available on
the agricultural land.
The evergreen forest has two stories of trees with
dense undergrowth and abundant Hanas. Small
woody vegetation (diameter at breast height [dbh]
1-10 cm) in 100 quadrats (see Collecting Proce-
dures) had a density of 18.05 stems/100 m 2. The
upper story trees, which form a closed canopy (Fig.
1), range in height from 20 to 35 m and in dbh
from 30 to 140 cm. Density of trees in that size range
in 100 quadrats was 1.01/100 m 2 • In March and
April, towards the end of the dry season, 17% of
the forest floor was in sunlight (average of 57 quad-
rats). The nature of floor cover was determined on
three 30-m transects through this forest (14-16
April 1969). Dead leaves accounted for 98% of the
cover and only 1% of the floor was exposed soil.
In contrast, the deciduous forest (Figs. 2-3) has
a single story of trees with a thin undergrowth of
grass, scattered seedlings, low shrubs, and cycads.
Much of the deciduous forest in northeastern Thai-
land is burned each year, destroying undergrowth
and litter. Small woody vegetation (dbh 1-10 cm)
in 100 quadrats had a density of 6.59/100 m 2 •
Mature trees, forming a discontinuous canopy, were
15-25 m high with a maximum dbh of 95 cm. The
density of trees with dbh >30 cm was 0.64/100 m 2
in 100 quadrats. In March and April, 48% of the
forest floor was in sunlight (average of 100 quad-
rats). Several 30-m transects were laid out on the
floor in this forest (14-16 April 1969), 3 in re-
cently burned-over areas and 1 in an area that
. had not been burned for 2 yr (Fig. 3). Dead leaves
accounted for only 20% of the floor surface, bare soil
and rocks for 37%, and erect dead grass for 41 %.
Summer 1977 COMMUNITIES OF AMPHIBIANS AND REPTILES 231

FIG. 2. Deciduous dipterocarp forest at Sakaerat, FIG. 3. Boundary between burned and unburned areas
Thailand during dry season (March). String marks part of deciduous dipterocarp forest at Sakaerat, Thailand dur-
of boundary of sample quadrat. ing dry season (March).

Although these 2 forests are clearly distinguish-
able, certain small areas in which we worked repre-
sented intermediate conditions. Several small patches
of evergreen forest had been so modified by the
removal of largO:!, mature, buttressed trees that their
climate appeared to approach that of the deciduous
forest. Movement of air and daily exposure of the
soil to the sun along a road through the evergreen
forest also produced conditions intermediate between
the 2 forest types.
In the deciduous forest, gallery vegetation rep-
resented another intermediate situation. Although
green all year, these gallery forests have more air
movement and are less humid than the evergreen
forest proper because of their narrowness and
broken canopies.
Most of our fieldwork in agricultural areas was
in relatively flat lands, mainly planted in rice during
the rainy season, with widely scattered trees, shrubs,
and dead wood. Though no meteorological data
were available for these agricultural sites, we assume,
because of their openness, that the climate approxi-
mates that of the deciduous forest.
March; 8-25 April). On the other hand, maximum
temperature for the entire year in the evergreen
forest was 34.5°C.
Daily minimum relative humidity was consistently
lower in the deciduous forest. During the hottest
part of the year (February through April) relative
humidity fell below 40% every day, whereas equally
low levels were reached in the evergreen forest only
twice. Total evaporation was consistently higher in
the deciduous forest (Fig. 5) and reached extremely
high levels during the February-April interval. In
contrast, total evaporation did not vary seasonally in
the evergreen forest and remained at low levels
throughout the year.
As defined by Colwell (1974), predictability is the
sum of 2 aspects of temporal pattern, called con-
stancy and contingency. An environmental variable
is predictable if it regularly tends toward a level
characteristic of each phase in its temporal cycle, for
example, each hour of the day or each month of the
year. If the levels characteristic of the different
phases in the cycle are equal to one another, the
predictability of the pattern is the consequence of a

Climates of the forests

38 • • •• • •
Although the 2 forest study sites at Sakaerat
are adjacent, their climates near the forest floor
differ considerably. Long-term weather records are
(/)3
::J
•• .. •
••• • •• •
.. • • .•.• • •..• •• • •
~ 30
not available for any of the 3 study sites, but w
standard hourly readings of temperature and total
evaporation were taken at stations in the forest sites
026
(/)

ttla:: 22 ""
•
° \j""
..
<> " "° ° "
RO,9.il\jo".zo
for an entire year (1969). We will use these data
to demonstrate differences between the forests in
@18 " " MAX MIN
{', "
"
microclimatic pattern in terms of absolute levels as
well as predictability.
0
14

10 "
/\
DECIDUOUS
EVERGREEN .
• t.
a €> £l

Temperatures (Fig. 4) in the deciduous forest JAN MAR JUN SEP DEC
equalled or exceeded 35.0°C on 88 days in 1969, FIG. 4. Biweekly temperature maxima and minima in
including 2 long, unbroken periods (12 February-3 evergreen and deciduous forests at Sakaerat, Thailand.
232 ROBERT F. INGER AND ROBERT K. COLWELL
100
. .
(/) 80
. DECIDUOUS ..
W
0:: 6
Iii .. . . .
EVERGREEN.
.. . ...... . ...
~40
...J
...J
~ 20
.. . .... .... .. . . ..
JUN SEP
FIG. 5. Biweekly total evaporation in evergreen and
deciduous forests at Sakaerat, Thailand.
high degree of constancy. If the characteristic
levels are unequal to one another, the pattern may
still be highly predictable, but as a result of the con-
tingency of level on phase.
Since only data of 1 yr are available, the pre-
dictahility, among years, of temperature and evapo-
ration through the seasonal cycle cannot be com-
puted. However, if biweekly means for several years
were known. we suggest that total evaporation would
prove to be less predictable in the deciduous forest,
judging from the great amplitude of fluctuations
within the dry season in that forest for even a single
year (Fig. 5). The predictahility among years for
biweekly temperature extremes cannot he reliably
guessed from Fig. 4.
At the level of daily cycles of temperature and
evaporation. adequate data were available to compute
predictability and its components using the method of
Colwell (1974). To reduce the confounding effect
of seasonal changes, computations were made for
each month based on frequency matrices derived
from readings at 3-h intervals (0300 h, 0600 h,
... , 2100 h, 2400 h) extracted from the raw data.
For each time of day, the frequency of each tem-
perature or evaporation level was compiled, pooling
all days of the month.
Differences in predictability between the two forest
types are striking and consistent (Fig. 6). For each
month. the daily pattern of both temperature and
evaporation is more predictable in the evergreen
than in the deciduous forest. Much higher constancy
in the evergreen forest overshadows the slightly
greater degree of contingency in the deciduous forest.
Annual cycles of predictability and what are probably
unseasonal anomalies (e.g., rains in March) in the re-
gional macroclimate are seen in both forest types
(Fig. 6), yet the predictability of both temperature
and evaporation is consistently greater in the ever-
green forest.
METHODS
Collecting procedures and data acquisition
Between February and December 1969, 4,404
amphibians and reptiles were collected. To provide a
Ecological Monographs
Vol. 47, No.3
framework for randomizing the location of collecting
activities, in each forest type a base line was laid
out and stations established along it at intervals. The
base line in evergreen forest was 0.9 km and the
interval between stations 30 m. In deciduous forest,
the base line was 1.9 km and the interval between
stations 100 m .
In each forest type randomly located quadrats
were thoroughly searched by 5 or 6 persons using
the procedure described by Lloyd et al. (1968). We
examined 479 quadrats, each 7.6 X 7.6 m, in the
evergreen forest and 173 quadrats, 15.2 X 15.2 m, in
the deciduous forest. Larger quadrats were used in
the deciduous forest to compensate for the greater
risk of frightening animals out of the more open
areas. Collecting in quadrats was done only during
daylight hours.
"Cruising collecting" was carried out in mornings,
afternoons, and early night. Five or six people
moved slowly for 2-3 h along a randomly placed
transect = 10m wide. The transects were perpendicu-
lar to the base line, except during night cruising in
the evergreen forest, when the collectors moved
parallel to the base trail. Floor debris was moved,
logs and rocks turned, and trees, shrubs, and bare
soil examined.
We treated streams as special transects because
they were not adequately exploited by the quadrats
or the cruising transects. Four short sections of
watercourses in the evergreen forest were visited
monthly. One was a permanent stream fed by a
spring, 2 were intermittent and 1 never had water
in the months it was visited.
Collections in agricultural areas were taken on
cruising transects with additional night collecting
concentrated on large temporary ponds. Although
collecting was not formally randomized, data from
agricultural land are included in this study to give at
least a rough comparison between this community
and those of less severely disturbed environments in
the same region.
Each captured animal was placed in a separate
plastic bag and its position when first observed re-
corded in terms of a complex microhabitat classifi-
cation (Appendix A). Each animal was killed,
tagged, identified, and preserved, usually within 3 h
of capture. Identifications were verified at the Field
Museum. The specimens are now housed in the
National Centre for Reference Collections, Bangkok,
and the Field Museum of Natural History, Chicago.
Computation of niche metrics
Resource matrices.-To allow quantitative com-
parisons of resource use among species and among
the three communities studied, microhabitat data
were cast into resource matrices, one for each com-
Summer 1977 COMMUNITIES OF AMPHIBIANS AND REPTILES
.8
PREDICTABILITY (P) OF EVAPORATION
DECIDUOUS.
EVERGREEN •
.2
.1~____~____~____~~__~~____~
o .1 .2.3 .5
CONTINGENCY (M)
FIG. 6. Predictability of evaporation (left) and temperature (dght) in evergreen and deciduous forests in Sakaerat,
Thailand.
munity (Colwell and Futuyma 1971). Resource
states (columns) of the matrix were defined by con-
sidering all possible combinations of the microhabitat
characteristics scored upon collection (Appendix A).
The number of possible combinations of the various
states of the habitat variables recognized is nearly
20,000, but only those combinations actually occu-
pied by one or more of the individuals collected
were included in the resource matrices. There were
137 occupied resource states in the evergreen forest,
83 in the deciduous forest, and 59 in the agricultural
land. Of these resource states, only 26 were common
to all 3 environments, the remainder being limited to
1 or 2. The total number of occupied resource
states for all 3 environments combined was 181.
The complete resource matrix for each environ-
ment consists of a column for each occupied resource
state, and a row for each species. The entries in a
given row represent the number of individuals of that
species found in each resource state.
Computations.-We examined the data of the
matrices for differences in resource variety, the
number of occupied microhabitats or resource states
in each environment and the pattern of their occupa-
tion; niche breadth, the degree to which microhabi-
tats jointly support each species; and niche overlap,
the degree to which pairs of species jointly utilize
microhabitats. Given a set of resource states, mea-
233
PREDICTABILITY (P) OF TEMPERATURE
DECIDUOUS.
EVERGREEN.
~
>
U
Z
~ .3
en
z
o
u
.1'-:----__~'::-________=:.::--_ _ _ ___.::.I
.1 .2 .3 .4
CONTINGENCY (M)
sures of niche breadth for a given species are based
on the evenness with which individuals of the species
are distributed among resource states. Indices of
niche overlap for a given pair of species measure the
degree to which the pattern of distribution of indi-
viduals among resource states is proportionally similar
in the two species.
A variety of problems arise in estimating niche
metrics from the data at hand. First, since the
microhabitats of our study are defined in part by
where organisms were actually collected, the number
of occupied resource states increases with the extent
of collecting activity. On the other hand, environ-
ments may actually differ in their essential complex-
ity; arboreal microhabitats, for example, are absent
in cleared areas. Thus a species which is a generalist
may appear to have a narrow niche in a simple en-
vironment or in a poorly collected complex environ-
ment. Certain specialists may appear to have a
relatively broad niche in a simple environment be-
cause few occupied resource states exist. Effects of
environmental complexity and collecting intensity on
estimates of niche overlap are even more difficult to
interpret.
To minimize the effects of these problems, we
limite<il our analysis in several ways. For between-
community comparison of niche metrics, we have
used only the data for those 26 occupied resource
234 ROBERT F. INGER AND ROBERT K. COLWELL Ecological Monographs
Vol. 47. No.3

TABLE 1. Frequency distribution of amphibians and rep- TABLE 1. Continued

tiles in major environmental types at Sakaerat, north-
eastern Thailand Agri- Center
Ever- Deeid- eul- of
Agri- Center green uous tural distri- Hab-
Ever- Deeid- eul- of Taxa forest forest land bution' its'
green uous tural distri- Hab-
Taxa forest forest land bution' its' Agamidae:
AcantllOsaurus 13 E AD
CAEClLIANS armatus
Ichthyophis 2 F? Calotes emma 95 3 E AD
glutinosus C. mystaceus 5 100 22 D AD
C. versicolor 13 152 78 D AD
FROGS Draco maculatus 51 5 E AD
Bufonidae: D. taeniopterus 4 AD
Bu/o 14 25 2 S TN Liolepis 29 6 D TD
me/allostictus belliana
Physignarhus 2 AD
Mierohylidae: cocincinus
Calluella 4 3 4 S FN
Seineidae:
guttulata
Glyphoglossus 10 D FN Dasia olil'acea 3 E' AD
molossus Davcwakcum 14 E F?
Kaloula pulchra 14 9 S TN miriamac
K. mediolineata 3 TN Lipinia 9 E AD
Mieroilyla 5 9 S TN vittigerum
berdmorei Mabuya 6 15 4 S TD
M. butleri 33 4 2 E TN longicaudata
M. heymollsi 71 12 4 E TN M. macularia 44 179 27 D TD
M. inornata 47 2 2 E TN M. multi/asciata 30 2 4 E TD
M. ornata 24 23 62 S TN Riopa howringi 370 339 14 S TD
M. pulehra 11 33 29 D TN Scinecl/a 81 10 E TD
reevesi
Ranidae: S. siamensis 133 15 E TD
Ooeidozyga 40 8 5 E RN Sphcnomorphus 1 TD
laevis indicus
O. lima 35 RN S. maeulatus 75 48 E' TD
Rana erythraea 5 5 RN Laeertidae:
R. lateralis 14 3 D TN
R. limllocharis 12 38 117 D TN Takydromus 4 TD
R. nigrovittata 122 4 7 E RN sexlincatus
R. pilmta 259 2 E RN SNAKES
R. tigrina 14 RN
Typhlopidae:
Rhaeophoridae:
Typhlops bramillus 12 D F?
Chirixalus 8 15 S AN T. diardi 2 F?
Ilollgkhorensis T. khoratellsis F?
C. vitta/us 13 2 20 S AN T. sp.A 1 F?
Poly pedates 35 20 41 S AN T. sp.B 4 F?
leucomystax
Rhacophorus 8 E AN Boidae:
bimaculatus Python
Theloderma 4 E' AN retieulatus A/TN
stellatum
Xenopeltidae:
LIZARDS Xenopeltis F/TN
unicolor
Gekkonidae:
Cosymhotus 2 167 7 D AN Colubridae:
platyurus Ahaetul/a picta 2 TD
Cyrtodactylus 29 E TN A. subocularis 10 D A/TD
angularis Boiga cyanea 6 1 2 S AN
C. intermedius 92 E TN B. multomaculata 2 AN
Gekko gecko 8 6 S AN B.ocel/ata 3 AN
H emidactylus 2 AN Calamaria F?
gamoti pavimentata
H. jrenatus 4 AN Clzrysopelea 2 4 S AD
Hemiphyl- 8 E AN ornata
lodactylus Dryocalamus 4 2 S AN
vunnanensis davisoni
Pe'/,opus 3 125 D AN Dryophis nasutus 3 AD
[aceratus D. prasinus 6 1 2 S AD
P. mutUatus 45 3 E AN Elaphe 1 T?
Phyllodactylus 9 198 49 D TN oxycepllllium
siamensis E. radiata 1 TN
PtycilOZOOIl 29 E AN Ellhydris 18 RN
liona/um plumbea
Summer 1977 COMMUNITIES OF AMPHIBIANS AND REPTILES 235
TABLE. 1. Continued states) in any particular 26-state resource matrix
were subject to computation of niche metrics for
Agri- Center that environment.
Ever- Decid- cul- of
green uous tural distri- Hab- To further minimize the biases of our human-de-
Taxa forest forest land bution' its' fined resource states, we have used the method of
Liopeltis 35 Colwell and Futuyma (1971) to derive a weighting
E T?
scriptus factor for each resource state in each matrix. The
Lycodon aulicus 5 TN weighting factor for a given resource state is pro-
L. fasciatus 2 TN
L.laoensis 3 TN portional to the distinctness of the complete her-
L. subcinctus TN petofaunal list for that state (including all species,
Natrix RN regardless of rarity) compared with the total her-
chrysarga
N. nigrocincta 4 1 1 RN petofauna for the matrix. As the relative abundance
N. piscator 2 3 RN of species in resource state j approaches the relative
N. stolata 1 R/T?
N. subminiata 3 abundance of species for all resource states pooled,
2 R/TD
Oligodoll cine reus 4 3 2 S TD the weighting factor for resource state j approaches
O. cyclurus 2 TD zero (Appendix B). By this weighting, a species
O. quadrilineatus 5 2 4 S TD
O. taeniatus 1 with equal frequency in several very distinct resource
1 TD
Pareas carinatus 3 T? states (high weighting factors) will yield a larger
P. mar- 7 E4 TD estimate of niche breadth than a species found with
garitophorus
Psammodynastes 5 TN equal frequency in the same number of resource
pulverulentus states indistinguishable to animals of the community
Ptyas korros TD as a whole (low weighting factors). Niche overlap in
Sibynophis 6 S TD
col/aris more-distinct resource states is also given extra
Elapidae: weight by this method of computation.
Bungarus TN The sum of all weighting factors for the resource
candidus states in each of our matrices is a measure of the
B. fasciatus 1 TN
Calliophis 2 2
total ecological range of microhabitats sampled. In
F?
macclellandi the extreme case, if the faunas of all resource states
Naja naja TN are proportionally identical, the total range is zero.
Ophiophagus TN
hannah Thus our use of weighting factors (the "absolute"
Viperidae: weighting factors of Colwell and Futuyma [1971])
Agkistrodon 8 D TN helps compensate for any differences in patterns of
rhodostoma microhabitat occupancy not accounted for by the
Trimeresurus 12 2 E AN reduction of the complete resource matrices to the
albolabris
T.popeorunl 21 3 E AN 26 state matrices. For all matrices, "noncircular"
niche metrics (Colwell and Futuyma 1971) were
TURTLES
computed. That is, data for the one (for niche
Cyclemys 16 2 E RN breadth) or two (for niche overlap) species for which
dentata
Testudo T? the metric was to be computed were excluded from
elongata the matrix during the computation of resource state
, Criteria for classification explained in text. E = pre- weighting factors to be used in the expressions for
dominantly or exclusively in evergreen forest. D = pre- niche breadth or overlap. Equations (21) and (24)
dominantly or exclusively in deciduous forest. S = of Colwell and Futuyma (1971) were used in com-
"shared," distribution not clearly centered in either forest.
2 The first symbol refers to habit type: A = arboreal, putations of niche breadths and overlaps, respec-
F = fossorial, R = riparian-aquatic, T =
terrestrial; the tively, with the niche breadth scaling constant k set
second symbol refers to active period: D = diurnal, N =
=
nocturnal, "?" unknown. equal to 10.
3 Though collected only in agricultural land, species al-
most certainly occurs in deciduous forests.
4 Shared according to chi-square criterion, but almost
COMPOSITION OF THE COMMUNITIES
certainly "evergreen" species.
General characteristics
The assemblage of 105 species of amphibians and
states common to all 3 environments, so that three reptiles at Sakaerat (Table 1) is typical of any large
new resource matrices are formed, each being a sub- sample drawn from the subtropical zone (ca. 15 0 -
set of the corresponding within-environment matrix. 25°N) of Southeast Asia. In describing them, refer-
In addition, only those species represented by at ence to ecological or habit types-terrestrial, arbo-
least 13 individuals (half the number of resource real, etc.-represents generalizations of our field
236 ROBERT F. INGER AND ROBERT K. COLWELL Ecological Monographs
Vo\. 47, No.3

TABLE 2. Summary of frequency distribution of amphibians and reptiles in three environments at Sakaerat Experi-
ment Station, northeastern Thailand. N =
no. of individuals

Frogs Lizards Snakes Turtles Totals

Habitat R.S.' Spp." N R.S. Spp. N R.S. Spp. N R.S. Spp. N R.S. Spp. N

Total collection
Evergreen
forest 84 19 725 112 28 1,175 44 29 142 4 16 137 77 2,058
Deciduous
forest 34 20 219 70 18 1,383 28 27 77 3 2 3 83 67 1,682
Agricultural
land 36 19 377 36 15 238 14 19 48 59 55 3 6643
totals 107 24 1,321 139 31 2,796 64 47 267 6 2 19 181 105 3 4,4043
Data from 26 resource states common to three environments
Evergreen
forest 21 15 210 24 24 509 13 24 79 3 15 26 64 813
Deciduous
forest 15 16 180 23 16 925 17 23 55 2 2 2 26 57 1,162
Agricultural
land 13 14 96 21 13 190 10 19 29 26 46 315
totals 23 21 486 26 29 1,624 21 43 163 4 2 17 26 95 2,290
1 R.S. =
number of resource states.
2 Spp. =
number of species.
"Including 2 individuals of the caecilian, Ichtlzyoplzis glu fiIlOSUS, not otherwise listed in this table.

observations supplemented in the cases of some rare
species by statements in the herpetological literature.
Two families, the Microhylidae and Ranidae,
dominate the Sakaerat frog fauna. The 24 species of
frogs, all nocturnal, include only 6 aquatic or
riparian forms, 4 species of Rana and 2 of the ranid
genus Ooeidozyga, reflecting the scarcity of perma-
nent streams and ponds. The same factor probably
accounts for the predominance of terrestrial, non-
riparian frogs, 11 species comprised of 2 of Rana, 6
of Microhyla, 2 of Kaloula (also a microhylid), and
1 Bu/o. In addition, there are 2 fossorial microhylids,
Calluella guttulata and Glyphoglossus molossus, and
5 arboreal species, all members of the Old World
family Rhacophoridae.
Temporary pools, many of them very shallow and
short lived, are important breeding sites for frogs at
Sakacrat (Heyer 1973). Only Rana nigrovittata and
Rana pi/eata, which are essentially confined to small
intermittent streams, seem not to use such situations.
With the exception of a single lacertid, the 31
species of lizards are members of three families:
Gekkonidae, Agamidae, and Scincidae. The 11 spe-
cies of geckos are all nocturnal and most are arbo-
real. Three geckos are primarily terrestrial, Phyl-
lodactylus siall1ensis and two species of Crytodactylus.
All 8 species of agamids are diurnal and all except 1
are arboreal. The exception, Liolepis belliana, is ter-
restrial, occupying cavities in the soil, usually de-
serted termite nests. The 11 species of skinks are
terrestrial except for 2 species: the arboreal Dasia
olivacea and the fossorial Davewakeum miriamae. As
far as is known, all the skinks are diurnal. The
lacertid, Takydroll1us sexlineatus, is diurnal and ter-
restrial though it climbs on grass and low seedlings.
Snakes are more diverse (47 species) at Sakaerat
than either frogs or lizards, but much less numerous
in terms of individuals (Table 2) as befits their
higher station in the trophic scheme. The snake
species are primarily members of the composite
family Colubridae, though 5 other families are also
represented. The 32 colubrids include 1 fossorial, 17
terrestrial, 6 aquatic, and 8 arboreal species. The
aquatic or riparian species, 1 Enhydris and 5 Natrix,
occur along intermittent streams and temporary
bodies of water where they feed on frogs. The 8
arboreal species (5 genera) eat frogs (Chrysopelea),
lizards (Dryophis) , and endothermic prey (Boiga).
The 17 terrestrial colubrid species represent 9 genera,
and include predators on invertebrates (Pareas and
Liope/tis) , frogs (Psammodynastes) , and endother-
mic vertebrates (Elaphe and Ptyas).
There are 8 venomous snakes in the sample, 5
species of Elapidae (cobras and allies), comprising
4 terrestrial and 1 semifossorial species; and 3 pit
vipers, including 2 arboreal species of Trill1eresurus.
The Sakaerat sample also includes 1 caecilian,
Ichthyophis glutinosus, and 2 turtles.
The sample has 2 taxonomic peculiarities. First,
the frog family Pelobatidae, which has 13 terrestrial
species in subtropical Southeast Asia (Bacon 1970),
·was not found at all. We have no explanation for its
absence. Secondly, the number of turtles is small,
considering the relatively great abundance of turtle
Summer 1977 COMMUNITIES OF AMPHIBIANS AND REPTILES 237

species in Southeast Asia, probably because of the TABLE 3. Distribution of species of frogs and lizards by
scarcity of permanent streams on the research site. habit types in three environments at 'Sakaerat Experi-
ment Station, northeastern Thailand
Emydids, the dominant group of turtles of the region,
are primarily aquatic. Riparian
or Ter- Arbo- Fos-
Habitat aquatic restrial real sorial Total
Comparison of communities
FROGS
The 3 samples of reptiles and amphibians differ in Ever-
species richness, taxonomic overlap, and relative green
abundance of species. The evergreen forest yielded forest 4(21%) 9(47%) 5(26%) 1( 5%) 19
77 species, the deciduous forest 67, and agricultural Decidu-
ous
lands 55. These differences are clearly not the forest 3(15%) 11(55%) 4(20%) 2(10%) 20
result of differences in sampling effort; in spite of Agricul-
continuous sampling in all 3 environments, only tural
land 5(26%) 10(53%) 3(16%) 1( 5%) 19
2 new species (of 105 total) turned up in the last
40% (1,762 individuals) of the total collections. LIZARDS
The list of species (Table 1) suggests much tax- Ever-
onomic overlap among the 3 communities: 15 of green
forest 11(39%) 16(57%) 1( 4%) 28
the 24 species of frogs, 10 of the 31 species of Decidu-
lizards, and 6 of the 47 species of snakes were col- ous
lected in all 3. Faunal overlap between the evergreen forest 9(50%) 8(44%) 1( 6%) 18
Agricul-
and deciduous forest is even greater: 17 species of tural
frogs, 17 species of lizards, and 14 species of snakes. land 8(53%) 7(47%) 0 15
The 3 environments are also similar in general
ecological distribution of frogs and lizards (Table 3),
the principal differences being a somewhat higher reliable assignment. Thus we have listed as shared
proportion of arboreal species in the evergreen forest (Table 1) only those species for which chi-square
and of terrestrial forms in the other 2 environments. is less than the autochthony criterion and for which
Our data on snakes as a group are insufficient to the pooled abundance in the 2 forests exceeds 5 indi-
make analogous comparisons with any confidence. viduals.
The data in Table 3 and the tabulation of species With these criteria for determining predominant
occurring in more than one environment exaggerate distributions, the distinction between the faunas of
the similarities among the 3 samples because they the evergreen and deciduous forests is clearly much
fail to take abundance into account. In fact, very greater than was suggested by the simple list of spe-
few species occurring in 2 or 3 environments had cies. Of the 43 species (17 frogs, 17 lizards, 8
similar abundances in each of them (Table 1). If we snakes, 1 turtle) occurring in both forests in numbers
first compare the 2 forest samples, we find that many sufficient for assignment, only 18 are shared (see
species were captured primarily in 1 environment fourth column of Table 1). The percentage of spe-
(e.g., Rana pileata and Peropus laceratus). We con- cies with distributions centered in one environment is
sider such species either predominantly evergreen or significantly higher (X2 = 4.51, P = .03) for ever-
predominantly deciduous, whereas the frog Kaloula green (50%) than for deciduous forest (29%).
pulchra, for example, is more correctly considered as Even if we add to the latter the species collected only
"shared" by the 2 forests (14 in the evergreen, 9 in in agricultural lands but likely to occur in deciduous
the deciduous forest). To make objective assign- forest (see below), the percentage is increased only
ments to these categories, we computed the chi-square to 33%.
contingency statistic for each species, comparing The fauna of the agricultural lands has 3 major
its abundance in the 2 forest types with the pooled components. The largest of these (8 frogs, 3 lizards,
abundance of all other species collected in each forest. 5 snakes) consists of species that appear to be
A high value of chi-square for a given species thus equally at home in all 3 environments. The next
indicates its predominance in 1 of the 2 forests and largest component (3 frogs, 7 lizards) consists of
takes into account the difference in total sample predominantly deciduous species, as defined above,
size (2,058 individuals in evergreen and 1,682 in or species collected only in agricultural land and
deciduous). To be conservative, we have taken P = deciduous forest. The third major component (5
.01, X2= 6.63, as the lower limit for autochthony. frogs, 2 lizards, 2 snakes) consists of predominantly
Species with chi-square <6.63 fall into two cate- evergreen species that were present in agricultural
gories, those which are truly shared between the 2 land only in small numbers and were largely restricted
forest environments, and those too rare to make any to the areas most similar to evergreen forest. Five
 ROBERT F. INGER AND ROBERT K. COLWELL Ecological Monographs
238 Vol. 47, No.3

TABLE 4. Faunal overlap of three communities of am- TABLE 5. Relative abundance of lizard species in cli-
phibians and reptiles from Sakaerat, northeastern Thai- matically intermediate portions! of two forests at
land, in terms of the index proposed by Horn (1966) Sakaer31t, ~ortheastern Thailand. Only the 14 species
occurnng In both forests are included
Communities Frogs Lizards Total '
Predominantly Predominantly
Evergreen-deciduous 0.56 0.53 0.49 evergreen deciduous
Evergreen-agricultural 0.43 0.39 0.39 species (8) species (6)2
Deciduous-agricultural 0.74 0.79 0.61 Number of individuals
1 Including snakes, turtles, and caecilians. in evergreen forest
In clearings 47 32
Elsewhere 425 44
species: the frogs, Rana tigrina and Ooeidozyga =
chi-square 54.92
lima; the lizards, Hemidactylus frenatus and Takydro- P < .001
mus sexlineatus; and the snake Enhydris plumbea, Number of individuals
were collected only in agricultural land but very in deciduous forest
likely occur in deciduous forest in this part of In galleries 71 3
Thailand. The frog, Rana erythraea, was collected Elsewhere 7 918
only in agricultural land and evergreen forest in =
chi-square 849.3
small numbers but almost certainly occurs in decidu- P < .001
ous forest as well. .' In .evergreen f?rc::st, clearings have some of the micro-
Our sample indicates that the deciduous and agri- clImatIc charactenstics of deciduous forest. In deciduous
cultural communities are more similar to each other forest, streamside galleries have some of the features of
evergreen forest.
than either is to the evergreen community and that • Predominant distribution as defined in text.
the last is more similar to the deciduous than to the
agricultural community. These semiquantitative con-
in the preceding comparisons because their habit
clusions are confirmed by application of Horn's
tends to keep them out of clearings and along
(1966) measure of overlap (Table 4) to the data
streams regardless of forest type.
of Table 1.
The 5 predominantly evergreen species of frogs
Collections from wetter parts of the deciduous
caught in agricultural areas also show this tendency
forest (gallery forest) and from drier parts of ever-
to seek out the right parts of an otherwise wrong en-
green forest (clearings) partially account for the
vironment. Of the 20 individuals collected in agri-
relatively high species overlap between the forests.
cultural land, half (7 Rana nigrovittata, 1 Microhyla
Predominantly deciduous species of lizards occur in
butleri, and 2 Microhyla inornata) were caught in
evergreen forest where it most resembles their usual
galleries along permanent streams. One lizard,
environment, while predominantly evergreen species
Scincella reevesi, occurring in agricultural land but
show the same pattern in deciduous forest (Table 5).
with clear predominance in evergreen forest (81: 10),
Table 5 demonstrates the significance of this trend on
was caught only in streamside galleries in agricultural
the basis of total individuals. To exclude the pos-
land.
sibility that a few very abundant species account for
The data in Table 5 also indicate that the move-
this result, we note that the median percentage of
ment of deciduous species within the evergreen forest
individuals in evergreen forest clearings is 49%
is less restricted than that of evergreen species within
for deciduous species and 15% for evergreen species.
the deciduous forest. Those data can be recast to
For galleries in the deciduous forest, the median
show the frequency confined to those portions of the
percentage of individuals is 58% for evergreen spe-
wrong environment resembling the home environ-
cies and 1% for deciduous species.
ment: 42% (32/76) for deciduous species in the
Frogs show analogous patterns, with out-of-place
evergreen forest and 91% (71/78) for evergreen
species living in parts of the "wrong" environment
species in the deciduous forest. The difference is
that are most like the "right" one. Of the four non-
riparian evergreen species, 6 out of 159 individuals
highly significant (X 2 = 41.5, P < .001), and is
supported by the medians of within-species frequen-
were caught in clearings within the evergreen forest
cies (see above): 49% for deciduous species in
as compared to 6 of 23 individuals of the two
clearings of the evergreen forest and 58% for ever-
deciduous species (both nonriparian). In the de-
green species in galleries of the deciduous forest.
ciduous forest, 6 out of 71 individuals of the two
nonriparian deciduous species were found in stream- Geographical affinities
side galleries as contrasted to 11 out of 19 of the four
predominantly evergreen species. To be conservative, Climatic differences between the two forests paral-
we have omitted the three riparian evergreen species lel similar differences on a geographical scale. In
Summer 1977 COMMUNITIES OF AMPHIBIANS AND REPTILES
East Asia, climatic extremes increase and predictabil-
ity decreases continuously from the tropics (0°_
15°N) through the subtropics (15°-25°N) to the
temperate zone (Bacon 1970) . If we divide the
fauna of Sakaerat (at 14°30'N) into "southern" spe-
cies, found no farther north than 20 N, and "north-
0
ern" species, occurring at least as far north as 25°N,
we have a coarse classification of species according
to their tolerance of harsh, unpredictable climates.
The communities of the two forests do not differ sig-
nificantly in terms of proportions of southern or
northern species, but they do differ markedly in
terms of total numbers of individuals of these two
classes. In the evergreen forest, 63% of the indi-
viduals belong to southern species compared to only
48% of individuals in deciduous forest (X 2 = 80.2,
P<.OOl).
Species richness of amphibians and reptiles in East
Asia declines from the tropics northward (Bacon
1970): tropics, 575 species; subtropics, 496 species;
temperate zone, 167 species. At Sakaerat also species
richness drops from the more predictable microcli-
mate to the lesser and at about the same rate.
Another parallel between the large scale geographic
pattern and relationships of the Sakaerat communities
involves percentages of species having distributions
restricted to or centered in a zone or community.
For the entire East Asian fauna, the percentage of
autochthons, in this sense, declines from zone to
zone as the climate becomes less predictable: tropics,
79%; subtropics, 65%; temperate zone, 53% (based
on data in Bacon 1970). Differences between ad-
jacent zones are significant: tropics vs. subtropics
X~ = 24.9, P < .001; subtropics vs. temperate X2 = A similar association explains the high frequency of
8.47, P < .001). At Sakaerat the evergreen forest
community, with the more predictable climate, has
a considerably higher percentage of autochthons
(50%) than does the deciduous forest community
(29%).
AN OVERVIEW OF MICROHABITAT PARTITIONING
Before turning to a purely quantitative and com-
parative analysis of community organization, much
can be learned about the qualitative nature of ecologi-
cal differences among species by grouping data from
the complete array of 181 finely subdivided micro-
habitat types (occupied resource states) into a much
smaller set of habitat categories. Each of the habitats
of Tables 6, 7, 8 and 9 contains the pooled data of
from 10 to 30 of the original resource states. In
Table 3 we divided the fauna inio general habit
types, terrestrial, arboreal, aquatic, or fossorial; here
we look at differences and similarities in microhabitat
use among the species comprising these groups.
Terrestrial lizard species differ considerably among
themselves (Table 6). The skinks can be ordered
239
according to their use of cover. Sphenomorphus
maculatus and the three species of Mabuya wander
freely over the forest floor; high percentages (46%-
74%) of individuals were caught in exposed situations
in both forests. The two species of Scincella are
somewhat more secretive (13%-33% exposed) and
Riopa bowringi the most secretive (4%-9% exposed).
The skink, Davewakeum miriamae, the only fossorial
lizard in our sample (not listed in Table 6), carries
this trend to its extreme. Looking more closely at
the two species of Scincella, Scincella siamensis was
significantly more active in exposed situations (X2
= 5.80; P = .02) and was more frequently associated
with rocks when captured under cover (x 2 = 26.6; P
=
< .001) or exposed (X2 6.97; P .01). =
Similar patterns of microhabitat occupancy by
some of the species of Table 6 conceal important dif-
ferences. For practical reasons, all animals captured
under cover were taken in the daytime. Thus the
diurnal skink Riopa bowringi was probably foraging
and the nocturnal gecko Phyllodactylus siamensis was
resting, although their tabled distributions are vir-
tually identical (x 2 = .04, P > .9) for individuals
under cover in the deciduous forest. Nonetheless,
behavioral interactions between these two species may
have some effect on their activities.
The 2 other geckos, Cyrtodactylus angularis and
Cyrtodactylus intermedius, were not found in suf-
ficient numbers for similar comparisons with diurnal
skinks. However, as clearly portrayed in Table 6, the
2 Cyrtodactylus species differ from one another, the
association of C. intermedius with rocky stream
courses accounting for its frequent capture on rocks.
capture of Sphenomorphus maculatus and Scincella
siamensis on rocks. Alone among these terrestrial
lizards, rather large percentages of C. angularis
(28%) and C. intermedius (36%) were caught on
tree trunks (Table 7).
The terrestrial frogs also differ among themselves,
though not as sharply as the lizards. Microhyla
inornata uses rocks and logs, both as shelters and as
foraging sites more often than its congeners in the
evergreen forest. As a group, the frogs appear to use
the full diversity of the floor structure less thoroughly
than do the lizards; the evenness of distribution
(H'jH'max) of frogs in the columns of Table 6 is less
than that of lizards both in the evergreen (0.82 vs.
0.92) and deciduous forests (0.66 vs. 0.87).
Arboreal species (Table 7) are as heterogeneous
as the terrestrial group. Certain lizards, such as the
species of Calotes, were found in shrubs as well as
trees, whereas others, such as Draco maculatus,
Co!'ymbotus platyurus, and Ptychozoon lionatum,
were essentially confined to medium and large-sized
trees. The pit-viper, Trimeresurus popeorum, and
240 ROBERT F. INGER AND ROBERT K. COLWELL Ecological Monographs
Vol. 47, No.3
TABLE 6. Distributions of abundant' terrestrial reptiles and amphibians at Sakaerat, northeastern Thailand. Indi-
viduals clearly caught in resting sites indicated by italics

Total Below Under cover of: Exposed on:

terres- surface Percent
Taxa trial of soil leaves rocks logs soil leaves rocks logs exposed'
Evergreen forest
FROGS:
Bufo melanostictus 14 1 1 4 5 2
Kaoula pulchra 11 1 4 5 1
Microhyla butleri 29 23 5 1
M. heymonsi 70 28 8 5 23 4 2
M. inornata 38 2 8 6 9 3 2 7 1
M. ornata 21 1 8 1 1 2 5 3
M. pulchra 11 6 4 1
Rana limnocharis 12 4 4 4
LIZARDS:
Cyrtodactylus angularis 21 1 9 5 1 5
C. intermedius 61 4 3 12 1 5 31 5
Scincella siamensis 132 6 32 37 13 8 11 24 1 33.3
S. reevesi 78 7 42 3 12 4 6 2 2 17.9
Mabuya macularia 43 2 11 1 7 1 14 2 5 51.2
M. multifasciata 30 3 2 1 6 1 9 4 4 60.0
Riopa bowringi 364 122 116 18 77 4 21 3 3 8.5
Sphenomorphus
maculatus 74 6 10 3 5 24 25 74.3
SNAKES:
Liope/tis scriptus 35 7 11 11 3 3 17.1

Deciduous forest
FROGS:
Bufo melanostictus 24 6 1 14 2
Microlzyla heymonsi 12 4 1 4 2
M. ornata 23 8 1 4 6 3
M. pulclzra 30 14 5 10 1
Rana limnocharis 37 16 1 4 16
R. lateralis 10 1 3 1 1 4
LIZARDS:
Liolepis belliana 28 22 1 3 1 1 17.9
Phyllodactylus siamensis 192 26 38 37 53 18 4 12 4
Scincella siamensis 15 5 7 1 2 13.3
Mabuya longicaudata 11 2 4 2 1 2 45.5
M. macularia 174 8 50 7 28 62 15 1 3 46.6
Riopa bowringi 332 20 91 84 124 6 4 2 1 3.9
Sphenomorplzus
maculatus 47 3 10 4 8 22 63.8
1 All species represented by 10 or more individuals.
2 For diurnal species.

the frog, Polypedates leucomystax, used seedlings, tall
grass, and shrubs more frequently than trees. Some
species, such as Cosymbotus platyurus and Calotes
emma, were caught primarily at moderate heights
(1-3 m) and others, such as Draco maculatus, mainly
in higher strata. Adults of Calotes mystaceus and
Calotes versicolor display slight, but significant dif-
ferences in vertical distribution (Table 8) analogous
to the interspecific differences mentioned earlier
within the terrestrial genera Scincella and Cyrtodacty-
Ius. As noted above, about one third of the indi-
viduals of the mainly terrestrial geckos, Cyrtodactylus
angularis and C. intermedius were caught low on the
trunks of trees and shrubs. The vertical distributions
of the arboreal capture sites of these two geckos are
distinctly lower than those of the genuinely arboreal
lizards of the evergreen forest (Table 7).
Three arboreal species were caught relatively often
in terrestrial situations or in seedlings or grass < 1 m
high, Calotes emma (25%), C. mystaceus (22%),
and C. versicolor (32%). Significantly greater pro-
portions of the individuals of these species caught on
or near the ground were juveniles (Table 8).
The riparian frogs of the evergreen forest have
differing distributions (Table 9). Ooeidozyga laevis
was found less often at the one permanent stream
than either Rana pileata (X~ = 6.73; P =
.01) or
Rana nigrovittata (X 2 = 3.90; P= .05). Rana pileata
Summer 1977 COMMUNITIES OF AMPHIBIANS AND REPTILES 241
TABLE 7. Distributions of abundant' arboreal reptiles and amphibians at Sakaerat, northeastern Thailand

Seed- Height (m) of capture

Total lings Trees (dbh-cm) in trees and shrubs
arbo- and
Taxa real grass Stumps Shrubs <10 11-30 31-60 >60 ? <1 1-3 >3 max
Evergreen forest
FROGS:
Polypedates
leucomystax 20 8 7 2 2 9 2 6m
LIZARDS:
Peropus mutilatus 24 8 II 5 5 16 2 5m
Ptychozoon lionatum 29 1 1 14 8 5 5 16 7 46m2
A canthosaurus armatus IO 4 2 3 1 1 8 I 3.5m
Calotes emma 71 7 II 22 27 3 15 38 10 7m
C. versicolor 12 1 I 2 7 4 5 6m
Draco maculatus 44 2 1 27 13 I I 2 39 15m
Cyrtodactylus angularis 8 I 3 2 2 6 2 I.5m
C. intermedius 31 9 11 4 4 2 21 10 2m
SNAKES:
Trimeresurus popeorum 14 6 4 3 4 2 5m

Deciduous forest
FROGS:
Polypedates
!eucomystax 14 12
LIZARDS:
Peropus laceratus 17 I 7 2 7 9 l.5m
Cosymbotus platyurus 158 1 2 4 7 89 45 10 40 96 18 5m
Calotes mystaceus 78 3 2 21 8 35 9 15 37 21 12m
C. versicolor 104 36 2 30 4 36 4 2 22 40 4 6m
1 All species represented by 10 or more individuals, except for Cyrtodactylus allgularis.
2 One lizard caught on observation tower in evergreen forest.

was caught oftener in water than R. nigrovittata
(X~ =8.89; P < .001).
The more rigorous microclimate of the deciduous
forest would be expected to force animals in that
forest to spend more time under shelter and less
time in exposed situations compared to animals in
evergreen forest. This expectation is confirmed by
the data of Table 6; 8 of the 9 species (4 lizards, 5
frogs) occurring in the terrestrial zone of both forests
were captured more frequently in exposed situations
in evergreen forest (sign test, P =
.02). Similar re-
sults (X~ = 67.7, P < .001) are obtained by compar-
ing the proportion of individuals in the total samples
caught on exposed surfaces in deciduous (31.4%)
and evergreen forests (46.8%). These results are all
the more significant in view of the fact that sheltering
floor cover is much less extensive in the deciduous
forest (see p. 230).
COMMUNITY ORGANIZATION AND SPECIES RICHNESS
The evergreen forest has more species (77) of
amphibians and reptiles than the deciduous forest
(67), which in turn is richer in species than agri-
cultural land (55) (Table 2). What differences in
environment or community structure permit a greater
number of species to exist III some habitats than in
others?
In comparing microhabitat use by the communities
in our study, correlates of greater species richness lie
in 4 nonexclusive categories: (1) greater diversity
of microhabitats; (2) smaller mean niche breadth;
(3) greater mean niche overlap; and (4) organization
of the community into a number of more or less dis-
crete sets of species, or guilds, with low overlap be-
tween sets, and relatively higher overlap within sets.
Microhabitat diversity
The number of microhabitat categories (resource
states) utilized (Table 2), and presumably the num-
ber available, is much larger in the evergreen forest
(137) than in the deciduous forest (83), and more
in the latter than in agricultural land (59). This pat-
tern alone might account for differences in species
richness, with the "extra" species occurring in the
extra resource states. To evaluate the contribution
of microhabitat diversity, we examine the reduced re-
source matrices described earlier, which include only
the 26 resource states occupied in all three environ-
ments.
242 ROBERT F. INGER AND ROBERT K. COLWELL Ecological Monographs
Vol. 47, No.3
TABLE 8. Vertical distribution of species of Calotes at Sakaerat, northeastern Thailand

Terrestrial Seedlings
Species Stage sites or grass Shrubs Trees x' P

Calotes emma' juveniles 9 5 2 8 12.99

adults 15 2 9 45 <.01
C. mystaceus" juveniles 12 2 12 14
9.03
adults 10 1 9 38 .03
C. versicolor" juveniles 23 21 10 7
adults 25 15 20 29 13.09 <.01

Height (m) of capture of adults

in shrubs and trees
<1 1-3 >3
Calotes mystaceus" 7 21 19 13.94 .001
C. versicolor" 9 36 4
, All individuals from evergreen forest.
" All individuals from deciduous forest.

If similar numbers of species were left in each
of the reduced resource matrices, there would be little
point in searching further for an explanation of the
observed differences in species richness in the three
communities. However, the same pattern of differ-
ences in richness exist among the reduced matrices as
among the original ones (Table 2), with 64 species
left in evergreen forest, 57 in deciduous forest, and 46
in agricultural land. Proportional reduction is not
significantly heterogeneous: X2 = .009, P =
.99.
Niche breadth
According to current theory (Levins 1968; Mac-
Arthur 1972; Leigh 1975), the greater buffering and
predictability of the evergreen forest microclimate
compared to that of the deciduous forest should lead
to a greater degree of specialization (narrower
niches) in the former. Although we lack climatic
data for agricultural lands, presumed extremes and
irregular fluctuations in temperature and humidity
are likely to be even more pronounced there than in
deciduous forest, so that theoretically, niches should
be broadest of all in the agricultural areas. The pre-
dicted pattern, if found, might then account for the
differences in species richness which remain in the
reduced resource matrices.
In fact, there is no significant difference in niche
breadth among the three communities considered
together, although the species of the agricultural com-
munity are significantly less specialized (have broader
niches) than those of either forest community, by
two-sample tests (Table 10). The two forest com-
munities are extremely similar in average niche
breadth. The same general pattern emerges when
niche breadths of lizards or of frogs are considered
separately (Table 10). Differences in niche breadth,
then, fail to account for the greater species richness
of the evergreen forest, although it appears that the
reduced number of species in agricultural lands may
at least in part be associated with broader niches.
These results are not biased by sample sizes (num-
ber of individuals collected per species). Niche
breadth is not significantly correlated with abundance
(P ~ .33 for each data set), and the distributions of
abundances do not differ significantly among the
three sets (Kruskal-Wallis analysis of variance, .05 <
P < .10).
Niche overlap
Several theoretical approaches suggest that more
predictable environments permit greater ecological
similarity (greater niche overlap) among coexisting
species (Levins 1966; MacArthur and Levins 1967;
MacArthur 1972; May 1973). Thus the differences in
climatic predictability documented in Figs. 4-6 sug-
gest greater niche overlap among the species in the
evergreen forest than in the deciduous forest, and
perhaps greater overlap in the latter than in agri-
cultural lands. Since microhabitat variety and aver-
age niche breadth are similar among our environ-
ments (for the reduced matrices), this pattern is also
predicted simply on the basis of differences in species
richness, since overlap should be greater where more
species share a given variety of resources, in the ab-
sence of compensating differences in niche breadth.
If niche overlap values for all possible pairs of
species within each reduced matrix are considered,
there are no significant differences among the three
environments (Table 11). However, not all these
overlaps are of equal importance biologically. Re-
source use by ecologically more similar pairs of spe-
cies is more likely to be subject to evolutionary ad-
justment through the combined effects of competition
and environmental fluctuation. In fact, the theoreti-
cal treatments cited above deal primarily with the
upper limit of niche overlap as a function of com-
petitive interactions and environment.
Summer 1977 COMMUNITIES OF AMPHIBIANS AND REPTILES 243
TABLE 9. Distributions of abundant riparian amphibians TABLE 10. Comparisons of niche breadths of reptiles
in evergreen forest' at Sakaerat, northeastern Thailand and amphibians in three adjacent Thai environments.
Analysis restricted to resource states common to the
Habitat Rana 3 environments, and to species with more than 12
and Rana nigrovit- Ooeidozyga individuals in these reduced resource matrices
occurrence pileata tata laevis
Ever- Decid- Agricul-
Total riparian 255 121 40 green uous tural
Statistic forest forest land
Permanent stream
in water 16 1 All species
on bank" 33 16
Median .2830 .2815 .3508
Temporary stream N' 16 12 7
in water 58 17 10 Analysis of vari-
on bank3 146 87 28 ance (Kruskal-
Wallis) P= .12
Temporary pond 2
Percent at tem- Pairwise tests:"
porary stream 80.0 85.9 95.0 Evergreen P>.lO P=.04
Deciduous P=.04
Percent in water 29.0 14.9 30.0
1 Not enough riparian frogs caught in deciduous forest Lizards only
to warrant tabulation. Median .2977 .2920 .4012
2 Includes exposed rocks in midstream. N 10 7 5
3 Includes exposed rocks in midstream and dried Analysis of
streambeds. variance P =.48
Pairwise tests:
Evergreen P=.50 P=.l1
Guild structure: Statistical analysis Deciduous P=.22

If we conceive of the niches of the species in a Frogs only

community as areas or volumes distributed in a re- Median .2597 .2570 .3359
source space (Hutchinson 1957), then each niche
N 4 5 2
Analysis of
will have a first, second, third, ... , ith nearest neigh- variance P= .21
bor. In general, overlap will be greatest with the Pairwise tests:
first nearest neighbor, less with the second, and so on, Evergreen
Deciduous
P = .36 P =.26
with distant neighbors having little or no overlap. P=.05
Turning the argument around for empirical purposes, 1 Number of species.

we may define the ith nearest neighbor of a given
species as that other species which has the ith greatest
niche overlap with the species in question. In Fig. 7,
we have plotted mean overlap with the first, sec-
ond, . . . , ith nearest neighbor (as defined above) for
the species of each of the three environments. Pro-
cedurally, starting with a symmetrical square matrix
of pairwise overlaps, the elements of each row are
rearranged in rank order; then the mean of the new
elements of each column are the ordinates in Fig. 7.
Looking at niche structure in the three communi-
ties in this way, species in the evergreen forest are
clearly packed more closely than those of the de-
ciduous forest; mean niche overlap is consistently
higher in the evergreen set at all orders of nearness.
In the particularly interesting case of the first nearest
neighbors, the difference is highly significant (.001 <
P < .01, one-tailed Mann-Whitney U-test). The fact
that the curve for agricultural land in Fig. 7 lies above
the curve for deciduous forest for first nearest neigh-
bors is not unexpected, since niche breadths are
greater among the species in the agricultural matrix.
Nearest neighbor analysis is commonly used to
investigate patterns of dispersion in sessile organisms
(Grieg-Smith 1964; Pie lou 1969), when densities are
"Mann-Whitney V-tests, one-tailed; a priori hy-
potheses: evergreen < deciduous; evergreen < agricul-
turalland; deciduous < agricultural land. Corresponding
t-tests yield very similar results.
known, using only first nearest neighbor distances.
The expected distribution of distance between ith
nearest neighbors for randomly placed points in a
plane is known (Thompson 1956), and has occa-
sionally been used in applied work (West 1969;
Hughes 1970). We use Thompson's (1956) formu-
lation to approximate the expected distribution of
overlaps between ith nearest neighbors in niche space,
assuming for the moment that our microhabitat re-
source space is two dimensional. We will assume
that the same area of resource space is occupied
by each of the three communities, a reasonable as-
sumption since all three reduced resource matrices
have the same 26 resource states. Since we have no
way to apply ordinary areal units to this space, we
arbitrarily consider it to be a circle with radius of
one, area 7T. Thus the niche density for each environ-
men.t is Sj7T where S is the number of species for
which overlaps were computed in that environment.
For density Sj7T, Thompson's (1956) distribution
244 ROBERT F. INGER AND ROBERT K. COLWELL
.4
~
~,'<- • (Evergreen)
"',
\ .. (Deciduous)
~
• (Agricultural)
'"
\ \~
'''~..
\
\
\ empirical curve, at second-nearest neighbor for the
3 5 7 9 11 13
NEARNESS ORDER (n)
FIG. 7. Overlap (empirical) of first through Sth near-
est neighbors for three communities of amphibians and
reptiles at Sakaerat, Thailand.
has the convenient property that the expected dis-
tance to the sth nearest neighbor is 1, the expected
distance to all closer neighbors of course being < 1.
Since we need an estimate of overlap rather than
distance, we use one-minus-distance (always between
zero and one) as an index of overlap. This index
may be visualized as the radial overlap between 2
circles of radius 0.5, with their centers on the
neighboring points in the plane.
The curves generated by this procedure are plotted
in Fig. 8. The ordinate, of course, is arbitrary, but
the relative position of the curves closely resembles
those derived from empirical data (Fig. 7). Since
the relationship among the theoretical curves (Fig.
8) is a necessary consequence of differences in
density (s, or species richness), the resemblance to
the empirical curves (Fig. 7) indicates that our as-
sumption of equal resource space for the three data
sets is approximately correct.
More important, the resemblance suggests that
niches in the real communities are arranged more or
less at random in resource space, i.e., that there is
little or no organization in the community. However,
this conclusion can be shown to be false by taking
a closer look at the empirical and theoretical distri-
butions of overlaps. So far we have considered only
observed and expected means. Figure 9 shows the
standard deviation of overlap with the ith nearest
neighbor for the empirical data and for the theoreti-
cal distribution. A very definite peak occurs in each
Ecological Monographs
Vol. 47, No.3
TABLE 11. Comparison of pairwise niche overlaps of
reptiles and amphibians in 3 adjacent Thai environ-
ments. Analysis restricted to resource states common
to the 3 environments
Ever- Decid- Agricul-
green uous tural
Statistic forest forest land
Median .1788 .1718 .2088
N Number of
pairs of species 120 66 21
Analysis of vari-
ance (Kruskal-
Wallis) P = .47
Pairwise tests: 1
Evergreen forest
Deciduous forest
P= .13 P= .24
P= .40
1 Mann-Whitney V-tests, one-tailed; see Table 10, Foot-
note 2.
agricultural set, third for the deciduous forest, and
15 fourth-nearest neighbor for the evergreen forest.
However, the standard deviation predicted by the
theoretical distribution of Thompson (1956) and ap-
plied to overlaps in a unit circle is essentially con-
stant (slightly monotonically increasing) for all or-
ders of nearness and for any niche density. The
difference between the theoretical and empirical
curves is striking, and, we believe, reveals two im-
portant aspects of community organization.
First, the low variance in niche overlap for very
close neighbors supports the idea of a limit to eco-
logical similarity, determined by competition. Sec-
ond, the existence of the peaks in the empirical
curves strongly suggest the existence of a definite
guild structure in our communities. If a small set of .
species are closely similar to one another, their niches
will form a closely overlapping cluster in resource
space. A different cluster in some other region of
the resource space will also have high overlaps
among its own members, but overlaps between each
member of the first cluster and each member of the
second will be very small. Now if clusters differ in
number of members, variance (or standard de-
viation) in niche overlap between pairs of species
should be small for low nearness orders, since most
pairs will share membership in the same cluster. As
the order of nearness begins to exceed the average
cluster size, the variance in pairwise niche overlap
should increase rapidly, as some overlaps are between
members of different clusters, while other overlaps
are still between co-members of larger clusters. When
the size of the larger clusters begins to be exceeded
at still higher orders of nearness, Variance in niche
.overlap should fall, because nearly all overlaps will
be between members of different clusters. This pat-
tern is precisely what we believe to be responsible for
iummer 1977 COMMUNITIES OF AMPHIBIANS AND REPTILES 245

.8 .14

i
1. 12

OVERLAP= 1- ( 1T)
S 'h [n(2nnf)2
(2n) I]
ri",I'/)~ . . . . . :S
II. .08 /7 \ ~ "", Theoretical

~
• s=16 (Evergreen)

"5.4
.. s=12 (Deciduous)
• s= 7 (Agricultural) Ii .06 1/, \
~
i.3 ~ .04
~ "~-\
~
.

II: j! \
~ .2 m.m \

.1
0~1----~3~--~5----~7~--~9----~1~1----~13~--~15
NEARNESS ORDER (n)

0~1----~3----~5----~7~--~9~--~1~1----~1~3--~15 FIG. 9. Standard deviation of niche overlaps (empiri-

NEARNESS ORDER (n) cal and theoretical) between first through Sth nearest
neighbors for three communities of amphibians and rep-
FIG. 8. Overlap of first through Sth nearest neighbors tiles at Sakaerat, Thailand. Ordinate for theoretical
for randomly placed niches in a unit circle. overlaps arbitrary.

the peaks in the empirical curves of Fig. 9. The overlaps) between ith nearest neighbors in n-space,
pattern of the three peaks suggests, moreover, that where i and n may take on any value, with n 2 =
guilds tend to be larger and tighter in the evergreen (Thompson 1956) a special case. The derivation and
forest community than in the deciduous, and larger resulting formulas will be published elsewhere. With
in the latter than in agricultural land. n =1, the variance increases rapidly and monotoni-
To see whether the pattern of peaks in Fig. 9 might cally; for n =
2, the variance is very nearly constant
be a trivial consequence of the method used to mea- (Fig. 9), and for n > 2, the variance decreases
sure niche overlap, "random" communities were monotonically, more and more steeply as n increases.
constructed from the data of the reduced resource Since no peaks appear at any dimensionality, our
matrices, and niche overlaps computed and analyzed
just as for the real matrices. To construct the ran-
domized resource matrices, the elements of each row
(the vector of abundance in the various resource iE
states, for a given species) were reassigned at random .g .10
to the 26 resource states, thereby destroying any in-
ternal organization that may have existed in the
!
• Evergreen
original matrix, while preserving relative abundance .. Deciduous
and the general pattern of niche breadths. With • Agricultural
randomization (Fig. 10), no trace remains of either
the low variance for the closest neighbors or of the z
o
ordered sharp peaks (Fig. 9), which must therefore 1i·04
represent internal organization in the real communi- ~
c
ties.
~.m
The theoretical curves discussed have all been ~
based on Thompson's (1956) derivation of the ex- z
j!
pected distribution for random points in a plane. m 0'~1----~3~--~5----~7~--~9~---1~1~--~1~3--~15
Could the empirical curves of Fig. 9 be closer to the NEARNESS ORDER (n)
expected curves for random points in some higher
FIG.. 10. Standard deviation of niche overlap between
dimensional space? To find out, it was necessary to first through Sth nearest neighbors for randomized em-
derive the expected distribution of distances (and pirical data.
246 ROBERT F. INGER AND ROBERT K. COLWELL Ecological Monographs
Vol. 47, No.3

FIG. 11. The evergreen forest community. Each "bal-

loon" represents the niche center for a species, while the FIG. 12. The deciduous forest community. Here
distance between two balloons is inversely proportional to guilds are less well defined, more linear, and looser.
the niche overlap of the species they represent. Notice the Notice the niche shift in species Wand Y relative to their
aggregation of species into closely packed guilds. The positions in the evergreen community. Species are identi-
axes are the first (horizontal in page plane), second fied in the caption of Fig. 11.
(horizontal behind page plane), and third (vertical)
principal components generated from the symmetrical
matrix of Colwell-Futuyma (1971) niche overlaps for
this community. The species in this figure and in Figs. deciduous forest, which in turn shows more evi-
12 and 13 are: A = Bu/o melanostictus, B = Microhyla dence of guild structure than the agricultural com-
heymonsi, C = Microhyla inornata, D = Microhyla munity.
ornata, E = Microhyla pulchra, F = Rana limnocharis,
Beginning with the evergreen forest community
G = Rana nigrovittata, H = Rana pileata, I = Polype-
dates leucomystax, J = Cyclemys dentata, K = Calotes (Fig. 11), we see a very tight terrestrial guild com-
emma, L = Calotes mystaceus, M = Calotes versicolor, prised of four lizards (U, V, W, Y), a frog (B), and
N = Draco maculatus, 0 = Cyrtodactylus angularis, P a snake (Z), with a second terrestrial frog (C) as a
= Cyrtodactylus intermedius, Q = Peropus laceratus, R satellite. Three other terrestrial lizards (0, P, X)
= Peropus mutilatus, S = Phyllodactylus siamensis, T = form another guild. Three riparian species, two
Cosymbotus platyurus, U = Scincella reevesi, V =
Scincella siamensis, W = Mabuya macularia, X = Ma- frogs (G, H), and a turtle (J), are well separated in
buya multi/asciata, Y = Riopa bowringi, Z = Liopeltis a tight guild. The three remaining species (K, N, R),
scriptus. Species A through I are frogs, species J is a all arboreal lizards, are singletons.
turtle. species K through Y are lizards, and Z is a snake. . The deciduous forest community (Fig. 12) in-
cludes a loose, linear array of terrestrial species, in
three not-very-distinct groups (frogs D, E and lizards
inferences about community structure from Fig. 9
S, Y; frog F and lizard W; frog A). The placement
stand as stated.
of the arboreal lizard Q with the terrestrial species
Guild structure: Graphical analysis
To provide an approximate visual representation
M
of guild structure, we treated the square matrix of
pairwise overlaps for each of our communities as a
matrix of similarity (correlation) coefficients to per-
form a principal components analysis (see p. 468-
508 in Nie et al. 1975). Figures 11-13 show the
niche centers for the species in each community,
plotted in the factor space defined by the first three
principal components. In these figures, Euclidean
distance between the centers of any pair of "balloons"
is inversely proportional to the niche overlap between
the species they represent in the original matrix. (The
size of the balloons is arbitrary.)
The inferences made from nearest neighbor anal-
ysis in the previous section are clearly supported by
FIG. 13. The agricultural community. Species are
Figs. 11-13, on inspection: guilds (clusters) in the widely spaced and not clearly aggregated into guilds.
evergreen forest are larger and tighter than in the Species are identified in the caption of Fig. 11.
Summer 1977 COMMUNITIES OF AMPHIBIANS AND REPTILES
TABLE 12. Numbers and kinds of sets of "similar spe-
cies" of reptiles and amphibians in three adjacent Thai
environments. Analysis limited to those species with
samples large enough (N ~ 13) to warrant calculation
of niche metrics
Ever- Decid- Agricul-
green uous tural
Group forest forest land
Major taxonomic groups:
frogs, lizards, snakes, turtles 4 2 2 between sets.
Congeneric pairs 5 2 2 We start by comparing niche overlap within and
Vertical distribution groups:
terrestrial, arboreal 2 2 2 turtle and 1 snake are included in the niche overlaps
Horizontal distribution goups:
riparian, nonriparian 2
Time-ot-activity groups:
nocturnal, diurnal 2 2 2 represented in the calculations for all 3 communities.
will be treated in the Discussion. The rest of the
species are arboreal; the lizards Land M form a
small guild, with lizard T as a satellite, while frog I
is a singleton.
The agricultural community (Fig. 13) has one
clear guild of terrestrial lizards (S, W, Y), and
otherwise not much structure, although the only two
arboreal species (the lizards Land M) are clearly
separated on the vertical axis from the other species.
Even the tightest clusters in Figs. 11-13 in-
volve species with significant microhabitat differences
among them. By referring back to our discussion on
p. 239-240, many examples can be found.
While n - 1 dimensions are always sufficient to
map n points exactly, given the n X n matrix of dis-
tances between each pair of points, projection of the
points into a smaller number of dimensions may be
possible without too much distortion. The principal
components technique orders the axes of the factor
spacc it creates in such a way that for an m-dimen-
sional representation (m < n - 1), the least distortion
is caused by using the first m axes. In the present
case, examination of planar projections for fourth
and higher dimensions reveals no qualitatively dif-
ferent patterns.
Guild structure: Biological analysis
We showed in the preceding two sections that the
evergreen forest community had larger and tighter
guilds than the other two. We now further cor-
roborate that finding by looking at guild structure in
a more biological way. In the previous section, guilds
were roughly delineated, a posteriori, on the basis of
measured overall microhabitat similarity. Here we
separate each community a priori, or at least not on
the basis of measured similarities, into sets of similar
species. Different sets are formed by applying dif-
ferent criteria, so that, taken together, the sets or
guilds defined on all criteria overlap considerably.
247
Table 12 shows the number of sets of similar species
for each community based on each of five criteria.
On the basis of this list, the evergreen forest com-
munity appears to have more internal organization
than the other two, which do not differ in this regard.
To the degree that the division of these communities
into such sets has ecological significance, its effect
should be evident in greater niche overlap within than
between taxonomic groups. Since only 1 species of
calculated for the evergreen forest community, we
ignore them and focus on the lizards and frogs,
Lizards and frogs considered here feed on inverte-
brates exclusively. Since their ecological roles are
alike in this way, there is some basis for considering
them one large set of similar species. Yet if the gross
differences between frogs and lizards in physiology
and behavior influence their distributions within en-
vironments, one would expect pairs of lizard species
and pairs of frog species to show greater microhabitat
niche overlap than mixed pairs of lizard and frog
species. This prediction is borne out only in the
evergreen community (Table 13).
Congeneric species are generally similar in mor-
phology and behavior and constitute another kind of
similar species. In the fauna as a whole, one would
expect them to have greater microhabitat niche over-
lap than noncongeneric species pairs. The congeneric
pairs in the evergreen community consist of two
groups of frogs (Microhyla and Rana) and three
groups of lizards (Cyrtodacrylus, Scincella, and Ma-
buya). In the deciduous forest and agricultural lands,
the congeneric pairs consist of 2 species of Microhyla
(though not the 2 in the evergreen forest) and 2 of
the lizard genus Calotes. In Table 13, the category
"noncongeneric pairs" is limited in several ways.
First, all pairs are frog X frog or lizard X lizard.
Secondly, at least one member of each pair belongs to
one of the congeneric pairs. Thirdly, all pairs con-
sist of terrestrial species in order to reduce some
of the complicating ecological components. Again,
we find the hypothesis of significantly greater over-
lap within matched than within mixed pairs sup-
ported only in the evergreen community.
We turn now to spatial and temporal distribution.
Some of the lizard species in each major set are
arboreal and some terrestrial, some are nocturnal and
some are diurnal. Again we expect niche overlaps of
mixed pairs to be lower on the average than those
of matched pairs. "Mixed pairs" here refers to ter-
restrial .X arboreal and noctural X diurnal. To avoid
interaction, we limit analysis of period of activity
to species pairs that are matched with respect to
248 ROBERT F. INGER AND ROBERT K. COLWELL Ecological Monographs
Vol. 47, No.3
TABLE 13. Niche overlap within and between taxonomic groups of amphibians and reptiles from three adjacent
Thai environments'

Evergreen forest Deciduous forest Agricultural land

Group and statistic median N median N median N
All taxonomic groups2
Matched species pairs3 .2217 51
Mixed species pairs .0637 69
Mann-Whitney V-test P < .001
Frogs and lizards only
Matched species pairs" .2217 51 .1703 31 .1819 11
Mixed species pairs .0574 40 .1853 35 .2227 10
Mann-Whitney V-test P < .001 P= .26 P= .34
Congeneric groups
Congeneric species pairs .3519 5 .3202 2 .3146 2
Noncongeneric pairs' .2325 22 .2454 8
Mann-Whitney V-test P = .035 P= .20
1 All tests single-tailed. A priori hypothesis--niche overlap of matched pairs greater than that of mixed pairs.
2 Frogs, lizards, turtles, snakes. Last two groups not included in calculations for deciduous or agricultural commu-
nities.
3 Matched species pairs: frog X frog; lizard X lizard .
• See text for limits on these mixed pairs.

vertical distribution. As Table 14 shows, the hy-
pothesis of greater overlap of matched pairs is sup-
ported consistently by the data from the evergreen
community, never by data from the deciduous, and
inconsistently by data from the agricultural com-
munity.
The frogs do not lend themselves to analogous
spatial and temporal comparisons. All of the frogs
are nocturnal. All of those present in the evergreen
sample in sufficient numbers for niche metric analysis
(in the reduced resource matrix) are terrestrial. Two
are riparian and two nonriparian, but testing the
effects of this difference is vitiated by the fact that
each pair is also a congeneric set. The five species
from the deciduous forest are nonriparian; one is
arboreal and four terrestrial. Niche overlap of the
arboreal species with the terrestrial ones is, as ex-
pected, lower (range 0.26-0.39) than overlaps among
terrestrial species (range 0.78-0.95). The two frogs
from agricultural areas are a congeneric pair of ter-
restrial species.
Tables 12-14 provide strong support for the pat-
terns discussed in previous sections. The evergreen
community, in contrast to the others, is subdivided
into tighter guilds characterized consistently by
greater niche overlap within than between guilds.
That the deciduous and agricultural communities
show virtually no significant differences in niche
overlap of mixed and matched pairs of species indi-
cates that they are, in this sense, less organized, and
that the sets of "similar species" in these two com-
munities may merely be convenient artifacts of the
biologist.
DISCUSSION
Because there is no such thing as a standard proto-
col for community analysis, we have had to make

TABLE 14. Niche overlap within and between ecological groupings of lizards from three adjacent Thai environments'

Evergreen Deciduous Agricultural

forest forest land
Vertical stratification2 median N median N median N
Matched species pairs .2362 24 .2053 9 .3430 4
Mixed species pairs .1889 21 .1412 12 .0092 6
Mann-Whitney V-test P= .005 P = .068 P = .005
Time of activity3
Matched species pairs .3586 12 .2826 3 .3471 2
Mixed species pairs .2312 12 .1878 6 .3898 2
Mann-Whitney V-test =
P .037 P= .45 P> .20
1See footnote 1, Table 13.
2 Matched pairs =terrestrial species X terrestrial species and arboreal species X arboreal species.
"Matched pairs = noctural X noctural and diurnal X diurnal. Analysis restricted to pairs of species that are
matched with respect to vertical distribution.
Summer 1977 COMMUNITIES OF AMPHIBIANS AND REPTILES
many simplifications, judgements, and choices of
technique along the way. It is well to ask to what
degree our conclusions depend upon these decisions,
and to inquire whether the sacrifice of precision has
violated reality as well.
In the Introduction and elsewhere, we noted that
our analysis of exploitation of microhabitats did not
separate resting, foraging, and breeding sites. For
some species, such as the abundant lizards Riopa
bowringi, Scincella reevesi, and Scincella siamen.l'i.l'
and the snake LiopeZti.l' scriptu.l', there is very likely
no sharp distinction among the three kinds of sites.
In other instances, though resting, foraging, and
breeding may occur in differl!nt microhabitats, a
group of species may use the same array for thl!se
respective functions, thereby making competition in
all sites possible. This almost certainly holds among
species within genera such as Microhyla, Rana,
CyrtodactyZu.l', Mabuya, and Scincella. In still other
cases, we can probably distinguish between resting
and foraging sites, but some uncertainty remains.
For example, the diurnal, terrestrial skinks, Spheno-
morphu.l' maculatus and the species of Mabuya, are
active foragers on exposed surfaces of forest floor
and litter; although individuals of these species
caught under cover were probably resting, we cannot
assume that these lizards never chase prey into
cover. Separation of microhabitats by function is
further complicated by the unknown effects of the ob-
servers. In some cases, an animal may have been
caught in a site to which it had retreated after seeing
the collectors, and without their knowledge.
Thus both practical and theoretical considerations
justify pooling all microhabitats in our analysis.
This procedure has almost certainly increased niche
breadths and overlaps compared to computations
based on foraging sites alone, with the greatest effect
on species that utilize different sites for different
functions. The greater microclimatic extremes and
the lower predictability of diurnal fluctuations in the
deciduous forest (Figs. 4-6) suggest a more pro-
nounced difference between foraging and resting
sites there than in the evergreen forest, particularly
for terrestrial foragers. Thus the effect of pooling
resting and foraging sites should be greater for the
deciduous forest species. However, this represents a
conservative bias with respect to our conclusions,
since a separation of foraging and resting sites would
differentially decrease niche overlap in the deciduous
forest, where it is already lower for pooled sites.
In the guild analysis, the pooling of all uses of
microhabitats produces some apparent peculiarities.
The most striking case is the placement of the arbo-
real gecko Peropus laceratu.l' at one end of the con-
catenated terrestrial guild in the deciduous forest
249
(species Q in Fig. 12). Thl! explanauon is simply
that this species rests in the floor stratum during the
day, and ascends TO forage at night. In the evergreen
forest, the flying lizard Draco maeulatus (species N
in Fig. 11) is placed near the main terrestrial guild.
That would be biological nonsense except for the fad
that this species, like virtually all agamids, oviposits
seasonally in the soil (Smith 1935; Pongsapipatana
1975). At the phase in the life cycle when females
move into the litter, digging into the soil and perhaps
affecting the lives of other species, this arboreal
species is temporarily a marginal membt:r of the
forest floor guild.
For quantitative analysis of community structure,
we removed from the full re~ource matrix for each
environment all but the 26 occupied resource statl!s
common to all three environments and thl!n limited
niche and guild analysis to only the most abundant
(N ?: 13) spl!cies in the reduced matricl!s. However
logically and statistically sound these procedures may
be, one might well a~k how biologically representative
are the resulting "subcommunities" of their parent
communities, and how truly comparable are the sets
of corresponding resource states.
The species included in niche and guild analysis
(those in Figs. 11-13) are a subset of the species in
the reduced matrices, which in turn are a subset of the
species in the full matrix for each environmt!J1t. Are
these successive samplings biologically biased? It
appears that in fact they are remarkably representa-
tive: there is no significant difference between "uni-
verse," sample, and subsample for anyone of the en-
vironments, or for all three poolt:d, in the proportion
of species by habit type (riparian, terrestrial, arbo-
real, fossorial) or by taxonomic group (lizards,
frogs), or in the total number of species in each
environment (all chi-square tests yield P > .71).
Furthermore, the reduced matrices are indeed
comparable. The distribution of individuals among
analogous resource states, for all species pooled, is
similar in pattern for all three reduced matrices
(Kendall coefficient of concordance W, P = .005).
This rather surprising result suggests that, even in
these contiguous communities, ecological replace-
ment and compensation are operating, since the
faunas of the three communities differ not only in
species composition, but in richness. It also reassures
us that collecting efforts were apportioned among
resource states in the same way in all three environ-
ments.
The Colwell-Futuyma measures of niche breadth
and overlap have been dismissed as unnecessary by
Sabath and Jones (1973) on empirical grounds,
because, for their data, niche metrics computed for
weighted and unweighted resource states were highly
250 ROBERT F. INGER AND ROBERT K. COLWELL
correlated for species in a single community of
drosophilid flies. This statistical result, first of all,
is probably an unavoidable consequence of their in-
clusion of many very rare species, which of course
cannot yield anything but small estimates of niche
breadths, regardless of method; the unweighted niche
breadths they present are correlated with the corre-
sponding relative abundances at P < .001, Spearman
rank correlation. Within a community, weighting
factors provide only the "fine tuning" in any case;
in Sabath and Jones' (1973) results, the rank order
for weighted and unweighted niche breadth differs
by up to five units (among 20 species), in spite of
the overall correlation, and in spite of the general
homogeneity of their resource states. Most important,
however, their results have no bearing on comparison
amung communities, since the summed heterogeneity
of resource states, which may differ greatly among
communities, directly affects the magnitude of all
weighted niche metrics.
In our data, the weighting factors ranged over
more than an order of magnitude in each community,
and in addition the total heterogeneity among re-
source states varied between communities (evergreen:
.47; deciduous: .37; agricultural: .49), so that un-
weighted measures would clearly have given quite
different distributions of niche breadth and overlap
than weighted measures, both within and between
communities. In general, resource states with the
largest number of individuals have the highest weight-
ing factors, because they usually represent a great
departure from the average fauna. This is a property
of nature, and not a necessary consequence of the
mathematics of weighting factors (Appendix B).
Thus, in a sense, resource states are weighted accord-
ing to their abundance (or habitability) in the en-
vironment (Schoener 1974). Since our resource
states were defined somewhat arbitrarily (if care-
fully) in the first place, these abundances and their
associated weighting factors could be changed simply
by pooling or further subdividing resource states,
a fact that simply points out the proper functioning
of the method. For example, one of the resource
states with the highest weighting factors (using all
species in the reduced matrix) was "away from
water, on a tree trunk, 11-30 cm dbh, between 1
and 3 m above ground level" (rank-order 1 in de-
ciduous forest, 4 in evergreen, and 5 in agricultural
land) . Had we split up this resource state into 2
vertical layers, for example, the weighting factors
for the 2 new resource states would likely be lower,
so that the niche breadth of a species not distinguish-
ing between them would be relatively unchanged.
Our conclusions concerning guild structure based
on nearest neighbor analysis are unaffected by any
monotonic scaling change in any or all of the com-
Ecological Monographs
Vol. 47, No.3
munities, as the pattern of peaks in Fig. 9 depends
only upon the relative magnitude of variation in
overlap with increasingly distant neighbors within
communities. Similarly, the biological analysis of
guilds by mixed and matched pairs is independent of
any change of scale, since it, too, concerns only
within-community comparisons.
The graphical portrayal of guild structure (Figs.
11-13), while not intended to be statistically rigorous,
quite accurately represents the relative distances to
the first few nearest neighbors based on niche over-
laps. Dendrograms have also been used to represent
niche overlap matrices (e.g., Cody 1974). However,
since communities, unlike taxa, are not fundamentally
hierarchical in organization, nonhierarchial methods
are called for (Everitt 1974).
In extensive comparative studies of desert lizard
communities, Pianka (1974, 1975) finds a negative
correlation between species richness and average
pairwise niche overlap. The correlation persists even
if only the top 10% of overlaps are considered. Since
this is not equivalent to examination of the overlap of
each species with its nearest neighbor, Pianka's re-
sults cannot be compared directly with our own.
However, because his conclusions as stated appear
to contradict ours, it is important to examine other
differences in some detail. Since resource diversity is
strongly correlated with species richness for desert
lizard communities, it is impossible to tell from
Pianka's published data what the pattern of overlaps
would look like (or even whether any differences in
richness would remain) if the effect of differences in
resource diversity were removed and niche metrics
computed over the domain of resource space com-
mon to all three continents. The effect of including
overlaps between distant neighbors is to lower aver-
age pairwise niche overlap, so that in a community
with peripheral niches beyond the bounds of this
common resource space, the average will be de-
pressed to an even greater degree, perhaps obscuring
an opposite effect of tighter packing among very
close neighbors.
Although Pianka (1975) finds a positive relation
between species richness and environmental predict-
ability, as we do, his conclusions concerning environ-
mental predictability and niche overlap cannot be
compared with our own, as a result of the problems
just discussed. Based on mathematical models, May
and MacArthur (1972) and May (1974) predict
that random environmental variability should have
little effect on maximum tolerance niche overlap.
We will argue in the final section that differences in
the predictability of rather extreme periodic fluctu-
ations (day-night and seasonal cycles) do have im-
portant effects on tolerable microhabitat overlap, and
on community organization as well.
Summer 1977 COMMUNITIES OF AMPHIBIANS AND REPTILES
CONCLUSION
The message of experimental community ecology
is that interactions among species are important in
the patterning of communities (Colwell and Fuentes
1975). In other words, while the physical-chemical
environment demands certain minimum physiological
c~edentials for entrance into a community, every spe-
cies must pass the much more demanding criteria
imposed by the biotic environment: the necessities of
finding food, coping with enemies, and reproducing
successfully.
In this study, the action of major climatic factors
as faunal filters is reflected in the composition of the
regional herpetofauna, in the distribution of species
among adjacent communities in the landscape (ever-
green forest, deciduous forest, agricultural land),
and in the association of smaller assemblages of spe-
cies with isolated macrohabitat patches (clearings in
evergreen forest, gallery vegetation in deciduous
forest). We showed that species of the deciduous
forest community, which has a northern cast, pene-
trate evergreen forest mainly in clearings. On the
other hand, species of the evergreen forest com-
munity, which has strong southern (or tropical)
affinities, penetrate deciduous forest almost ex-
clusively in gallery vegetation.
Biotic interactions have their major effect within
communities. As community development involves
the simultaneous interaction of many species, we may
envision a process of iterative adjustment of the na-
ture and intensity of interactions, with the interactive
network eventually approximating some equilibrium
structure. However, for organisms with a generation
time commensurate with the period of major fluctu-
ations in the physical environment (e.g., small verte-
brates in a highly seasonal environment) the pattern
of interactions may change continuously over an
annual cycle, and probably varies from year to year
to a degree dependent on the seasonal predictability
of the physical environment.
In this study we have used detailed data on micro-
habitat use, pooled over seasons, to examine the
pattern of association of species in three environ-
ments differing in predictability. Thus. in our results
a very low niche overlap between species means that
they rarely use the same microhabitats, either se-
quentially or simultaneously. Therefore, such species
very likely have neutral functional interactions. A set
of highly overlapping species, on the other hand,
may have a variety of functional interactions: com-
petition, predation, mutualism. or more complex re-
lationships, but are much less likely to be neutral.
We showed that the pattern of microhabitat over-
lap is more structured in climatically more predict-
able environments, with more discrete sets (guilds)
of highly overlapping species. The significance of
251
these results, in our view, is that greater predictability
permits an approach to a unique equilibrium of
spatial and functional relationships. On the other
hand, the continual perturbations inflicted by a less
predictable environment impose a never-ending series
of ecological adjustments, or at best a series of
transitions from one equilibrium to another, so that
each species must cope with nearly every other spe-
cies in the community at some time and in some way.
The coalescence of a small set of species .into the
same pattern of microhabitat use in a predictable en-
vironment has two consequences. First, if other sets
of species in the same community converge on other
characteristic patterns of microhabitat use, then each
species has the overwhelming proportion of its non-
neutral interactions with only the members of its own
guild. Second, the organization of a community in
this manner apparently permits a greater total num-
ber of species to coexist, given an equivalent array
of resources. This is an empirical result, which does
not follow necessarily from geometric arguments.
MacArthur's idea of "diffuse competition" (Mac-
Arthur 1972; Pianka 1974) suggests a trade-off be-
tween the number and the intensity of interactions
among coexisting species. Imagine an unstable com-
munity with many species, in which each species
has strong interactions of some kind with every other
species. We contend that the outcome of adjustment
of the interactive network, by extinctions and niche
shifts, takes a course dependent on environmental
predictability. In an unpredictable environment, after
a large number of extinctions from our hypothetical
community, most species would still interact with
each other, but only to a moderate degree. At
equilibrium in a predictable environment, on the
other hand, there would be a coalescence of sets of
species into guilds, with strong interactions within
and weak interactions between members of different
guilds.
We envision these alternative patterns of com-
munity organization as ends of a continuum corre-
lated with environmental predictability. Validation
or refutation of this model awaits further empirical
studies, as well as theoretical investigation of the sta-
bility properties of communities structured in dif-
ferent ways.
ACKNOWLEDGMENTS
During most of the field year, the crew was under the
direction of Dr. W. Ronald Heyer: we are indebted to him
for the energy and care expended in the collection of
specimens and data. The Applied Scientific Research
Corporation of Thailand (ASRCT) kindly granted us
permission to use the Sakaerat Experiment Station. We
also received transportation support from the Earth
Sciences Laboratory. U.S. Army Natick Lahoratories
(NLABS).
We are particularly grateful to Dr. Prasert Lo-
252 ROBERT F. INGER AND ROBERT K. COLWELL
hav:J.nijaya (ASRCT), and Messrs. George Immish,
Rohert Fegley, and Joseph Zabransky (NLABS) for
many personal kindnesses while the party was in Thai-
land. We also thank Dr. Mary-Claire King for her help
with extending the nearest neighbor distribution to n
dimensions. Miss Betty Peyton for her patient typing of
numerOllS drafts of the manuscript, Mr. Shahid Naeem
for drawing Figs. 11-13, and Mr. Robert Magnaval for
assist.ance with data analysis.
This work was partially supported by grants from
the National Science Foundation (GB 7845X and GB
31195) ilnd by the University of California at Berkeley
Comonter Center.
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Appendix A: Habitat classification
Based upon previous experience in rain forest and
other environments in Borneo (1950-63), and with
the inspiration of Elton's (1966) habitat classifica-
tion, we prepared a habitat code prior to fieldwork
in 1969. Since the general system was designed for
application in any vegetation in tropical Asia, flexi-
bility was built into the system by making the cate-
gories expandable. For example, we have used an
expanded version of this system in rain forest en-
vironments of Malaya and Borneo. As presented
here, only those elements encountered at Sakaerat
are listed.
In the following list, each general microhabitat
variable, in italics, is followed by a list of alternative
categories for that variable. Certain of the categories
listed below were further subdivided; we made
arbitrary groupings of tree diameters (dbh < 10 cm,
11-30 cm, 31-60 cm, >60 em), heights above
ground «1 m, 1-3 m, >3 m), and depths below
surface of soil «20 em, >20 cm). In general, each
individual collected was assigned to one of the al-
ternative states for each variable.
VeRetation: deciduous dipterocarp forest; dry
evergreen forest; agricultural land. Horizontal po-
sition: in water of permanent stream; mid-stream on
Summer 1977 COMMUNITIES OF AMPHIBIANS AND REPTILES 253
bar, rock, or snag in permanent stream; on bank of Appendix B:
permanent stream; in water of intermittent stream; Weighting factors for resource states
mid-stream on bar, rock, or snag of intermittent Demonstration that if the total fauna of a resource
stream; on bank of intermittent stream; in dried bed matrix is a multiple of the fauna of resource state j,
of intermittent stream; in water of temporary pond; then weighting factor OJ = 0. Notation as in Colwell
on bank of temporary pond; above water at tempo- and Futuyma (1971).
rary pond; away from streams or ponds. Vertical The condition of the premise is that Nij/Xj =
position: on surface of soil; under surface of soil; on ¥jZ for all i, with j constant. Transposing, we have
the surface on dead leaves; under leaves; on rock; N;iY i = Xi/Z, so that we may define A = N;i¥i =
under rock; on log; under log; in decaying log; on Xi/Z. The expression for the weighting factor OJ
stump; in grass; on seedling or herbaceous plant (from Eqs. 16 and 17 of Colwell and Futuyma 1971)
«1.0 m height); on shrub or young sapling (woody is
plant <7 m); on tree or large vine (plant >7 m).
Substrate: on leaf; on stem of herbaceous plant; on
OJ = (~i 7Ti}Og Pij - Pjlog Pj)/H(X);
twig or branch of woody plant; on trunk of shrub,
OJ = [~i (N;iZ)log(N;iYi )
- (Xj/Z)log(Xi/Z)]jH(X).
tree, or stump; on epiphyte; under bark of log or
stump. Special: permanent pond away from streams; Under the premise,
temporary rain pool away from streams; tree hole;
OJ = [~i (Nii/Z) log A - (Xj/Z)logA]jH(X);
earthworm burrow or other burrow; small forest
OJ = [(log A)/Z)](~i Nij-Xj)/H(X).
clearing.
A microfiche copy of the data set may be obtained However, since by definition Xj = ~i N ij , we obtain
by writing the senior author. OJ = 0, which was to be demonstrated.

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