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Transcript of A Comparison of Amphibian Communities Through Time and Place to Place
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Journal of Trop ical Ecology (1993) 9:409-433. With 2 figures
A ciimparison of amphibian communities
through time and from place to place in Bornean
forests
ROBERT F. INGER
and
HAROLD K. VORIS
Field Museum of Natural History, Chicago, Illinois, USA
ABSTR ACT. We sampled riparian frogs along 18 streams at eight localities in Borneo. At four of
these sites we samp led during more than one year. Altogether 49 spe cies were includ ed in our
study an d total sam ple size was 13,249. We measured overlap in spe cies occurrence s and arrays
of abund ances within and among loca lities. Variation over the time span of our study was minor
within com mun ities. Overlaps between streams at a locality were generally higher than overlaps
of pairs of streams from different loca lities. Environmental variation, particularly in stream width
and gradient, had a clear effect on both intra-and inter-locality overlaps. Although rainfall varied
between loca lities and within loca lities over time, that variation did not seem to affect overlaps
among or within com mun ities. Environmental factors did not accoun t for all differences in overlaps
between comm unities. Instead, regional proce sses, perhaps the timing of barriers or spec iation
events, appear to have been respon sible for geograph ic restrictions of several specie s, leading to
variation in overlap values.
KEY WORDS: anurans, Borneo, commun ity structure, community variation, Malaysia, rain
forests.
INTRODUCTION
When one looks at community structure within a relatively small area, the
significance ecologists attach to local phenomena, such as biotic interactions
and species-specific responses to environmental features, is understandable.
These are the most apparent phenomena, even if they are not always easy to
tease apart and evaluate. However, as Ricklefs (1987) observed, features of local
community structure may be strongly affected by regional processes, such as
barriers to dispersal, the history of speciation, and other phylogenetic events.
For certain aspects of community structure viewed at a regional scale, Hanski
( 1982) proposed a model relating distribution of species across sites to their
abundances. This model assumes that species populations fluctuate in stochastic
fashion over time in response to biotic interactions and environmental fluctu-
ations. In effect, local processes lead to regional patterns. A study by Gascon
( 1991) on larvae of Amazonian frogs shows the inter-play of local and regional
processes. These tadpoles had species-specific responses to abiotic habitat char-
acteristics and Gascon’s data give hints of biotic interactions, e.g. scarcity of
409
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410
ROBERT F. INGER AND HAROLD K. VORIS
co-occurrence of larval species, a typical local process. Yet the restriction of
‘
two larval forms to stream microhabitats is clearly a product of phylogenetic *
history. These two, Atelopus p&her and Centrolenella oyampiensis, belong to genera
J al l of whose larvae develop in streams (Duellman & Lynch 1969, McDiarmid,
1978) regardless of the distributions of other sympatric tadpoles of other genera.
4
Most studies of community structure in tropical amphibians have concen-
trated on a single, relatively restricted area (e.g. Crump 1971, Gascon 1991,
Heyer 1973, Inger 1969, Inger & Colwell 1977, Toft & Duellman 1979). That
concentration perhaps explains the primary focus on local processes. In this
paper we examine variation among Bornean amphibian communities across an
area approximately 700 km wide, which obliges us to consider regional processes
as well as local ones.
Our analysis is restricted to that segment of the Bornean amphibian commun-
ity occurring along rain forest streams, species that spend their entire li fe cycles
in this riparian habitat or that utilize streams for breeding and larval develop-
ment. We begin with examination of variation over a 22-year period at one site,
Nanga Tekalit, Sarawak (Figure 1, site 1 ), which provides a standard with
which to evaluate variation over space. We then consider factors associated
with variation from place to place.
1100
114O
1180
I
I I
I
1 I
Figure 1. Map show ing the eight local ities in the Malaysian states of Sarawak (numbers l-4) and Sabah
(numbers 5-8) on the island of Borneo. The localities are (1) Nanga Tekalit, (2) Segaham, (3) Pesu, (4)
Labang, (5) Mendolong, (6) Purulon , (7) Marak Parak, and (8) Danum.
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Variatibn among frog communities
411
SAMPLING SITES
Our sampling of stream frogs was conducted at eight localities in rain forests
below 800 m in Borneo in the Malaysian states of Sabah and Sarawak (Figure
1, Table 1). Distances between localities range from 35 to 640 km (Table 2).
Full details of these sites are given in Appendix A.
Sites differed in several ways:
(i) Topography. The only flat area was Labang (site 4, Figure 1). The rest
were hilly (e.g. Danum, site 8) to steep (Purulon and Segaham, sites 2 and
6).
(ii) Elevation. All except Purulon and Mendolong (site 5) were below 300 m
asl. The streams at Purulon are at 320-370 m and the one at Mendolong
at 750 m.
(iii) Vegetation. The entire area of Mendolong included in this study had been
selectively logged. The forest at Marak-Parak (site 7) was old (45-50 y)
secondary growth that now has a high closed canopy. Labang was covered
with flat, alluvia l forest. Most of the area at Pesu and all the area at the
remaining sites was well-drained, hil ly, and covered with primary diptero-
carp forest. There was a small area of swamp forest at Pesu (site 3), and
both Danum and Nanga Tekalit (site 1) had a few flat areas.
(iv) Rainfall. Amount of rainfall during sampling periods varied greatly from
site to site. The four in Sarawak had the most precipitation, with few
months having 300
mm.
(v) Streams. Regardless of amount of rainfall, al l streams in this study were
perennial. All became turbid after heavy rain, but only Sungai Seran at
Labang was continually turbid. Sungai Seran was also the only low gradi-
ent stream and the only one with a silt bottom. Stream widths vary from
3 to 25 m. Streams less than 8 m wide were under canopy, those 8-10 m
partially under canopy. Banks of even the largest streams were under trees.
MATERIALS AND METHODS
Field procedures
At each site work was divided between effort along streams and in the forest
proper. Data from the non-riparian work, which consisted of forest floor plots,
search of buttress-enclosed areas, and night transects through the forest, are
not used in this paper, but contribute to an understanding of the total fauna at
each site. The riparian work included collecting and observing tadpoles by day
and collecting and observing frogs during night transects. The data from the
night transects along fixed segments (250600 m) of streams form the basis of
this paper. On streams at four of the localities, we marked stations at 15-30 m
intervals with plastic flagging and recorded the position of each frog observed
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Table 1. The number of night stream transects and the number of anurans observed each year on each stream.
Locality Stream
First sample period Second sample period Third sample period
Year Transects Anurans Year Transects Anurans Year Transects Anurans
Labang Seran 1963 29 477
Nanga Tekalit Ensurai 1962 36 747 1970 5 295 1984 5 241
Sekentut 1962 36 772 1970 5 206 1984 5 247
Selubok 1962 15 640 1970 5 232
Set-bong 1962 36 1038 1970 5 377 1984 5 252
Lawan 1962 17 469
Wong 1962 12 319 1984 2 51
PeSU Pesu 1964 78 1775
Segaham Segaham 1984 13 612
Marok 1984 10 331
Danum Cabin 1989
4 97 1990 7 179 fz
P. Tambun 1986 5 476 1989 4 88
1990 7 306
S.Kalison 1989 7 194 1990 9 267
E;
W6S5 1986
4 213 1989 4 ’ 71 1990 4 122
206
F
Surinsin 1988 4
c
c
Marak Parak Mendolong 1987 5 234 1989 5 327 1990 5 291 ;
6 284
;:
Mendolong Kilampo n 1989
7 322 1990
Purulon Purulon 1989
6 257 1990 7 234
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Variation among frog communities
413
Table 2. Distances (km) between local ities in Sarawak and Sabah.
Nanga
Tekalit Pesu Segaham Danum
Marak ‘.
Parak Mendolong Purulon
Labang 180 35 77 565
485 295 340
Nanga Tekalit 166 125 640 615 425
475
Pew 56 540
475 285 330
Segaham 555 500 305
350
Danum 210 275
250
Marak Parak
185 145
Mendolong
45
relative to the stations. We recorded or captured every frog seen as we waded
upstream (see Appendix B). The Sungai Seran at Labang (see Appendix A)
was too silty for wading and was searched from boats.
At Nanga Tekalit in 1962/63 we marked, released, and recaptured frogs on
three streams - Ensurai, Sekentut, and Serbong. To avoid the bias of counting
the same frog many times, we used the number of individuals seen (either in a
quarter or during the entire year) rather than number of observations of those
species marked and released.
Table 1 gives the number of transects and number of individuals per stream
per year. Intervals between transects for Nanga Tekalit in 1962/63 are given
elsewhere (Inger & Greenberg 1966). Intervals in other years and at other
places varied according to the lengths of the sampling periods (see Appendix
A).
Statistical analysis
We relied on two measures of overlap to assess the similarity between
samples. The first of these, overlap of abundance arrays, uses Morisita’s index
as modified by Horn (1966)
C = 2CnunJ(h, + h,)N,Nz
where nji = number of individuals of species i in sample j and
hj = cnj,‘/Nj2
Wolda (1981) found this index to be relatively independent of sample size and
diversity.
The second measure of overlap, of species present in two samples, uses the
Czekanowski coefficient (Wolda 198 1) :
overlap (or similarity) = 2c/(S + L)
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414
ROBERT F. INGER AND HAROLD K. VORIS
where c = number of species common to the two samples
S = number of species in smaller sample
L = number of species in larger sample
As Wolda pointed out, most of the binary similarity coefficients have some
weaknesses, but the Czekanowski increases linearly from zero to 1.0 as c
increases, and is simple to calculate.
We have the impression that our estimate of the abundance array of the
community on a stream approaches the ‘true value’ after about five transects.
We have tested this assumption in the following way. We calculated the overlap
between two of the most heavily sampled streams at Nanga Tekalit (site 1,
Figure I), Serbong, and Ensurai, using one transect drawn at random from the
36 on each stream in 1962/63. We then recalculated the overlap after adding a
second transect drawn at random, repeating the process until we had overlaps
based on five randomly drawn transects. We followed this procedure twice.
Overlaps based on a single transect from each stream were 0.555 and 0.549 in
the two sets, increasing to 0.774 and 0.842, respectively, when two transects
were drawn from each stream, and reaching 0.812 and 0.839 after five transects.
As the last values are 95-98 of the value (0.85) obtained from the full data
set from each stream, we believe that our samples from every stream (see Table
1) in the study were adequate for our purposes.
For inter-locality measures of overlap, we summed data for a stream across all
years of sampling before calculating overlaps of abundance arrays with another
stream. We assessed the effect of this procedure on the overlaps between the
Palum Tambun at Danum, Sabah (site 8, Figure l), and each of the three
streams at Nanga Tekalit, Sarawak, sampled during three years. Consequently,
for the overlap of Palum Tambun (sampled 1986, 1989, 1990) with Sekentut
(sampled 1962, 1970, 1984) we had nine new overlap values, and nine between
Palum Tambun and Ensurai and between Palum Tambun and Serbong. For
the Palum Tambun/Sekentut pair, the mean of the nine overlaps was 0.61
(SD = 0.11) compared with 0.65 when all years for a stream were combined.
For Palum Tambun/Serbong the mean of nine was 0.5 1 (SD = 0.14) compared
to 0.51 for years combined. The mean of nine for Palum Tambun/Ensurai, 0.44
(SD = 0.1 l), diverged further from the value for years combined, 0.54, but still
close enough to justify combining years.
RESULTS AND DISCUSSION
Over short (within locality) or long distances (between localities), variation
among these riparian frog communities might appear because of differences in
general topography or vegetation, rainfall regimes (between localities), or phys-
ical characteristics of streams (gradient, width, bottom characteristics, etc.).
However, before accepting any observed differences between a pair of commu-
nities as reflecting inter-community variation, it is necessary to consider that a
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Variation among frog communities
415
single community might vary over time. If the variation between two communit-
ies is no greater than the variation over time within one of them, then the
observed difference between the two communities may have only transient bio-
logical significance. We therefore begin our analysis with an examination of
variation within communities.
Our analysis has the following organization: (i) Variation over time within
a stream, thus holding stream characteristics (width, gradient, etc.) and general
environment constant, but allowing for the effects of temporal variation in rain-
fall and species’ behaviour over shorter (year) periods.
(ii) Variation between streams at a locality, thus holding general environment
(topography and vegetation) relatively constant but allowing for effects of differ-
ences in stream characteristics. (iii) Comparison of the effects of time and dis-
tance within a locality. This comparison is restricted to four streams at Nanga
Tekalit that are similar in gradient, width, and bottom types, thus minimizing
the effects of stream characteristics. (iv) Variation between communities over
greater distances (i.e. at different localities), thus allowing for effects of differ-
ences in general environment (topography and vegetation, rainfall) and in
stream characteristics (width, gradient, etc.).
Variation within localities
Nanga Tekalit (site 1, Figure 1). This is our largest data set in terms of numbers
of individuals, numbers of streams, numbers of transects, and length of observa-
tion time (Table 1). We use variation in it, among other things, as a standard
against which to evaluate variation at other localities and overlaps between
localities.
(i) Variation over time. If the assemblage of species and individuals on a
stream varies over time, there should be gradual, if not continuous, divergence
from an initial observed state. Under this hypothesis, within-stream overlaps
of abundance arrays and species occurrences between adjacent quarters of a
year should be larger than overlaps between non-adjacent quarters. We test
this prediction with the data from 1962/63 on the four larger streams at Nanga
Tekalit (Table 3), the streams for which we have most data during that year.
Table 3. O verlap of abundance arrays and of spe cies occurrences between quarters within streams at Nanga
Tekalit, Sarawak, in 1962 and 1963.
Overlap of Abundan ce Arrays Overlap of Spe cies Occurrences
Quarters
Ensurai
Sekentut Serbong Selubok Ensurai Sekentut Serbong Selubok
1x2 0.92 0.93 0.98 0.84 0.81 0.73 0.83 0.76
2X3 0.98 0.93 0.82 0.84 0.80 0.86 0.92 0.84
3x4 0.96 0.96 0.91 0.93 0.87 0.96
1X3 0.91 0.94 0.86 0.90 0.73 0.74 0.81 0.85
2X4 0.94 0.91 0.81 0.73 0.79 0.88
1x4 0.90 0.89 0.86 0.73 0.80
0.79
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416
ROBERT F. INGER AND HAROLD K. VORIS
These are also the streams most similar in width, bottom types, and gradient.
Overlaps of abundance arrays between adjacent quarters were slightly greater
than between non-adjacent quarters,
but the difference is not significant
(Mann-Whitney test, single-tailed, P = 0.09). However, overlaps of species
occurrences between adjacent quarters are significantly larger than those
between non-adjacent quarters (Mann-Whitney test, single-tailed, P = 0.025).
Of the 16 species absent from these streams in one or several quarters (i.e.
those that depressed between-quarter overlaps of species occurrences), nine
were represented by
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Variation among frog communities 417
Tab le 5. Overlap of abundan ce arrays and of spe cies occurrence s between streams at Nanga T ekalit, Sara-
wak. The rows for Selubok and Lawan are omitted due to empty cel ls in 1970 and 198 4.
Overlap of Abund ance Arrays Overlap of Spe cies Occurrences
1962 Sekentut Serbong Selubok Lawan Wong Sekentut Serbong Selubok Lawan Wong
Ensurai
0.96 0.85 0.92 0.52 0.38
0.76
0.78 0.76 0.74
0.89
Sekentut 0.87
0.88 0.53 0.29 0.78 0.80 0.64 0.70
Serbong
0.85 0.69 0.29
0.80 0.62 0.73
Selubok
0.45 0.28
0.71 0.76
Lawan
0.42
0.82
Overlap of Abund ance Arrays
Overlap of Spe cies Occurrences
1970 Sekentut Serbong Selubok Lawan Wong Sekentut Serbong Selubok Lawan Wong
Ensurai 0.71 0.71 0.77 0.73 0.89 0.67
Sekentut 0.68
0.84
0.80 0.79
Serbong 0.77
0.79
Overlap of Abund ance Arrays
Overlap of Spe cies Occurrences
1984 Sekentut Serbong Selubok Lawan Wong
Sekentut Serbong Selubok Lawan Wong
Ensura i 0.82 0.79
0.55 0.81
0.87 0.59
Sekentut 0.95
0.41
0.76
0.69
Serbong 0.42
0.58
Tab le 6. Overlap of abunda nce arrays and of spe cies occurrence s between streams at Nanga Tekalit, Sara-
wak. Observations for each stream are summ ed over all years.
Overlap of Abund ance Arrays
Sekentut Serbong Selubok Lawan Wong
Overlap of Spe cies Occurrences
Sekentut Serbong Selubok Lawan Wong
Ensura i 0.92 0.83
0.95 0.43 0.44 0.83 0.88
0.79 0.85 0.94
Sekentut 0.88 0.90 0.42 0.35 0.91 0.86 0.73 0.85
Serbong
0.87 0.38 0.33
0.86 0.84 0.90
Selubok
0.45 0.37 0.73 0.81
Lawan
0.45 0.83
Serbong (Tables 5 and 6), despite the fact that Selubok and Sekentut were
much closer to Wong (see Appendix A).
Wong and Lawan were only half the widths of the other four streams and
had much lower overlaps of abundance arrays with the four larger ones than
the last had with each other (Table 6). Except for width, Lawan was very
similar to Selubok and Sekentut, having a bottom mainly of sand and gravel,
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418
ROBERT F. INGER AND HAROLD K. VORIS
with open pools, rifIles, and side pools. In terms of microhabitats known to be
used by five dominant species (Inger 1969), Rana blythi, R. ibanorum, R. chalconota,
R. signata,
and
Pedostibes hosei,
for larval development (Inger
et al.
1986) and for
adult perch sites (Inger 1969), Lawan should have had very high overlap of
abundance arrays with Sekentut and Selubok. Its low overlap of abundance
arrays with those streams suggests that it provided a significantly different
environment, particularly for the larger species, such as R. b&hi and R. ibanorum.
These two species constituted 6 and 5 , respectively, of the Lawan sample
but 23-25 and 12-22 of the samples from Sekentut and Selubok (see
Appendix B; authorization for species’ names is also given here). These large
species also had low abundances (3 and 7 ) respectively) on Wong.
However, more than stream size was involved, for the two small streams,
approximately the same size, had a low overlap of abundance arrays with each
other. Wong had the steepest gradient of all the streams at Nanga Tekalit,
which may explain why
Lefitobrachella mjobergi,
a riflle breeding species (Inger
1985), was one of the more abundant species on Wong but was seen only once
on three of the larger streams and not on the others. Staurois natator, a frog that
in our experience perches on rocks or leaves overhanging streams, constituted
25 of the sample from Wong, but less than 1 of the other five samples.
Overlaps of species occurrences did not show a dichotomy between the small
and large streams. Although there was not complete correspondence between
the species seen on the small streams with those on the large ones, the streams
were close enough for any species in the area to reach them. Species that did
not or could not maintain sizeable populations on the small streams, i.e. those
that depressed overlap of abundance arrays, none the less contributed to overlap
of species occurrences.
(iii) Comparison of the effects of time and the unique qualities of each stream.
The Nanga Tekalit data set allows comparison of small-scale time and distance
(as represented by the distinct assemblage on each stream) effects. For this
purpose we use data only from the four larger streams to increase the time
range (Table 1) and to avoid the complicating effects of differences in stream
size. Overlaps of abundance arrays between streams within quarters (range
0.69-0.91, median = 0.84) were smaller than overlaps between quarters within
streams (median = 0.91, Table 3) (Mann-Whitney test, two-tailed, P C 0.01).
At these scales, between-stream effects contributed more to variation than time.
Expanding the time scale changed the results. The overlaps between years
within streams (Table 4) did not differ from overlaps between streams within
years (Table 5) (M ann-Whitney test, two-tailed, P > 0.10). Overlaps of species
occurrences between streams within time intervals (quarters or years) did not
differ from overlaps between time intervals within streams (Mann-Whitney
tests, P > 0.10).
Other multi-year and multi-stream samples.
Within-stream overlaps of abundances
at l-4 y at Danum (site 8) (Table 7), Purulon (site 6; 0.87, 0.93), and Men-
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Variation among frog communities
419
Table 7. Overlap of abundance arrays and of spec ies occurrences between years within
streams at Danum, Sarah.
Years
Overlap of Abund ance Arrays
Palum Sapat
Cabin Tambun Kalison W6S5
Overlap of Spec ies Occurrences
Palum Sapat
Cabin Tambun Kalison W6S5
1986x 1989 0.85 0.68 0.77 0.72
1986x 1990 0.96 0.72 0.83 0.76
1969x 1990 0.90 0.80 0.74 0.64 0.83 0.76 0.86 0.83
dolong (site 5; 0.84, 0.93, 0.94) ranged from 0.64 to 0.96 (median = 0.85). As
a set, they are smaller than the overlaps between quarters at Nanga Tekalit
(Mann-Whitney test, single-tailed, P = 0.03), but did not differ from those
between longer intervals at Nanga Tekalit (Mann-Whitney test, P > 0.20).
Within-stream overlaps of species occurrences between years at Danum, Puru-
lon (0.74, 0.84), and Mendolong (0.65-0.86) did not differ significantly from
those at Nanga Tekalit (Mann-Whitney tests, P > 0.10).
Between-stream overlaps of abundances and of species occurrences at Puru-
lon (0.97 and 0.89, respectively) were much higher than between streams at
Danum (Table 8). The two streams at Purulon joined near the origins of the
surveyed sections and were very smiliar in size, gradient, bottom substrate, and
frequency of microhabitat types.
At Danum between-stream overlaps of both abundance arrays and species
occurrences (Table 8) fell within the range of those among streams at Nanga
Tekalit. As at Nanga Tekalit, distance between streams at Danum, varying
between 1 and 12 km, did not account for differences in overlaps of abundances
between streams. The Palum Tambun was much farther from the Cabin stream
and Sepat Kalisan than from the stream at W6S5, yet had higher overlaps with
the first two. Physical differences between these streams are not clearly related
to their overlaps. All streams had mixtures of pools, riffles, and torrents, though
W6S5 had a steeper gradient. Danum was the only site besides Nanga Tekalit
where the data permitted a comparison of time and distance effects. Between-
Table 8. Overlap of abundance arrays and of spec ies occurrences between streams at
Danum, Sabah.
Overlap of Abund ance Arrays
Palum Sapat
Tambum Kalison W6S5
Overlap of Spec ies Occurrences
Palum Sapat
Tambun Kalison
W6S5
Cabin 0.73 0.65 0.84 0.76 0.83 0.74
Palum Tambur 0.83 0.65 0.83 0.90
Sapat Kaliso n 0.57 0.82
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ROBERT F. INGER AND HAROLD K. VORIS
year, within-stream overlaps both of abundance arrays and species occurrences
(Table 7) were significantly higher than between-stream, within-year overlaps
(Mann-Whitney tests, two-tailed, P < 0.05).
Summa? of in&a-locality variation
(1) Within-stream overlaps (abundance arrays and species’ occurrences)
between short intervals (3 months) were very high (tested at Nanga Tekalit
only).
(2) Within-stream overlaps between years were also high (four localities), but
at Nanga Tekalit were not as high as overlaps between 3-month intervals.
(3) Between-stream overlaps (abundance arrays and species’ occurrences) were
strongly affected by stream width and gradient.
(4) If stream characteristics did not vary, overlaps (abundance arrays and
species’ occurrences) between streams within years did not differ from over-
laps between years within streams. This comparison was possible only at
Nanga Tekalit.
Variation between localities
Values of inter-locality, between-stream overlaps of abundance arrays had a
very broad range, 0.01-0.83 (Tables 9, 10, ll), and represent a significant
shift downward from intra-locality overlaps. Inter-locality overlaps of species
occurrences followed the same pattern; the range was broad and 81 of the
values fell below the range of intra-locality overlaps.
Effects of stream character.
Between-stream overlaps of abundance arrays at
Nanga Tekalit were affected by stream width. Stream width appears to have
been an important factor in inter-locality overlaps as well. We have grouped
streams (Table 1) into four size categories (map numbers in parentheses): width
3-6 m: Marok (2), Wong (l), Lawan (1); width 7-9 m: Purulon (6), Kilampon
Table 9. Overlap of abundance arrays and of spec ies occurrences between streams between loca lities in
Sarawak. Stream data were sum med over all years.
Overlap of Abundance Arrays Overlap of Spec ies Occurrences
Segaham Pesu Labang Segaham Pesu Labang
Locality Stream Segaham Marok
Pesu Seran
Segaham Marok Pesu Seran
Nanga Ensurai 0.37 0.10 0.54 0.28 0.72 0.60 0.60 0.47
Tekalit Sekentut 0.29 0.12 0.70 0.36 0.78 0.64 0.63 0.49
Selubok 0.29 0.12 0.44 0.30 0.68 0.59 0.59 0.43
Serbong 0.24 0.09 0.59 0.26 0.70 0.57 0.60 0.46
Lawan 0.45 0.22 0.25 0.26 0.60 0.42 0.55 0.39
Wong 0.22 0.27 0.22
0.16 0.71 0.66 0.62 0.38
Segaham Segaham 0.26 0.12 0.60 0.44
Marok 0.07
0.04 0.56 0.41
Pesu Pesu
0.67 0.61
.
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Table 10. Overlap of abundance arrays and of species occurrences between streams between localities in Sabah. Stream data were summed over all years.
s
2.
OVERLAP OF ABUNDANCE ARRAYS OVERLAP OF SPECIES OCCURRENCES
z?.
s
Mmak Mendolong Purulon Ma&& Mendolong
Purulon s
Locality Stream Surinsin Mendolong Purulon Kilampon surinsin
Mendolong Pundon Kilampon
s
T
Danum Cabin 0.21 0.65
0.83 0.83 0.45 0.67 0.59 0.61 2
P. Tambun 0.19
0.39 0.57 0.55 0.54 0.63 0.67 0.63
S. Ka.lison 0.09 0.37 0.56 0.54
0.40
6
0.65 0.58
w6S5 0.18
0.60
0.49
0.80 0.78 0.48 0.72 0.65
g
Marak Parak
0.67
0.15 0.29 0.24
0.67 0.71 s
Mendolong
0.60
0.68 0.63 0.84
N.
0.80 -g.
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Table 11. Overlap of abundance arrays and of species occurrences between streams between localities in Sabah and Sarawak. Stream data were summed over all
years.
OVERLAP OF ABUNDANCE ARRAYS
SABAH LOCALITIES
Danum
Mamk Mendolong Purulon
Sarawak
Localities Stream Cabin P. Tambun
S. Kaiison W6S5 Surinson Mendolong Purulon Kilampon
Labang Seran 0.19 0.21 0.20 0.08 0.01
Nanga
0.07 0.02
Ensurai 0.30 0.54
0.01
0.48 0.15 0.02 0.09
Tekalit
0.03
Sekentut 0.34
0.02
0.65 0.63 0.18 0.03 0.09
Selubok
0.04
0.30 0.54
0.04
0.45 0.16 0.02 0.09
Serbong 0.30
0.03 0.02
0.51 0.57 0.16 0.02 0.09
Lawan
0.04
0.38 0.29
0.04
0.30 0.33 0.02 0.15
Wong
0.10
0.27
0.07
0.27 0.52 0.30
Pesu
0.04
Pesu
0.28
0.24
0.25
0.42
0.18
0.54 0.14 0.03 0.06 0.04
0.06
Segaham Segaham 0.16 0.14 0.17 0.11 0.09 0.07
Marok 0.20
0.10
0.27
0.08
0.16 0.44 0.03 0.09 0.17
0.14
OVERLAP OF SPECIE S OCCURRENCES
SABAH LOCALITIES
Danum
Mart& Mendolong Purulon
Sarawak
Localities Stream Cabin P. Tambun S. Kalison
W6S5 Surinson Mendolong Purulon Kilampon
Labang Seran
0.38 0.43 0.44 0.40
Nanga
0.15
Ensurai
0.65
0.28
0.67
0.24
0.68
0.28
0.70 0.43
Tekalit
0.60
Sekentut
0.65
0.53
0.67 0.63
0.60
0.60 0.52 0.54
Selubok
0.49 0.62
0.56 0.54
0.59 0.55 0.39 0.49
Serbong
0.62
0.46
0.59
0.65
0.49
0.57 0.42 0.56
Lawan
0.61
0.49
0.59
0.51
0.65 0.56 0.40
Wong
0.68
0.50
0.65
0.42
0.71
0.50
Pew
0.64
Pesu
0.46 0.62
0.56
0.60
0.49
0.66
0.60 0.51 0.33
Segaham
0.45
Segaham 0.61
0.42
0.59
0.55
0.40
0.51 0.40
Marok
0.50
0.41
0.53
0.50
0.45
0.51 0.52 0.36 0.51 0.54
0.47
I
rlrt
~.
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Variation among frog communities
423
(6), Cabin (8), W6S5 (8), Sepat Kalisan (8); width 10-14 m: Pesu (3), Seran
(4), Mendolong (5)) P
a urn Tambun (8), and the four larger streams at Nanga
Tekalit (1); width 25 m: Segaham (2). If stream width accounts for a significant
part of the inter-locality variation, overlaps between streams within size categor-
ies should be greater than overlaps between streams across size categories.
Overlaps (grouped into four classes:
cO.21, 0.21-0.40, 0.41-0.60, BO.60)
within-stie categories were significantly greater than those across size categories
(chi-square = 19.4, df = 3, P < 0.01); 57 of the within-size category overlaps
(n = 30) exceeded 0.40 compared to 17 of the between-size overlaps (n =
82).
Between-stream overlaps at Nanga Tekalit showed some effect of differences
in stream gradients. If differences in gradients affect inter-locality overlaps,
overlaps between streams of similar gradients should exceed those between
streams of differing gradients. We grouped streams into the following gradient
classes from steepest (A) to flat (E); A: M arok, Purulon, Kilampon, Mendolong;
B: Surinsin, Segaham; C: W6S5, Wong; D: Pesu, Lawan, Ensurai, Sekentut,
Selubok, Serbong, Cabin, Palum Tambun, Sepat Kalisan; E: Seran. Overlaps
were grouped as in the preceding paragraph. Overlaps within gradient classes
were significantly larger than those across classes (chi-square = 26.31, P <
0.01); 53 of the within-class overlaps (n = 30) exceeded 0.40, in contrast to
14 (n = 99) of the between-class overlaps.
Inter-locality overlaps of species occurrences gave slightly different results.
Overlaps within-and between-width classes did not differ (chi-square = 4.94,
P = 0.30). However, overlaps within gradient classes were significantly greater
than overlaps between those classes (chi-square = 12.44, P < 0.02). We inter-
pret these results as indicating that stream gradient had a greater effect than
stream width on the occurrences of species.
The effect of stream gradient is best seen on the Seran, the only stream
flowing through flat forest and the only one with turbid water and a silt bottom.
These circumstances almost certainly account for the absence of any of the 15
species (27 of 55) in our data set that breed at riffles and torrents and have
larvae that either attach to rocks or wriggle into interstices between rocks and
gravel on stream bottoms. These 15 include all five species of
Amolops
(Inger
1966), three species of Ansonia (Inger 1992) four species of Leptobrachella, and
three species scattered through other genera (Inger 1985, and unpublished
data). An additional five species absent along the Seran have been observed
elsewhere only at turbulent, rocky areas of other streams: Micrixalus baluensis,
Philautus hosei, Rana hosei, Staurois latopalmatus, S. tuberilinguis (unpublished data).
The two streams at Segaham (Figure 1, locality 2) show interaction of stream
width and gradient on overlap. Although the two streams, Marok (3 m) and
Segaham (25 m), joined where they were sampled, their overlap of abundance
arrays was only 0.07 and, though their overlap of species occurrences was higher
(0.65) it was still lower than the overlaps at Nanga Tekalit. The five largest
species at that locality were much more abundant on the Segaham (41 of the
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424
ROBERT F. INGER AND HAROLD K. VORIS
sample) than on the Marok (8 ). Both are high gradient streams. However,
the Marok consists of a series of small waterfalls separated by short pools over
gravel and rock, whereas the Segaham has long stretches of foaming rapids over
large boulders, the kind of habitat that supports populations of adult Rana hosei
and Amolops cavitym~anum and is used by tadpoles of A. cavi~mpanum and Bufo
juxtasper. The last two species were five times more abundant and Rana hosei 20
times more abundant on the Segaham than on the Marok (Appendix B).
Effects of rainfall patterns. Another environmental factor that may have played
a role is the pattern of local rainfall. Variation in rainfall locally may have
affected overlaps over time within streams, and systematic differences between
localities in amount and seasonal distribution of rainfall may have affected
inter-locality overlaps. Several features of the climate of Borneo are important
here: (1) At any given site, in the great majority of years there are no months
totally lacking rain. For example, over a 30-year period at Melalap Estate, a
recording station 10 km from Purulon (site 6), no month lacked rain and only
two had 25 mm, which was enough
to cause spates on the observation streams, were distributed at random through
the year (Lloyd et al. 1968). Rainfall was moderately heavy during the other
sampling periods at Nanga Tekalit and overlaps of abundance arrays between
years remained high (Table 4). The fact that sampling periods at Nanga Tekalit
over the years did not occur during the same calendar intervals appears to have
had little effect. Within-stream, between-year overlaps using data from the
entire year 1962/63 did not differ from overlaps using data only from those
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Variation among frog communities
RAIN IN
mm (1887)
500
w MELALA P u MENDOLONG w DANUM
J F MAMJJASOND
MONTH
RAIN IN mm
(1908)
1000
+ Nanga T ekallt 1963 q MELALAP
JFMAMJJASOND
MONTH
RAIN IN m m (WSO)
700
td MELALA P i-i MENDOLONG
n
DANUM
J F M A M J JASOND
MONTH
4 5
Figure 2. Monthly rainfall data for three years at three localiti es in Sabah. Melalap is 8 km from our
sampled locality Purulon. The middle graph gives the monthly rainfall for Nanga Tekalit (Sarawak) in
1962163.
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426
ROBERT F. INGER AND HAROLD K. VORIS
parts of 1962/63 matching the periods of work in 1970 and 1984 (Wilcoxon
matched-pairs signed-ranks test, P > 0.05).
Although rainfall varied appreciably from year to year at Danum (Figure 2)
and during sampling periods, there was no association between rainfall and
amount of overlap within streams (Table 7). At Purulon rainfall during the
second sampling period was less than 30 that in the first period (see above),
yet overlaps between years were very high (0.87, 0.92). Similarly, at Mendolong
between-year overlaps were high (0.84-0.94) despite variation in rainfall.
If differing amounts of rainfall (either at the time of sampling or annually)
affect inter-locality overlaps significantly, overlaps between localities having
similar rainfall should be larger than those between localities differing in
amounts of rain. To test this hypothesis, holding stream gradient and width
constant, we compare overlaps of abundance arrays between the four larger
streams at Nanga Tekalit and Pesu (site 3), both areas with high rainfall, with
overlaps between those streams and Palum Tambun at Danum (about half as
much rain). Differences between the two sets of overlaps (see Tables 9 and 11)
do not differ significantly (Mann-Whitney test, P > 0.20). All other inter-
locality tests of rainfall effects are confounded by differences in gradient or
stream width. However, overlaps of abundance arrays between Purulon and
Mendolong (twice as much rain) were high (BO.60) despite differences in
stream width. Also overlaps between Purulon streams and those at Danum
(twice as much rain) were also high (0.54-0.83, Table 10) despite differences
in gradient.
Restricted geographic distributions.
In all the preceding analyses, a tacit assump-
tion is that all species are available at all localities. There is evidence, on the
contrary, that some species have limited geographic distributions within
Borneo. For this part of the analysis, we use all frogs observed at a locality,
whether on a surveyed stream or not, to establish presence in an area. General
distribution is based on Inger ( 1966), Inger & Dring ( 1988), Inger & Stuebing
(1992) and Matsui (1986). Ab un d antes and distributions of species mentioned
below are given in Appendix B.
One of the dominant species along streams in hilly areas of Sarawak,
Rana
ibanorum,
is currently known only from Sarawak. Intensive, repeated search at
Danum (site 8) in streams that provide appropriate habitats (clear streams
having beds of sand, gravel, and rock) and at other similar streams 100 km
south and 175 km west of Danum (field work by Inger) have failed to uncover
this species in eastern Sabah.
Bufo asper,
abundant on streams at all four localit-
ies in Sarawak, was also absent at Danum and the eastern Sabah localities
referred to above, again, despite the presence of appropriate habitats. Other
abundant stream-side species that appear to have geographically restricted dis-
tributions, despite wider availability of suitable habitat, include Amolops phaeom-
eluS (found in central Sarawak) and
A. whiteheadi
(western Sabah). Altogether,
11 species (20 )
in our data set depress inter-locality overlaps because of
geographically, as distinguished from ecologically, restricted distributions.
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Variation among frog communities
427
Do these restricted ranges reflect historical processes or competitive interac-
tions with ecologically similar species? We look for evidence of biotic interac-
tions within groups of closely related species, which, as they are usually ecolo-
gically homogeneous, are most likely to show competition. There were eight
such groups in our sample, four of which were present in sufficient numbers
for statistical analysis.
1. Rana ibanorum co-occurred with two related large species along streams at
Nanga Tekalit (site 1) (Inger & Greenberg 1966). All three were absent from
four streams in western Sabah that lacked suitable microhabitat. Correlations
of relative abundances of these species in the 14 streams where at least one
was present were positive. This suggests that the absence of
R. ibanorum
from
appropriate habitat in eastern Sabah is independent of the distribution of the
other two species.
2. The two large species of
Bufo,
usper and
juxtasper,
which form a distinct
species group (Inger 1972), co-occurred on seven streams. As their abundances
were negatively correlated (Spearman r, = -0.57, P < O.Ol), the absence of B.
asper from its usual habitat in eastern Sabah could be the result of competition.
3.. The two species of Amolops having geographically restricted distributions
(see above),.each co-occurred with an abundant congener, poecilus in the case of
phaeomeru&nd orphnocnemis in the case of whiteheadi. Larvae of phaeomerus and
poecilus
have been collected together in tadpole stations (Inger 1985) as have
tadpoles of the other two (unpublished data). Relative abundances of
whiteheadi
and
orphnocnemis
were positively correlated, whereas abundances of
phaeomerus
and
poecilus
were negatively correlated though not at a significant level (P >
0.05). The distributions of
whiteheadi
and
phaeomerus
were not completely comple-
mentary as neither occurred in eastern Sabah.
4. The three species of Staurois (Appendix B) co-occurred in four streams and
in pairs or singly in the remaining 14 streams. Their relative abundances were
positively correlated, which makes competition an unlikely explanation of the
absence of S. tuberilinguis from streams at Danum and other streams in eastern
Sabah that lie outside our study area.
The other groups having one or more species with geographically restricted
ranges had small sample sizes (Appendix B). They include: three species of
slender toads,
Ansonia,
one widely distributed and two absent from eastern
Sabah though sympatric at two localities in Sarawak; two species of horned
frogs,
Megophy,
one apparently restricted to central Sarawak and sympatric
with the second which was observed across the study area; and three species
of small pelobatid toads, Leptobrachella, with one restricted to Sabah and two
to Sarawak. As a genus, Leptobrachella exhibits strong altitudinal stratification
(Inger & Stuebing 1992), suggesting that in this group competition may be
important, if not universal. The other two groups show no evidence that biotic
interactions play a role in their geographic restrictions.
According to Hanski (1982)) for many organisms there is a positive correla-
tion between local abundance and regional distribution; i.e. species tend to be
either locally abundant and widespread (‘core species’) or locally rare and
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428
ROBERT F. INGER AND HAROLD K. VORIS
regionally scarce (‘satellite species’). An earlier paper (Inger 1969) on the frogs
of the streams at Nanga Tekalit noted that nine species accounted for >85
of observations and each contributed >3.5 of the sample. Hanski’s model
predicts that these nine species should have been equally abundant at Danum,
where streams were very similar to those at Nanga Tekalit in width, gradient,
and array of microhabitats. Yet at Danum these nine species constituted
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Variation among frog communities 429
breed in clear, turbulent water, i.e. roughly a quarter of the species in our
study.
There were significant differences within and between localities in rainfall,
both on an annual basis and at the times of sampling. Yet this variation did
not account for differences in overlaps among communities.
Although environmental factors had strong effects on inter-locality overlaps,
they do not account for all of the variation. At least 11 species have geographic-
ally restricted ranges that cannot be explained on the basis of distribution of
suitable microhabitats. Nine of these co-occur with at least one similar congener
at one or more localities. Regional processes (sensu Ricklefs 1987), perhaps the
existence or timing of barriers to dispersal or the timing of speciation events,
appear to be responsible for their geographic restrictions rather than biotic
interactions.
ACKNOWLEDGEMENTS
We wish to express our thanks to men of the Iban longhouse, Rumah Jimbong.
Without their able assistance, none of the work at Nanga Tekalit would have
taken place. We also wish to acknowledge Lucas Chin, Director, and Charles
Leh, Zoologist, both of the Sarawak Museum, who arranged for government
permits and eased many logistical problems. We are grateful to the authorities
of Sabah Parks, Sabah Forest Industries, and Yayasan Sabah for permission to
work in areas under their respective jurisdictions and for living accommoda-
tions. We are also grateful to Sabah Parks and Universiti Kebangsaan Malaysia
(Kampus Sabah) for provision of camping equipment, transportation, and
logistical assistance. We thank R. B. Stuebing, F. L. Tan, and P. Yambun for
assistance in the field and for many kindnesses. Professional colleagues, J. P.
Bacon, S. Emerson, K. J. Frogner, W. Hosmer, D. Karns, F. W. King, J. C.
Murphy, and P. Walker, helped us collect at various times. We are grateful to
M. Lloyd and B. Zimmerman for helpful comments on the manuscript. We
received valued technical assistance from A. Resetar. Field and laboratory work
were partially supported by grants from the National Science Foundation, the
Allen-Heath Memorial Foundation, and the National Geographic Society.
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CRUMP, M. L. 1971. Quantitative analysis of the eco logica l distribution of a tropical herpetofauna. Occa sional
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Natur Histoy, University of Kan sas 3~1-62.
DUELLMAN, W. E. & LYNCH, J. D. 1969. Description s
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tadpoles and their relevance to atelopo-
did class ificatio n. Herpetologica 25~231-240.
GASCON, C. 1991. Popu lations and community-level analyses of spe cies occurrences of Central Amazonian
rainforest tadpoles. Ecology 72: 1731-l 746.
HANSK I, I. 1982. Dynamics of regional distribution: the core and satellite spec ies hypothesis. Oikos 38:210-
221.
HEYER , W. R. 1973. Eco logica l interactions of frog larvae at a seaso nal tropical location in Thailand.Jo anzal
of HerpGtologr 7~337-3 61.
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HORN, H. 1966. The measurement of ‘overlap’ in comparative ecolo gical studies . American Naturalist 100:419-
424.
INGER, R. F. 1966. The systema tics and zoogeography of the Amp hibia of Borneo. Ficldiuna : Zoology 52:1-
4Q2.
INGER, R. F. 1969. Organization of comm unities of frogs along sma ll rain forest streams in Sarawak. Journal
of Anim al Ecology 38123-148.
INGER, R. F. 1972. Bufo of Eurasia. Pp. 108-l 18, 357-360 in Blair, F. W. (ed.). Evolution in the genus Bufo.
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University of Texas Press, Austin.
INGER, R. F. 1985. Tadp oles of the forested regions of Borneo. Fieldiana: Zoology (n.s.) 26:1-89.
INGER, R. F. 1992. Variation of apomorph ic characters in stream-dwelling tadpoles of the bufonid genus
Anso nia (Amphibia: Anura). Zoolog ical Journal of the Linncan Society 105:225-237.
INGER, R. F. & COLWELL, R. K. 1977. Organization of contiguo us comm unities of amp hibians and
reptiles in Thailan d. Eco logic al Morwgraphr 47~229-253.
INGER, R. F. & DRING, J. 1988. Taxonom ic and ecolo gical relations of Bornean stream toads allied to
Anso nia leptopus (Guenther) (Anura: Bufonidae). Malayan Nature Journal 419:461-471.
INGER, R. F. & GREENBERG, B. 1966. Eco logic al and competitive relations among three spec ies of frog
(genus Rana). E cology 47~746-759.
INGER, R. F. & STUEBING , R. B. 1991. Frogs of Sabah. Sabah Parks Pub lication, no. 10.
INGER, R. F. & STUEBING , R. B. 1992. The montane amph ibian fauna of northwestern Borneo. Muluyun
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INGER, R. F., VORIS, H. K. & FROGNER, K. J, 1986. Organization of a community of tadpoles in rain
forest streams in Borneo. Journal of Trop ical Ecolog y 2: 193-205.
LLOYD, M., INGER, R. F. & KING, F. W. 1968. On the diversity of reptile and amph ibian spe cies in a
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MATSUI, M. 1986. Three new spec ies of Amo lops from Borneo. Copeis 1986:623-630.
McDIARMID, R. W. 1978. Evolution of parental care in frogs. Pp. 127-147 in Burkhardt, G. M. & Bekoff,
M. (eds). The development of behavior: comparative and evolutionary aspec ts. STPM Press, New York.
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Accepted 27 April 1993
c
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Variation among frog communities
431
APPENDIX A. Site chara cteristics. Upper four in Sarawak, lower four in Sabah.
Site Labang Nanga Tekalit Pesu Segaham
Coordinates 3” 21’N/113” 27’E
Elevation (m) Cl00
Topography
flat
Vegetation primary forest
Streams:
number 1
widths (m) 10
gradient low
clarity turbid
bottom
mud
Dates:
first Ott 63/Feb 64 128 d
second
third
Locality 4
(Figure 1)
1” 37’N/113” 35’E
loo-230
hilly
primary forest
3“ 7’/113” 48’E
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432
ROBERT F. INGER AND HAROLD K. VORIS
APPENDIX B. List of spec ies and specim ens. Authorities for spec ies names given in Inger (1966, 1985) and
Inger & Stuebing (1991).
ANURAN SPECIES
Nanga
Tekalit Segaham
Ensurai Sekentut Sex-bong Selubok Wong Lawan Segaham Marok
Amolops cavitympanum
Amolops orphnocnemis
Amolops phaeomerus
Amolops poecilus
Amolops whiteheadi
Ansonia albomaculata
Anronia leptopus
Ansonia longidigita
Ansonia spinulafer .
Bufo asper
Bufo divergens
Bufo juxtasper
Chaperina furca
Leptobrachella baluensis
Leptobrachella mjobergi
Leptobrachella parva
Leptobrachella serasanae
Leptobrachium hendricksoni
Leptobrachium montanum
Leptobrachium nigrops
Leptolalax gracillis
Megophrys edwardinae
Megophrys nasuta
Micrixalus baluensis
Microphyia petrigena
Occidoqga baluensis
Pedostibes hosei
Pedostibes rugosus
Philautus disgregus
Philautus hosei
Philautus tectus
Rana baramica
Rana bbthi
Rana chalconota
Rana glandulosa
Rana hosei
Rana ibanorum
Rana ingeri
Rana kuhli
Rana laticeps
Rana malesiana
Rana paramacrodon
Rana sign&a
Rhacophorus bimaculatus
Rhacophorus gauni
Rhacophorus pardalis
Staurois latopalmatus
Staurois natator
Staurois tuberlinguis
TOTALS
0
0 0 0
0
0 0 0
90 83 330 107
13 4
28 13
0 0 0 0
0
0 0 0
32 12 21 20
0
0 0 0
0 0
0 0
107
121 71
52 0
11
2 2 0
0 0 0
0 0 0
0 1 1
0 0 0
0 0 0
0
0 0
8
20
9
0 0
0
3
0
0
5
0
0
45
0
0
0
0
1
0
0
0
5
0
3
0
4
0 0
28
3
0 0 0 0
2 0 1 1
12 143 76 3
0 0 0 0
0 0 137
2
5 2 1 0
0 0 0
0
0 0 0 3
15 14 13 1
48 1 0 0
1
0 83 8
0 0 0 0
0 0 0 0
17 0 0 96
0 0 0 4
0 0 0 0
0 0 0 0
3 0 0 20
0 0 0 0
7 1
3 0
2 1 0
2
3 2 0 0
1
14
0
2
0
1
0
0
1
1
10 0 5 0 1
50
48 106
29
1
0 0 0 0 0
0 0 0 0 0
6 0 4 5 27
0 0 0
1
0
0 1
12
2 0 0
4 0 0
0 6 1
0 2 0
0 0 0
8 0 0
0 0 5
0 0 0 0 0
0
257
307
292 200 10 30
171 156
200 125
45 135
0 0 0 0 0
0
100 56 36 20 4 0
247 141 199 191 25 22
12 82 43 44 1 18
12 14 9 17 16 55
0 0 0 0 0 0
0 0 0 0 0 0
0 0 0 0 0 0
69 150 272 23 10 11
1
3
18
0
1
11
0
1
0
0
1
1
1
2
0
1281
0
7
2
7
3
0
1221
6
3
0
1664
0
10
4
3
0
0
870
4 2 5
80 15 2
7 0 0
349 468 605
0
18
70
0
117
26
3
0
0
0
0
2
0
0
11
0
14
5
0
2
0
0
&
78
2
0
0
1
0
0
1
0
7
21
292
-
8/9/2019 A Comparison of Amphibian Communities Through Time and Place to Place
25/25
Variation among frog communities
Marak
Pew Labang Parak
Purulon Danum Mendolong
Pesu Seran Surinsin Purulon Kilampon Cabin P.Tambun SKalison W6S5 Mendolong
0 0
2 2
0 0 31
218
0 0 0
0
0 0 0 0
0 0 0
29
25:
0
0
5
0 0 0 0 3
96 161 66 108 216
0 0 0 0 0
0
0
0 0 0
0 0 0 0 161
0 0
0 0 0 0
1 0 0
0 0 1
0 0
0 0 0 0
0 0 0 0
0 0
217 27 0 1
2
0
2 0 0 0 0
1
6 0 13 53 60 3
0 0 0
3 3 0
0 0 0 0
0 0
0 00 0 0
0 0
0 0 0 0
12 4
0 0 0 0
0 0
1
0 0 0
0 0
0 2 0 6
5 0
0 2 0 0
0 0
0 0 3 18 34
14
0
0
0
0
0
0
0
0
0 0 0
5
0
6 0 4
0
0
0
0
0
0
0
0
0
60
0
0
0
1
1
241
118
315
187
5
181
10
0
T
0
0
379
18 0
0 0
0 0
0
0
0 0
14 0
38 0
57 1
155 0
0 14
0 0
61 0
0
5
0 0
1 0
9 0
7 0
5 2 3 3 8
0 0 0 0 0
2 0 0 0 0
0
15 42 12
1
0 0
0
0 0
0 0 1 0 4
0 0 0 5 0
0 0 0 0 0
0 0 0 0 0
0
20
235 83 10
1 50 48 37 33
0 0 0 0 0
1 0 0 0 0
0 0 0 0 0
0 6
1
1 0
48 30
91 17 113
0 0
0
0 0
0 0
0 0 0
0 0
0 0 0
24
14 56
82
21
5
0
19
0
11
1
1761
11
0
7
0
2
0
411
0
1
0
0
3
0
0
9
0
136
1
1
207
0
0
2
0
0
0
0
0
0
0
0
0
9
0
4
0
0
50
0
0
0
10
0
1
0
20
73
15
514
6 5
63 6
11 0
549 276
0
20
2:
0
2:
6
0
0
0
0
0
4
0
16
0
5
0
2
0
1
22
0
0
0
3
0
0
1
0
18
0
0
0 0
0
124
3
0
0
0
2 2
18 1
0 0
0 0
0 0
0 60
0 0
0 0
3
2
0 0
32 8
0 0 0
42 19 7
2 1 3
32 0
4
39 73 26
0 0
0
851 451 400
0
0
2
0
0
0
0
0
1
0
0
0
73
0
0
20
11
22
177
844