Res Popul Ecol (1999) 41:195–202 © The Society of Population Ecology and Springer-Verlag Tokyo 1999

ORIGINAL ARTICLE

Teruyoshi Nagamitsu · Kuniyasu Momose · Tamiji Inoue

David W. Roubik

Preference in flower visits and partitioning in pollen diets of stingless bees

in an Asian tropical rain forest

Received: May 14, 1997 / Accepted: April 23, 1999

Abstract Floral resource partitioning among stingless bees

Introduction
(Trigona, Meliponini, Apidae) in a lowland rain forest in
Sarawak, Malaysia, was investigated using tree towers and
walkways in a 4-year study that included a general flowering Stingless bees (Meliponini, Apidae, Hymenoptera) are
period. We obtained 100 collections of insect visitors to perennial eusocial insects and the most abundant flower
flowers of varying floral location and shape representing 81 visitors in the tropics (Roubik 1989). Many species of sting-
plant species. The tendency of 11 species of stingless bees less bees often coexist in the same habitat even though they
to visit specific flowers with a particular floral location depend on common foods, pollen and nectar. Availability
and shape was analyzed by logistic regression analysis. of either foods or nest sites limits the colony density of
This analysis showed that the proportion of flower visitor the bees (Hubbell and Johnson 1977; Inoue et al. 1993).
collections containing Trigona fuscobalteata and T. However, interference and exploitation by foragers sharing
melanocephala differed according to floral location. The feeding sites reduces the foraging efficiency of stingless
former was frequently collected at canopy and gap flowers, bees (Johnson and Hubbell 1974; Roubik 1980; Roubik
whereas the latter was most often collected at understory et al. 1986). Thus, foragers of cooccurring species may
flowers. The analysis also suggested that T. erythrogastra compete with each other for pollen and nectar. As a con-
was more rarely collected at shallow flowers than at deep sequence of the competition, floral resource partitioning is
flowers. Analysis of the pollen diets of T. collina, T. expected to occur.
fuscobalteata, T. melanocephala, and T. melina revealed Results from previous field experiments suggest that
that similarity of pollen sources differed among the six coexistence of these competing species is fostered by
permutated pairs of the four species. The lowest mean rank spatial and temporal variation in floral resource avai-
of similarity found was between T. fuscobalteata and T. lability of certain plant species (Johnson and Hubbell
melanocephala. This result supports the hypothesis that 1975; Hubbell and Johnson 1978; Johnson 1981; Nagamitsu
preference in visiting flowers in different locations leads to and Inoue 1997). Species that aggressively monopolize
pollen resource partitioning. feeding sites tend to use clumped flowers, whereas non-
aggressive species excluded from the clumped flowers
Key words Flower visitation · Foraging · Pollen preference · use scattered flowers (Johnson and Hubbell 1975;
Resource partitioning · Sarawak · Trigona Johnson 1981). Species that rapidly discover flowers also
coexist with species that discover flowers more slowly,
but aggressively monopolize flowers used by the rapid
T. Nagamitsu (*) flower-discovering species when they eventually locate
Hokkaido Research Center, Forestry and Forest Products Research them (Hubbell and Johnson 1978; Nagamitsu and Inoue
Institute, 7 Hitsujigaoka, Toyohira, Sapporo 062-8516, Japan 1997).
Tel. 181-11-851-4131 (ext. 246); Fax 181-11-851-4167
e-mail: [email protected] Such partitioning of common foods in time and space
may be a unique feature of eusocial insects, which can
K. Momose
Graduate School for Asian and African Area Studies, Kyoto
evaluate changing resources by communicating the loca-
University, Kyoto, Japan tions of optimal feeding sites (Seely 1985; Davidson 1998).
T. Inoue
However, there are also other factors that may enable sting-
Center for Ecological Research, Kyoto University, less bees to partition floral resources from plant taxa with
Otsu, Japan differing floral traits. First, tropical rain forests are vertically
D.W. Roubik structured, and thus stingless bees may be specialized in
Smithsonian Tropical Research Institute, Balboa, Republic Panama visiting flowers in different types of locations in the forest.
196

Second, flowers of tropical rain forests are morphologically

Materials and methods
diverse, and stingless bees may be specialized in feeding
on nectar and pollen of flowers with particular morpho-
Study site
logies. Previous studies on flower visits and pollen diets
of stingless bees have not been well designed to detect
The study was conducted in the Canopy Biology Plot (8 ha,
floral resource partitioning according to flower location and
200 3 400 m) and a belt transect along the Waterfall Trail
floral morphology (Heithaus 1979; Roubik et al. 1986;
(5 ha, 1 km 3 50 m) established in a lowland mixed diptero-
Inoue et al. 1990; Martinez-Hernandes et al. 1994; Wilms
carp forest in Lambir Hills National Park, Sarawak, Malay-
and Wiechers 1997). Although Roubik (1993) found that
sia (4°209 N, 113°509 E; 150–250 m in altitude). At the center
Trigona fulviventris was more frequently light-trapped in
of the Canopy Biology Plot, a canopy access system with
the understory than in the canopy, it remains uncertain
two tree towers (one 50 m and the other 55 m in height) and
whether this finding reflects specialization in foraging for
nine aerial walkways (total length, 300 m; 15–35 m above
understory flowers or is attributable to other, unknown,
ground) was constructed (Inoue et al. 1995; Fig. 1). The
factors.
Waterfall Trail was located along a stream from the head-
The purpose of this study was to examine floral resource
quarters of the park to the Operation Raleigh Tower, a
partitioning of stingless bees according to flower location
structure built in a previous research program. In the study
and morphology. This issue remains open mainly because
site, the flowering phenology of 576 individually marked
access to the canopy of tropical rain forests is difficult.
plants of 310 species was monitored from August 1992.
To resolve this problem, we built two tree towers (one
Results from the monitoring show that general flowering
50 m tall, and the other 55 m) in a lowland rain forest in
occurred from March to December in 1996, when the pro-
Sarawak, Malaysia (Inoue et al. 1995). These towers were
portion of flowering individuals among the marked plants
connected by a series of aerial walkways that extended
continuously exceeded 10% (Sakai et al., in press).
300 m through the forest. Using these towers, we have al-
In the study site, 27 species of stingless bees
ready shown that six species of stingless bees visited honey-
(Hypotrigona and Trigona, Meliponini, Apidae) were col-
water feeders at different heights (Nagamitsu and Inoue
lected (Inoue et al. 1994). In October 1994, 14 colonies of
1997). We showed that 2 of 17 coexisting species had much
seven stingless bee species were found in the Canopy Biol-
longer tongues than the other species (Nagamitsu and
ogy Plot. In August 1996, the number of colonies increased
Inoue 1998).
to 26. We found 10 colonies at trunk bases, 2 in the cavities
In the study presented here, we collected flower
of tree trunks, and 14 in wooden nest boxes (10 3 10 3
visitors and analyzed the pollen diets of stingless bees in a
20 cm) located on the tree towers. These boxes had been set
4-year research program. To examine whether preference
in place for another study on population dynamics.
in visiting flowers with different floral traits leads to pollen
resource partitioning, we analyzed our data by the following
two methods. First, using logistic regression analysis, we Flower visits
evaluated whether flower location and floral morphology
affect the tendency of each stingless bee species to visit From June 1993 to October 1996, we surveyed flowers every
specific flowers. Second, we examined whether pollen diet month from the forest floor in the Canopy Biology Plot and
similarity was lowest between pairs of species showing the along the Waterfall Trail, and every 2 months we surveyed
least similar floral preferences. flowers from the canopy access system and from the

Fig. 1. Canopy access system

and colonies of stingless bees for
which pollen diets were studied
in the Canopy Biology Plot.
Closed circles, colonies studied in
both 1994 and 1996; open circles,
colonies studied only in 1996
 197

Operation Raleigh Tower. On census days surveys usually August 4–8, and 19–23 August, in 1996 during the general
started at 0800. When we found flowering plants, we col- flowering, we collected pollen loads carried by returning
lected flower visitors for at least 10 min at individual foragers from three colonies of Trigona collina, two colo-
flowering plants. At first, we caught visitors flying around nies of T. fuscobalteata, one colony of T. melanocephala,
and coming to flowers, and later we swept completely visi- and one colony of T. melina in the Canopy Biology Plot
tors on and in flowers. When few flower visitors were col- (see Fig. 1). We also studied two additional colonies of
lected, we collected additional visitors at various times on T. fuscobalteata in 1996. We collected pollen loads of re-
following days until the plants finished flowering. Among turning foragers with hand nets at 0730, 1030, and 1430 on
the collected flower visitors, we examined only stingless single days in front of a nest entrance to each colony, which
bees in the present study. Stingless bee specimens were was closed by a cotton cloth for the preceding 20 min.
pinned and identified following the classifications of After the standard procedure of acetolysis (Erdtman
Schwarz (1939), Sakagami (1975, 1978), and Sakagami et al. 1960), 200 pollen grains from each forager were examined
(1990). We designated each group of collected individuals and classified by morphology into taxonomically distinct
of stingless bees visiting flowers of each plant species in pollen types (Huang 1972; Roubik and Moreno 1991; Tissot
each year as ‘a collection.’ et al. 1994). We regarded the most abundant types of pollen
Specimens of surveyed flowering plants were collected as legitimate pollen sources of the foragers.
and identified in SAR (Sarawak Herbarium, Forest Depart- We obtained the number of foragers, Nijk, of stingless bee
ment, Sarawak). We recorded the location, shape, and color species, i, with pollen type, j, of legitimate pollen source in
of flowers of each surveyed plants, following the methods of collection period, k. We then calculated Morishita’s similar-
Momose et al. (1998). The locations of the flowers in the ity index (Morishita 1959), Cλkl, in period, k, for pair, l,
forest structure were divided into three categories: (1) un- (l 5 1–6) of species, i and i9, as follows:
derstory (,12.5 m high in the closed forest), (2) canopy
($12.5 m high, above the understory), and (3) gap (outside
the closed forest). Flower shape was classified as either
( ) (λ 1 λ9) Â (N )Â ( N 9 )
Cλkl 5 2 Â j Nijk Ni 9jk i i j ijk j i jk

shallow (cup, rotate, or brush-shaped), or deep (including λi 5 Â {N ( N 2 1)} Â ( N ) {Â ( N ) 2 1}

j ijk ijk j ijk j ijk
papilionaceous, Caesalpinia-like, campanulate, bilabiate,
urceolate, tubular, and chamber shapes). Floral color was λ i9 5 Â {N 9 ( N 9 2 1)} Â ( N 9 ) {Â ( N 9 ) 2 1}
j i jk i jk j i jk j i jk
categorized as (1) white and green, (2) yellow, or (3) red-
dish (including orange, brown, red, pink, and purple). We tested a null hypothesis that there were no differences
Logistic regression analysis was performed to examine in the similarity indices among the species pairs, H0: Cλ1 5
effects of floral traits on the tendency of each stingless bee Cλ2 5 Cλ3 5 Cλ4 5 Cλ5 5 Cλ6, by Friedmann’s test.
species to visit specific flowers (SAS Institute 1994). Effects
of flower location, L, and shape, S, on the proportion of
collections, Pi, containing at least one individual of stingless
bee species, i, was tested. We added effects of flower color, C, Results
and collection year, Y, to models of the analysis as block
factors. Collection years were grouped into (1) 1993–1995, Flower visits
before the general flowering and (2) 1996, when the general
flowering occurred. A linear model of the analysis is given by From June 1993 to October 1996, we obtained a total of 100
collections of stingless bees visiting flowers (of 81 plant
logit(Pi )klmn 5 µi 1 Yk 1 Ll 1 Sm 1 Cn species representing 37 plant families; see Appendix),
which were classified according to floral location, shape, and
(k 5 1, 2; l 5 1,2,3; m 5 1,2; n 5 1, 2,3) color (Table 1). These collections consisted of 3279 stingless
where µi is mean logit-transformed proportion of collec- bees of 21 species (Table 2). Trigona itama was the most
tions that contained stingless bee species, i. We tested abundant species (comprising 1016 individuals), although T.
the null hypotheses that flower location and shape had no fuscobalteata was most frequently collected, appearing in 34
effect: of the 100 collections. We examined 11 stingless bee species
that appeared in more than 10 collections in the following
H0 : L1 5 L2 5 L3 5 0, and analysis.
H0 : S1 5 S2 5 0 The linear model of logistic regression analysis was not
by Wald’s ø2 test. The probabilities of type I error were rejected for any species, suggesting that there were negli-
adjusted by the sequential Bonferoni method for multiple gible interactions among year, location, shape, and color
tests for i species (Sokal and Rohlf 1981). (Table 3; Wald’s ø2 test, P . 0.930, adjusted by the sequen-
tial Bonferoni method for results of tests for the 11 species).
The effects of flower location on the proportion of collec-
Pollen diets tions containing T. fuscobalteata and T. melanocephala
were significant (P 5 0.045 and 0.002, respectively, adjusted
On June 22–23, August 10–13, and September 17–19, as above). T. fuscobalteata was frequently collected at
in 1994 before the general flowering, and on May 15–18, canopy and gap flowers, whereas T. melanocephala often
198

Table 1. Sample numbers in collections of stingless bees visiting flowers representing three types of location, two types of shape, and three types
of color
Shape Color 1993–1995 1996 Subtotal Total

Canopy Gap Understory Canopy Gap Understory

Deep White and green 1 0 4 4 1 2 12

Yellow 0 2 1 1 0 0 4
Reddish 2 9 5 2 2 1 21 37
Shallow White and green 3 9 5 6 1 3 27
Yellow 3 7 3 8 2 1 24
Reddish 0 6 1 2 1 2 12 63
Subtotal 9 33 19 23 7 9
Total 61 39 100

Table 2. Stingless bee species found in 100 collections of flower visi-

tors, showing the numbers of collected individuals and the number of
collections including each stingless bee species
Stingless bees Code No. of bees No. of collections

Species

Trigona fuscobalteata fus 380 34

T. itama ita 1016 32
T. laeviceps lae 74 26
T. melanocephala mla 58 25
T. apicalis api 151 22
T. erythrogastra ery 113 18
T. ventralis ven 278 17
T. fimbriata fim 310 16
T. canifrons can 405 15
T. collina col 186 14
T. melina mli 30 13
T. nitidiventris 45 9
T. rufibasalis 29 7
T. thoracica 28 6
T. reepeni 11 6
T. terminata 46 5
T. haematoptera 46 4
Hypotrigona scintillans 8 4
T. melanoleuca 34 2
T. moorei 29 2
T. geissleri 2 2
Fig. 2. Proportion (bars) of collections containing Trigona fusco-
Total 3279
balteata and T. melanocephala individuals visiting canopy, gap, and
understory flowers. The number of collections of stingless bees visiting
flowers at each type of location is shown
appeared in collections from understory flowers (Fig. 2). An
effect of flower shape on the proportion of collections that
fuscobalteata, 132 collected pollen from euphorbiaceous
contained T. erythrogastra was marginally significant (P 5
species. Pollen from species of Araceae, Annonaceae, and
0.079, adjusted as above). T. erythrogastra was more often
Leguminosae was collected by 49 of 135 T. melina foragers,
collected at deep flowers than at shallow flowers (Fig. 3).
41 of 121 T. melanocephala foragers, and 37 of 226 T. collina
foragers, respectively.
Pollen diets The analysis of flower visits (see earlier) suggested that
among the four species of which pollen diets were exam-
We collected pollen loads from 104, 105, and 65 foragers in ined, T. fuscobalteata and T. melanocephala preferred flow-
three periods in 1994, and from 124, 148, and 147 foragers in ers in the most divergent locations. Thus, if differences in
three periods in 1996. In pollen loads from the 693 foragers flower location preference causes partitioning between
sampled, 102 types of pollen grains were morphologically plant taxa acting as pollen sources, the pollen diet should be
distinguishable. Among the 102 types of pollen, 55 were less similar between T. fuscobalteata and T. melanocephala
identified to the plant family level. The most abundant fam- than those between any of the other five permutated pairs
ily was Euphorbiaceae, observed in pollen loads from 198 from the four species.
foragers, followed by Araceae (52), Annonaceae (51), and In accordance with this hypothesis, pollen diet similarity
Leguminosae (38). The main pollen sources differed among significantly differed among the six permutated pairs of the
the stingless bee species studied. Among 211 foragers of T. four stingless bee species (Friedmann’s test, P 5 0.048), and
 199

Table 3. Results of logistic regression analysis testing effects of collection year, flower location, flower shape, and flower color on the proportion
of collections including each stingless bee species
Stingless bee species

fus ita lae mla api ery ven fim can col mli

Year 0.737 0.869 0.973 0.361 0.832 0.600 0.849 1.000 0.982 0.991 0.529
Location 0.045 0.152 0.694 0.002 0.833 0.620 0.732 1.000 — 0.641 0.918
Shape 0.978 1.000 0.894 0.755 0.272 0.079 0.974 0.999 1.000 1.000 1.000
Color 0.999 0.632 0.994 1.000 0.631 1.000 — 0.858 1.000 0.999 1.000
Model 0.984 0.930 1.000 1.000 0.998 0.944 1.000 0.986 0.986 1.000 0.993
—, No test was done because no corresponding samples were collected
Species codes are shown in Table 2. The probabilities of type I errors occurring, adjusted by the sequential Bonferoni method for the four factors
and linear models of the analysis, are also shown

Fig. 4. Pollen diet similarity measured by Morishita’s similarity index,

Cλ, between pairs of four stingless bee species. Species codes are as
Fig. 3. Proportion (bars) of collections containing Trigona erythro- shown in Table 2. Similarities were found in three periods in 1994 and
gastra individuals visiting shallow and deep flowers. The number of in three periods in 1996
collections of stingless bees visiting flowers of each type of shape is
shown

the mean rank of the T. fuscobalteata and T. melanocephala and Hubbell 1974; Roubik 1980; Roubik et al. 1986). Avail-
pairing was lowest in the pollen diet similarity (Fig. 4). able evidence so far accumulated suggests that stingless bee
However, the range of similarity strongly shifted, from species are able to partition floral resources of the same
0.00–0.10 during June 22–23, 1994 to 0.51–0.85 during 19– plant species through temporal and spatial variation in the
23, August 1996. In all test periods, except for 19–23, floral resources (Johnson and Hubbell 1975; Hubbell and
August 1996, when the pollen diet similarity of every pair Johnson 1978; Johnson 1981; Nagamitsu and Inoue 1997).
was extremely high, the similarity between T. fuscobalteata Pollen and nectar sources of diverse plant species are thus
and T. melanocephala was less than 0.02, indicating almost shared among stingless bee species (Heithaus 1979; Roubik
complete partitioning of pollen sources. Thus, the foregoing et al. 1986; Inoue et al. 1990; Martinez-Hernandes et al.
prediction was largely supported. 1994; Wilms and Wiechers 1997).
Stingless bees have also been regarded as generalists in
floral resource use that rarely specialize in particular plant
taxa with unique floral traits. However, several lines of
Discussion evidence suggest that floral resource partitioning according
to flower location and morphology does occur (Roubik
Floral resource partitioning has been proposed to occur 1993; Nagamitsu and Inoue 1997, 1998). The results pre-
among stingless bee species, because foragers of these spe- sented here show that two stingless bee species visited flow-
cies are likely to compete for pollen and nectar (Johnson ers in different types of locations within the forest structure,
200

and suggest that these two species did partition pollen ac- the results obtained in this study show that pollen diet simi-
cording to taxonomic source. Further, one species visited larity varied considerably, both seasonally and annually.
morphologically specialized flowers with enclosed petals, There was a period when pollen diet similarity was ex-
although this result was only marginally significant. These tremely high between all pairs of studied stingless bee spe-
results suggest that floral resource partitioning of stingless cies. This result suggests that pollen resource partitioning
bees arises not only from variation in floral resources within depends on pollen resource availability, which seemed to
plant species, but also from variation in floral traits among fluctuate during this study because of the general flowering
plant species. that occurred in 1996. However, we did not evaluate tempo-
The observed preference of Trigona melanocephala for ral changes in floral resource abundance and foraging effi-
visiting understory flowers is consistent with our previous ciency in the present study. These issues remain to be
observation that this species tended to visit the lowest addressed in further studies.
among honey-water feeders placed at heights ranging from
1 to 50 m above ground level (Nagamitsu and Inoue 1997). Acknowledgments We thank Dr. H. S. Lee, Mr. A. A. Hamid (Forest
However, height above the ground alone cannot explain the Department, Sarawak), and Dr. K. Ogino (University of Shiga Prefec-
ture) for kindly supporting the present study; Mrs. R. Johan, B.
different preferences for gap and understory flowers at simi- Nyambong, R. Rapi (Forest Department, Sarawak), and all members
lar heights. Factors common to both the canopy and gap, of the Canopy Biology Program in Sarawak for helping collect flower
but differing in the understory, may affect the observed visitors; and anonymous reviewers for improving this manuscript. This
preference in visiting flowers in different locations. study was partly supported by grants-in-aid from the Japanese Ministry
of Education, Science and Culture (grant numbers 04041067, 06041013,
Possible factors involved include environmental condi- and 09N1501).
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Appendix. Plant species of flowers visited by stingless bees

Plant Floral trait Collection code No. of collection

Family Species L C S 1993 1994 1995 1996

Anacardiaceae Buchanania sessilifolia u w d 62 1

Annonaceae Enicosanthum coriaceum u w d 63 1
Enicosanthum macranthum u w d 1 34 2
Polyalthia cauliflora u y d 2 1
Uvaria elmeri u w s 35 1
Ascrepiadaceae Gongonema g y s 3 1
Burseraceae Dacriodes laxa c r s 64 1
Santiria laevigata c w s 4 65 2
Celastraceae Lophopetarum glabrum c r s 66 1
Compositae Eupatrium odoratum g w s 36 1
Mikania micrantha g w s 5 1
Vernonia arborea g w s 67 1
Convolvulaceae Ipomoea pes-carpae g r d 6 1
Cyperaceae Carex sp. g w s 7 37 2
Dilleniaceae Dillenia exelsa g y s 68 1
Dillenia suffruticosa g y s 8 1
Dipterocarpaceae Dryobalanops aromatica c w d 69 1
Dryobalanops lanceolata c w s 9 1
Shorea beccariana c y s 70 1
Elaeocarpaceae Elaeocarpus stipula g w s 10 1
Euphorbiaceae Cleistanthus pseudopodocarpus c y s 11 1
Cleistanthus sumatranus c y s 73 1
Dimorphocalyx denticulatum u r s 74 1
Drypetes longifolia u w s 14 38 51 3
Drypetes xanthophyloides u w s 75 1
Homalanthus populneus g y s 15 39 2
Koilodepas laevigatum c y s 16 76 2
Macaranga winkleri g w s 52 1
Mallotus grifithii u y s 77 1
Mallotus penangensis c y s 12 71 2
Mallotus wrayi u y s 13 1
Tapoides sp. c w s 72 1
Fagaceae Lithocarpus lucidus g y s 78 1
Gesneriaceae Didisandra sp. u w d 17 40 2
Guttiferae Mesua oblongifolia u w s 79 1
202

Appendix. Continued
Plant Floral trait Collection code No. of collection

Family Species L C S 1993 1994 1995 1996

Hypericaceae Cratoxylum sumatranum g r s 41 53 2

Lauraceae Litsea sp. u w s 80 1
Leguminosae Acacia mangium g y s 54 1
Callerya nieuwenhuisii g r d 18 81 2
Cassia sp. g y d 19 1
Fordia splendissima u r d 20 82 2
Mimosa pudica g r s 42 1
Peltophorum pterocarpum g y d 43 1
Sindora beccariana c r d 21 55 2
Sindora irpicina c r d 83 1
Spathorobus ferrugineus g r d 84 1
Melastomataceae Melastoma beccarianum g r d 22 44 2
Melastoma malabatricum g r d 23 45 56 3
Memecyron sp. g r s 85 1
Pternandra multiflora g y s 24 57 2
Meliaceae Walsura sp. c w s 86 1
Musaceae Musa campestris g r d 46 1
Myrtaceae Eugenia sp. c w s 47 1
Eugenia subrufa c w s 87 1
Olacaceae Anacolosa frutescens u r s 25 1
Scrodocarpus borneensis c w d 58 1
Palmae Calyota sp. u y s 26 1
Ponteridaceae Monochoria sp. g r s 59 1
Rhamnaceae Ventilago malaccensis c y s 88 1
Rubiaceae Ixora woodii u r d 27 48 60 3
Uncaria longifolia g w d 89 1
Rutaceae Glycosmis sp. u y s 28 1
Sapindaceae Nephelium cuspidatum c w s 90 1
Pometia pinnata c w s 91 1
Simaroubaceae Quassia borneensis c y s 92 1
Sterculiaceae Heritiera borneensis c w d 93 1
Pterocymbium tubulatum c w d 94 1
Scaphium borneensis c w d 95 1
Scaphium longipetiolatum c r d 96 1
Sterculia stipulata u r d 29 1
Theaceae Eurya acuminata g w s 49 1
Tiliaceae Grewia latistipula c y s 97 1
Grewia stylocarpa c y s 98 1
Ulmaceae Trema tomentosa g w s 30 1
Urticaceae Dendrocnide stimulans u w s 31 1
Pipturus argentens g w s 32 1
Verbenaceae Calicarpa havirandii g r s 50 1
Calicarpa pendula g r s 61 1
Clerodendron megaphyla u r s 99 1
Stachytapheta indica g r d 33 1
Xanthophyllaceae Xanthophyllum velutinum c y d 100 1
Total 33 17 11 39 100
L, location; c, canopy; g, gap; u, understory; C, color; w, green and white; y, yellow; r, reddish; S, shape; s, shallow; d, deep
Floral traits (L, location; C, color; S, shape) are shown for each plant species. Collections of stingless bees visiting flowers of each plant species
are shown for each study year