The Xanthomonas oryzae pv. oryzae raxP and - Pamela Ronald

BlOTROPlCA 36(4): 522-533 2004

Temporal Variation in the Relative Abundance of Fruit Bats (Megachiroptera: Pteropodidae) in Relation to the Availability of Food in a Lowland Malaysian Rain Forest’

Robert Hodgkison2, Sharon T. Balding

School of Biological Sciences, Zoology Building, University of Aberdeen, Tillydrone Avenue, Aberdeen AB24 2TZ, Scotland

Akbar Zubaid

Department of Zoology, Universiti Kebangsaan Malaysia, 43600 UKM Bangi, Selangor, Malaysia

and

Thomas H. Kunz

Center for Ecology and Conservation Biology, Department of Biology, Boston University, Boston, Massachusetts 02215, U.S.A.

ABSTRACT The aims of this study were to investigate the diet and relative abundance of fruit bats in a lowland Malaysian rain forest and to test the hypothesis that the local assemblage structure of fruit bats varies significantly over time in relation to the availability of food. In total, 352 fruit bats of eight species were captured during 72,306 m2 mist-net hours of sampling between February 1996 and September 1777. Three species of fruit bats (Balionycteris macukzta, Chironaw rnekznocephalw, and Cynopterw brachyotis) that fed on a wide range of “steady state” and “big b a n g food resources were captured continuously throughout the study period, with no significant variation in capture rates over time. In contrast, five species that fed exclusively or almost exclusively on “big b a n g food resources were sampled intermittently, with significant temporal variation in the capture rates of two species (Cynopterw horsjeLdi and Megaerops ecaudatus). Significant variation in the capture rates of the remaining three species (Dyacopterw spadiceus, Eonycteris spekzea, and R o w e m amphicaudztw) could not be detected due to small sample sizes. Since ephemeral “big b a n g food resources were only sporadically available within the study area and were associated with large canopy trees and strangler figs, these results suggest that food abundance, or the availability of specific food items, may be important factors limiting local fruit bat species diversity in old-growth Paleotropical rain forest. Thus, only three fruit bat species were locally resident within the forest throughout the study period. Therefore, further studies on the ranging behavior and habitat requirements of Malaysian fruit bats are required to assess the adequacy of existing reserves and protected areas.

Ki word: Southeast Asia.

Chiroptera; dipterocarp forest; foraging ecology; migration; nomadism; phenology; Pteropodidae; refigins

IT HAS LONG BEEN RECOGNIZED THAT OLD WORLD FRUIT BATS (Chiroptera: Pteropodidae), like many other frugivores, often migrate or move nomadi- cally in response to temporal variation in the availability of food (Nelson 1965; Thomas 1982, 1983). Thus, local population abundance of some species can fluctuate considerably over time, particularly in highly seasonal habitats (e.g., subtropi- cal Australia and parts of Africa). In contrast, the foraging movements of fruit bats in less seasonal parts of the world are less clearly understood. Al- though several species forage over large areas of habitat (Start 1974; Marshall 1983, 1985), no

’ Received 21 October 2003; revision accepted 24 April 2004.

Corresponding author.

study has quantified variation in local fruit bat population abundance and assemblage structure over time, particularly in relation to the availability of food. Thus, the influence of food abundance on the maintenance of fruit bat species diversity is poorly understood.

The aim of this study was to investigate the diet and population abundance of fruit bats in a lowland Malaysian rain forest and to test the hypothesis that the local assemblage structure of fruit bats varies significantly over time in relation to the availability of food. Based on studies on birds and diurnal mammals in Southeast Asia (Fogden 1972, Leighton & Leighton 1983, Curran & Leighton 2000) and studies on bats in other regions of the world (Fleming et al. 1972, Bonaccorso 1979), we predicted that fruit bat species that exploit aseaso-

522

Fruit Bat Abundance 523

nal “steady state” food resources would be captured continuously throughout the year, whereas species that exploit ephemeral “big bang” food resources would be captured intermittently; however, due to the supra-annual fruiting phenologies of many Southeast Asian trees (Leighton & Leighton 1983, Appanah 1985), we suggest that the foraging movements of some species of bat may vary between years. Therefore, we investigated the diet and relative abundance of fruit bats during 33 months of sampling.

METHODS STUDY sm.-This study was conducted at Kuala Lompat (3”43‘N, 102”17’E), within the Krau Wildlife Reserve, Pahang, Peninsular Malaysia during two 12-month field seasons between May 1997 and November 1999. Additional data were also collected during a pilot project between February and October 1996.

The Krau Wildlife Reserve consists of a large area of old-growth forest (Clark 1996), which rises from 50 m at Kuala Lompat to over 2000 m at the summit of Gunung Benom. Because it lies in the rain shadow of Gunung Benom, Kuala Lompat is comparatively dry for the region, averaging ca 1982 mm of rain annually (Raemaekers et al. 1980). Although the rainfall pattern is variable, there is a slight trend toward a relatively dry season at the beginning of the year, particularly in Feb- ruary, followed by a wet season from October to December (Raemaekers et al. 1980, Hodgkison 2001). Temperature is stable throughout the year, with maximum and minimum daily air tempera- tures ranging from 30 to 35 “C and 20 to 25 “C, respectively (Raemaekers et al. 1980, Hodgkison 2001).

The vegetation at Kuala Lompat can be classified as lowland evergreen mixed dipterocarp forest; however, because it is relatively poor in dip- terocarps and unusually rich in large Leguminosae, the site is not characteristic of most dipterocarp forest sites within the region (Raemaekers et al. 1980).

The fruit bat fauna at Kuala Lompat includes at least 1 1 species (Medway & Wells 1971; Francis 1990, 1994; Zubaid 1993), including: Balionycteris maculata (Thomas), Chironax melanocephalus (Temminck), Cynopterus brachyotis (Muller), Cynopterus hor$eMi (Gray), Dyacopterus spadiceus (Thomas), Eonycteris spelaea (Dobson), Macroglos- sw sobrinw (Andersen), Megaerops ecaudatus (Tem- minck), Megaerops wetmorei (Taylor), Penthetor lu-

casi (Dobson), and Rousettus amplexicaudatus (Geoffroy). At least two of these species (E. spelaea and R. amplexicaudatus) are known to forage over large areas of habitat from central refuges and can travel at least 38 km to feed within a single night (Start 1974; Start & Marshall 1976; Marshall 1983, 1985).

FRUIT BAT ASSEMBLAGE.-FrUit bats were sampled throughout the 33-month study period using two- ply, 50 denier, 35 mm mesh mist nets. To reduce sampling bias in relation to net height, these nets were stacked together in rigs to create continuous walls of net, extending from ground level to the lower forest canopy (Hodgkison et al. 2002). Two pulley systems (one at either end of the net) were used to hoist the nets into position, and the entire rig was held in place by a main support rope positioned across two sturdy branches in the canopy.

A total of 24 rope-and-pulley rigs were estab- lished at sites located throughout the 1.5 km2 study area (Hodgkison et al. 2004). All rigs were positioned away from forest trails and known food plants, and were high enough to deploy between five and nine 6 x 3 m mist nets. To reduce the risk of bats learning the positions of the mist nets, the 24 sites were netted on rotation. Only very rarely was any single net operated on two consecutive nights. Some nets, however, were eventually run more frequently than others.

Netting protocol was influenced by both weather, and lunar cycles. In the interest of safety, nets were never opened during or immediately after high winds and heavy rains. Several rope pulley systems were partially or completely destroyed by falling trees and branches, particularly after wet weather. Netting was also avoided during brightly moonlit nights because it appeared, but was not demonstrated, that bats may have avoided the nets more easily on such occasions.

Nets were opened after dusk (1930-2000 h) and inspected every 15-30 minutes for bats until they were closed, between midnight and 0300 h. Captured fruit bats were identified to species, following Payne et al. (1985) and Medway (1983), and records were made of forearm length, body mass, sex, age, and reproductive status. All individuals were marked with stainless steel ball-chain necklaces (Ball Chain Manufacturing Company, Mount Vernon, New York) for the larger species (Cy brachyotis, Cy hor$eMi, M. ecaudatus, D. spadiceus, and R. amplexicaudatus) or monel wing bands (Lambournes Ltd., Birmingham, England) for smaller species and the nectarivorous Macro-

524 Hodgkison, Balding, Zubaid, and Kunz

glossinae (Ch. mekznocephalus and E. spekzea; Kunz 1996). Recaptured B. mucukzta were recognized by their distinctive wing markings (Hodgkison 2001). Individuals of larger species that were too small ( ie . , immature) to be fitted with a necklace were marked temporarily by wing punching. Feces and fruit parts collected during mist netting were placed in paper envelopes and used to identify food plants (see Fruit Bat Diets below).

hALysls.-Fruit bat abundance was estimated at quarterly (3 mo) intervals and was measured as the number of individual fruit bats captured per 100 m2 mist net hours (mnh) of sampling (Zubaid 1994). This figure was calculated separately for each of the mist net rigs that were operated inde- pendently during 1 1 quarterly sampling periods. Temporal variation in the abundance of fruit bats was then analyzed for all species combined, and then each species individually, using a Kruskal- Wallis test ( S o u & Rohlf 1995).

FRUIT BAT DIErs.-Three main methods were used to investigate fruit bat diets within the study area: (1) regular collection of fruit bat feces and rejected fruit parts at day roosts and feeding perches (B. macuhta only); (2) mist netting at fruiting/flowering plants; and (3) opportunistic collection of food items or food remains from netted bats (Thomas 1988).

REGULAR COLLECTION OF FECES AND REJECTED FRUIT

rmTs.-The feces and rejected fruit parts of B. macuhta were sampled throughout the study period using seed collection trays. These trays, which were positioned directly below feeding perches and diurnal roost sites, consisted of fine-weave black ny- lon fabric, spread loosely over a 1 x 1 m square wooden frame. A wooden stand, ca 0.5 m tall, provided a level base to support each tray and helped reduce seed removal by ground-foraging animals. All trays were collected from the forest once a month and inspected for the presence of seeds and pollen. All feeding perches and roost sites used in this study were located during a concurrent study on the roosting ecology and social organization of B. macuhta within the same study area (Hodgkison eta/. 2003).

MIST NETTING AT FRUITING~FLOWERING PLANTS.-

The forest was continuously monitored throughout the study period for signs of flowering and fruiting. Particular attention was paid to flowers and fruits that displayed any of the characteristic features as-

sociated with the attraction of bats (van der Pijl 1957). Food sources were also located through di- rect observations of fruit trees at night and by searching the area surrounding fruiting and flowering plants for signs of fruit bat feeding activity, such as fruit bat feces, tooth and claw-marked flowers and fruits, and piles of rejected seeds and fruit parts. When a plant was judged to offer a potential source of food, the presence and identity of foraging bats were investigated using mist nets. De- pending on the size of the individual fruiting/flowering plant, these nets were either deployed hori- zontally, on poles, or hoisted into the subcanopy/ canopy on the end of a rope (Munn 1991). Captured fruit bats were then placed in cloth bags and provided with fruit samples collected from the tree species under investigation. If these fruit samples were eaten, we determined that these food items were included in the diet of the fruit bat species concerned.

OPPORTUNISTIC COLLECTIONS OF SEEDS AND FRUITS.-

Dietary information was occasionally collected by chance, either when a bat carried a fruit or other food remained directly into a mist net or when a captured individual excreted seeds or pollen in its feces during handling. To increase the occurrence of the latter, captured individuals were held in cloth bags for at least 30 minutes prior to their release.

All unknown seeds collected from fruit bats were germinated for identification. To avoid con- fusion with seeds within the soil, small seeds were germinated in petri dishes lined with moistened fil- ter paper. The seedlings were then transferred to pots and grown in soil when their cotyledons had developed. Large seeds were planted directly into pots. All plants were grown under shading, watered liberally, and kept free of weeds and pests. Pollen samples from fur and feces were dried and stored in paper envelopes or were mounted and stored as slides using a gelatin-fuchsin stain (Beattie 1971). Some samples were also stored in 70 percent ethanol.

IDENTIFICATION OF FOOD rmTs.-Seeds, fruits, and other plant remains were identified by two methods: ( 1 ) comparison with reference collections of fruits, seeds, seedlings, and pollen; and (2) germination.

REFERENCE coLLEcTIoNs.-Wet and dry collections were made of all fruit, seed, and pollen specimens collected from within the forest. Wet samples were stored in 70 percent ethanol. Dry specimens were


a) Bafionycteris maculata (N = 105)

FIGURE 1. Temporal variation in the mean capture rates (CR) of five s ecies of fruit bat (a-e), in an old-growth lowland rain forest at Kuala Lompat, Malaysia. Ca ture rates are expresselas bats per 100 m2 mist net hour (mnh) and were measured at quarterly (3 mo) intervals Lorn February 1996 to October 1999. The number of mist net sample sites operated durin each three-month interval is indicated in parenthesis on the x-axis, and eriods without sampling are indicated by skading. Bars indicate standard deviations and Nindicates the sample size i!r each species. Statistics refer to Kruskal-Wallis tests.

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stored in paper envelopes. Some seeds were also germinated and grown to the seedling stage. To- gether these specimens provided a reference collection by which all fruits and seeds and some pollen collections recovered from bats could be compared and matched for identification. Seeds and seedlings were examined with the aid of a hand lens and binocular microscope. Pollen specimens were mounted in a gelatin-fuchsin stain (Beattie 1971) and examined microscopically using a compound light microscope. Preliminary identifications based on seed morphology alone were confirmed by seed germination or by netting around the plants from which the reference samples had been collected. The former method was especially important for verifying the identity of small seeds from large gen- era (particularly Ficus), which could easily be con- fused at the species level.

GERMINATIoN.-Seeds and seedlings that could not be matched with the reference collection were grown in pots until they were large enough to be keyed out and identified in the field. All specimens were identified following Whitmore (1972, 1973) and Ng (1978, 1989). Once a full description of the plant had been obtained, it was then possible to locate mature specimens in the forest, to mon- itor their phenology and collect leaves, flowers, and fruits. Many of these plants then provided new tar- get species for the sampling of bats.

PHENoLoGY.-The production of fruits and flowers was monitored at monthly intervals for all fruit bat food plant species identified within the study area, measuring the frequency and timing of crop production and crop size and the degree of synchrony in the maturation of fruits and/or flowers on an individual plant (Gentry 1974, Newstrom et al. 1994). Crop size was recorded as the percent area in which fruiting and flowering activity was estimated to occur within the crown of an individual plant (Wheelwright 1986); degree of synchrony was estimated as the time interval that separated the ripening of the first and last fruits or the open- ing of the first and last flowers. Hence, in this classification, the degree of synchrony was inversely proportional to the duration of fruiting and flowering activity on an individual plant. Species with large crops (>50% crown cover) that matured syn- chronously (< 1 mo) at annual or supra-annual intervals were classified as “big bang,” whereas species with small crops (<50% crown cover) that matured gradually (>1 mo) at sub-annual intervals were classified as “steady state” (Gentry 1974). All


other species were classified as “intermediate”. These species were generally characterized by small crops (<50% crown cover) that matured gradually (> 1 mo) at annual or supra-annual intervals; however, because many food plant species remained unidentified until the final year of study, interannual variation in crop production was not systematically recorded.

VERIFiCATioN.-Herbarium specimens of all fruit bat food plant species identified in the forest were submitted to two of Malaysia’s national herbaria for verification, Universiti Kebangsaan Malaysia (UKM) and the Forest Research Institute of Ma- laysia (FRIM).

RESULTS FRUIT BAT muND.mcE.-Three hundred and fifty- two fruit bats of eight species were captured in the stacked mist net rigs during 72,306 m2 mist net hours (mnh) of sampling, which included a sampling effort of 12,852 m2 mnh in year 1 (February 199GOctober 1997), 40,691 m2 mnh in year 2 (May 1997-April 1998), and 18,763 m2 mnh in year 3 (November 1998-October 1999). The mean quarterly capture rate in stacked mist net rigs ranged from 0.26 to 0.86 fruit bats per 100 m2 mnh, with no significant variation over time (H = 13.52, P > 0.05).

Three species of fruit bats (B. maculata, Ch. melanocephalus, and Cy brachyotis) were captured continuously throughout the study period, no significant variation in capture rate over time (Fig. 1). In contrast, five species (Cy horsjeka’i, D. spadiceus, E. spelaea, M. ecaudutw, and R. amplexicaudatw) were captured intermittently, with significant temporal variation in the capture rates of two species (Cy. horsjeka’i and M. ecaudutw). Significant temporal variation in the capture rates of the remaining three species could not be tested due to small sample sizes (<5 captures).

FRUIT BAT DlET.-Twenty-nine plant species from 14 families were identified within the combined diets of the 8 fruit bat species captured within the study area (Appendix). Twenty-five of these plant species were exploited by fruit bats for fruits, 3 for flowers, and 1 for fruits and flowers. The fruiting and flowering phenologies of these food plant species are shown in Table 2. Seven more food plant species (unidentified in the forest) were also recorded by their seeds, pollen, and leaves (Table 1).

The fruit bat fauna at Kuala Lompat could be

divided into two groups in relation to the fruiting/ flowering phenologies of the plant species included in their diets (Fig. 2). Five species of fruit bats (Cy horsjeldi, D. spadiceus, E. spelaea, M. ecaudatus, and R. amplexicaudatus) that consistently formed large feeding aggregations almost exclusively exploited “big bang” crops associated with large canopy trees and strangler figs (Ficus spp.; Appendix). In contrast, the remaining three species (B. maculata, ch. melanocephalus, and Cy brachyotis), were the only fruit bat species that also exploited the truly asea- sonal “steady state” crops associated with certain species of subcanopy tree and one species of scram- bling fig (Ficus globosa; Appendix). These plants produced small quantities of fruits that were available to fruit bats throughout the year (Fig. 2).

DISCUSSION The fruit bat assemblage at Kuala Lompat consisted of eight species, which collectively fed on the fruits and flowers of at least 35 species of plant. Three species of fruit bats (B. maculata, Ch. melanocephalus, and Cy brachyotis) that fed on a wide range of “steady state” and “big bang food resources were captured continuously throughout the study period, with no significant variation in capture rate over time. In contrast, five species that fed exclusively or almost exclusively on relatively ephemeral “big bang” food resources were sampled intermittently, with significant temporal variation in the capture rates of two species (Cy horsjekfi and M. ecaudatw). Significant temporal variation in the capture rates of the remaining three species (D. spadiceus, E. spelaea, and R. amplexicaudatw) could not be detected due to small sample sizes; however, since “big bang food resources were available only sporadically within the study area and were associated with large canopy trees and strangler figs, these results suggest that food abundance or the availability of specific food items may be important factors limiting local fruit bat species diversity within old-growth Paleotropical rain forest. Thus, only three fruit bat species were locally resident within the forest throughout the study period (B. maculata, Ch. melanocephalus, and Cy brachyotis) .

These results therefore have important impli- cations for the conservation of fruit bat diversity in Malaysia, since many species of Malaysian fruit bats require relatively large areas of habitat in order to locate continuous supplies of food. For example, E. spelaea, which forages within a wide range of forest and non-forest habitats, can travel up to 38 km to


b) Chironax melanocephalus 10 1

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______-_

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0 1 J F M A M J J A S O N D

Month 0 Steady state rn Intermediate rn Big bang rn Unknown

FIGURE 2. Tempord variation in the availability of fruits and flowers consumed by eight species of fruit bats (a- h) in an old-growth lowland rain forest at Kuala Lompat, Malaysia. This chart shows the number of known “steady state”, “intermediate”, and “big bang phenology food items consumed by each species of fruit bat throughout the year, based on more than one year of study. See methods for definitions of phenology pattern.


feed in a single night (Start 1974, Start & Marshall 1976). T h e foraging ranges and habitat requirements of many other species are largely unknown; however, at least two species that were nonresident within the study area at Kuala Lompat (D. spadi- cells and M. ecaudatus), appeared to be strongly associated with old-growth lowland and montane rain forest (Lim 1966, Medway 1983, Payne eta/. 1985, Zubaid 1993, Francis 1994). Thus, these species are likely to be particularly vulnerable to local extinction due to habitat destruction and forest degradation. Further studies on the ranging behavior and habitat requirements of these bats are clearly required in order to assess the adequacy of existing reserves and protected areas.

ACKNOWLEDGMENTS We wish to thank the Department of Wildlife and Na- tional Parks (Malaysia) for their generous support throughout this project and providing us with the use of the field station facilities at Kuala Lompat. This study would not have been possible without the permission of the Economic Planning Unit of Malaysia. Valuable assis- tance and encouragement in the field was given by A. Bin Dagu, M. Salleh Kamarudin, T. Kingston, M. A. Bin Ngah Sanah, T. Ong, K. Thomsen, and Z. Bin Zainal. Constructive comments on the manuscript were offered by I? A. Racey and two anonymous referees. This study was supported by a grant from the Lubee Bat Conser- vancy, Florida, to T. H. Kunz. Additional sponsorship to R. Hodgkison was provided by the Carnegie Trust for the Universities of Scotland, the Royal Society South East Asia Rain Forest Regeneration and Recovery Programme (RS-112), and Bat Conservation International.

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APPENDIX. Food resources consumed by fruit bats in a lowland Malaysian rainforest, KuaIa Lompat. Plant nomenclature follows Whinnore (1972, 1973) and Ng (1978, 1989), unless otherwise stated.

Growth Food Fruit bat Plant species forma VPe speciesb

Annonaceae Cyathocalyx scortechinii (King) Sinclair Polyalthia obliqua Hk. f. et Thorns. Polyalthia sp. Pseuduvaria setosa var. major (King) Sinclair

Diospyros sumatrana Miq.

Elaeocarpus stipulark var. stipularis B1.

Parkia javanica (Larnk.) Merr. I! speciosa Hassk.

Fagraea racemosa Jack ex Wall. Sqchnos avillaris Co1ebr.c

Memecylon rnegacarpum Furado Pternandra echinata Jack

Ficus annulata B1. B depressa BI. I? globosa BI. E magnokaefolia BI. E scortechinii King B sunduica BI.

Ebenaceae

Elaeocarpaceae

Legurninosae

Loganiaceae

Melastornataceae

Moraceae

Myrtaceae Eugenia chlorantha Duthie E. grzfithii Duthie Eugenia sp. 1 Eugenia sp. 2

Strombosia javanica BI.

Pellacalyx saccardianus Scort.

Prunuspolystachya (Hk. f.) Kalkman

Diplospora malaccensis Hook. f. Nauclea oficinalis (Pierre ex Pitard)

Olacaceae

Rhiwphoraceae

Rosaceae

Rubiaceae

Merr. & Chun Sapotaceae

Palaquium hispidum Lam I! obovatum (Griffith) Engler I! obovatum (Griffith) Engler Payena lucih (G. Don) DC.

Adinandra sarosanthma Mia. Theaceae

S U U U

S

C

C C

U L

U S

L L L C U L

C U C -

C

S

C

S S

C C C S

C

Fruit Fruit Fruit Fruit

Fruit

Fruit

Flowers Flowers

Fruit Fruit

Fruit Fruit

Fruit Fruit Fruit Fruit Fruit Fruit

Fruit Fruit Fruit Fruit

Fruit

Fruit

Fruit

Fruit Fruit

Flowers Flowers Fruit Fruit

Fruit

Ba.ma', CybB Ba.ma'9 2, Ch.mZ Ba. ma' Ba. ma'

Ba. ma', Ch. m& 3

Cyb?, Cy.hoZ, DyspZ, Me.e$, Ro.an?

Ba. ma'* Ba. ma', 3

Ba. ma' Ba.ma'. 3, CybP

Dysp2. DyspZ. * Ba.rnal.3, Ch.mB, Cy.bP Cyb?, Me.e$ Ba.ma'* 2- 3, Ch.mZ* 3, Cyb? Ba. ma'

Cyb?, Me.e$ Ba.mal Cy b? Ba.mal

Cy b?

Ba.ma', 3, Ch.mB, Cy.bP

Cyb?, Dysp2, Me.e$

Ba. ma' Ba.ma'. 3, CybB

Eo.sp2. Eo.s$ Cyb?, Eo.s$~ 4, Mr.e$, Me.wZ Cy btA, Cy ho4

Ba.ma', Ch.m4, Gbt2


APPENDIX Continued.

Growth Food Fruit bat Plant species forma w e speciesb

Unidentified seed 1 - Fruit Ba. ma' Unidentified seed 2 - Fruit Ba. ma' Unidentified seed 3 - Fruit Ba. ma1 Unidentified pollen - Flowers Ba. m d Unidentified leaf - Leaf- Cy 6 8

Unknown

a Plant growth form: U = understory tree; S = subcanopy tree; C = canopy tree; and L = liandstrangler. Fruit bar species: Ba.ma (Balionycteris rnanrlata), Chme (Chironax melanocephalus), Cy. br (Cynopterus brachyotis),

Qho (Cynopterus horsfiekizl, Dysp (Dyacopterus spadiceus), E0.p (Eonycteris spekzea), Me.ec (Megaerops ecauaktus), Me. we (Megaerops wetmorez), Ro.am (Rousem ampkxicauaktus). Numbers refer to methods of dietary analysis described in the text: 1 = roost and feeding perch collections; 2 = mist netting; 3 = chance collections; and 4 = nighttime observations.

Nomenclature follows Leenhouts (1972).

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