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234 Journal of Tropical Forest Science 14(2): 234-263 (2002) THE ROLE OF SELECTED ANIMALS IN POLLINATION AND DISPERSAL OF TREES IN THE FOREST: IMPLICATIONS FOR CONSERVATION AND MANAGEMENT S. S. Lee, Norsham Suhaina Yaakob, K. S. Boon & L. S. L. Chua Forest Research Institute Malaysia, 52109 Kepong, Kuala Lumpur, Malaysia Received January 2001_______________________________________________ LEE, S. S., NORSHAM SUHAINA, Y., BOON, K. S. & CHUA, L. S. L. 2002. The role of selected animals in pollination and dispersal of trees in the forest: implications for conservation and management. The current state of knowledge of the role of some plant-animal interactions in forest regeneration and conservation in Peninsular Malaysia is discussed. In particular, we focused on selected insect pollinators and animals which act as seed dispersers, for example, bats, primates, squirrels and birds. Tree species, including timber trees and endemic tree species utilised by bats, primates, squirrels, birds and selected insect pollinators are listed by their scientific as well as local names. The forest types where such species can be found are also identified. Some of the shortcomings of the currently available data are highlighted and means to overcome the situation are suggested. Keywords: Plant-animal interactions - pollinators - seed dispersers - conservation - forest regeneration LEE, S. S., NORSHAM SUHAINA, Y., BOON, K. S. & CHUA, L. S. L. 2002. Peranan sesetengah haiwan dalam pendebungaan dan penyebaran pokok di hutan: implikasiuntukpemuliharaandanpengurusan. Artikel ini membincangkan maklumat terkini mengenai peranan beberapa interaksi tumbuhan-haiwan terhadap regenerasi dan pemuliharaan hutan di Semenanjung Malaysia. Khususnya, kami menumpukan perhatian kepada beberapa serangga pendebungaan dan haiwan yang berperanan sebagai agen penyebaran biji benih, contohnya, kelawar, primal, tupai dan burung. Spesies pokok, termasuk pokok kayu balak dan jenis endemik yang digunakan oleh kelawar, primal, tupai dan burung disenaraikan mengikul nama saintifik serta nama tempalan. Jeni&jenis hutan yang terdapat pokok-pokok lersebul juga dikenal pasti. Beberapa kekurangan dari segi data semasa dilekankan dan cara-cara unluk mengatasinya dicadangkan. Introduction Plants have evolved a variety of mechanisms to ensure their successful pollination, dispersal and colonisation of suitable habitats. Some plants depend on wind while others depend on insects and animals for seed dispersal. An animal species may be the pollinator or dispersal agent of a specific plant species only or it may pollinate and disperse many different plant species. Plants, on the other hand, not only provide sources of food for insects and animals but also function as nesting and roosting sites. Less obvious, but more importantly, plants and trees provide organic matter and channel energy into the ecosystem through photosynthesis while animals play a very important role in decomposition and distribution of organic Journal of Tropical Forest Science 14(2): 234-263 (2002) 235 matter. However, the myriad and complex plant-animal relationships are not the main topic of this paper and are best elaborated elsewhere. Suffice to say that these relationships are even more complex in the tropical rain forests where the diversity of flora and fauna is very high. It is estimated that there are about 15 000 species of flowering plants in Malaysia. Pollinating and seed dispersal agents are called "mobile link" species (Gilbert 1980, Whitmore 1990) and they perform highly critical functions in the forest as they bridge otherwise unconnected components of the forest ecosystem (Whitmore 1990). In contrast to mobile link species, "keystone species" are plants that provide critical food resources for animals during annual periods of food scarcity which, if prolonged, may cause the extinction of those species which depend on these resources (Leighton & Leighton 1983, Terborgh 1986, Whitmore 1990). Logging changes the forest structure, thereby altering the availability of food and other resources for the animals. Reduced food resources may impact animal abundance and foraging behaviour, which in turn could have an effect on forest regeneration. Reduced populations or the disappearance of such frugivore species may result in the disappearance of plants that depend on these animals for seed dispersal. This paper focuses on the currentstate of knowledge of selected insect pollinators and animals that act as seed dispersers and their implications for forest conservation and regeneration. The paper also highlights some of the shortcomings of currently available data and suggests means to overcome the situation. Insect pollinators of trees in the forest Insects are considered the most diverse group of living organisms in the world and an estimated 150 000 species are known from Malaysia (Anonymous 1997a). Insects play a variety of roles in the forest, ranging from pests of plants to agents of biological control. They also play a very important role as pollinators of many forest tree species. The bees, in particular the trigonids, and aphids are among the major insect groups responsible for the pollination of many plants in Malaysian forests; 65 species of timber trees and 19 species of non-timber trees have been reported to be pollinated by bees (Appendix 1). Thrips form another important group of insect pollinators (Appanah & Chan 1982, Momose et al 1997). They are known to pollinate 23 species of timber trees and 13 species of non-timber trees in Malaysian rain forests (Appendix 1). Beetles are the third most important group of insect pollinators, they are reported to pollinate 35 species of timber trees and 3 species of non-timber trees in Malaysia (Appendix 1). Butterflies and moths (lepidopterans), insects in the fly order (dipterans), grasshopper order (orthopterans), cockroaches (blattodeans) and true bugs (hemipterans) are also pollinators of some forest trees. In view of the high plant diversity and the limited number of studies that have been carried out in Malaysian forests, the figures reported here represent only a fraction of the insect pollinators to be found in our forests. Some insects are host specific, pollinating only one species of plant, e.g. the •gaonid wasps which pollinate figs, but others, like the bees, are generalists. Bees are known to be important pollinators of a number of timber trees, including 236 Journal of Tropical Forest Science 14(2): 234-263 (2002) DryobaUmops spp., Neobalanocarpus heimii, Vatica spp., Shorea section Ovaks (Appanah 1987),Sindoravelutina(Appanah 1985) andKoompassiaspp. (Sakai^at 1997a).In a study of 43 timber tree species at Lambir, Sarawak, about 19 species were reported to be pollinated by bees (Sakai et al 1997a). One species of bee can pollinate many species of plants and sometimes one plant species can be pollinated by many species of bees, e.g. Dryobalanops aromatica (kapur) is known to be pollinated by 10 different species of bees (Sakai et al. 1997a). However, such detailed information is only available for a limited number of tree species. Information on the foraging patterns and feeding behaviour of the insects remains fragmentary. Bats and birds as pollinators of trees in the forest Some species of bats are known to be the main pollinators for species of economically important forest trees. Pteropodid bats, Eonycteris spelaeaand Macroglossus minimus, are known as the main pollinators ofDurio spp. which are important both for fruit as well as for timber (Start & Marshall 1976, Soepadmo & Eow 1976, Gould 1977, 1978, Marshall 1983). Two other species of pteropodids, namely, Pteropus vampyrus and Cynopterussp., have also been reported to feed on flowers of Duriospp. (Gould 1977, Fujita & Tuttle 1991) but they are probably not important as pollinators as their visits are irregular and destructive. Pollen from 32 plant species have been found in the droppings of the three nectarivorous bats, E. spelaea, M. mMimus and M. sobrinus (Start 1974, Start & Marshall 1976, Soepadmo & Eow 1976). These include pollen from the timber tree species Bombax valetonii, D. zibethinus, Parkia spp. including P. speciosa (petai), Artocarpus spp., Palaquium spp. (nyatoh) and Sonneratiaspp., indicating that these bats are important pollinators for these tree species. Among the birds, sunbirds and spider hunters (Nectariniidae) as well as white eyes/spectacle birds (Zosteropidae) are known as pollinators. These birds are small, active, arboreal birds that feed on nectar and small insects, as well as spiders. Wells (1988) recorded 10 species of sunbirds from the lowland forest of Peninsular Malaysia and observed that all spider hunters and Aethopyga as well as Nectarinia sunbirds of the lowland forest also range deeply into montane forests. However, although the characteristics of bird-pollinated flowers are well known (e.g. Meeuse 1961, Faegri & van der Pijl 1966), comprehensive data on the species of trees pollinated by birds is not available. Birds are known to pollinate at least 34 species of timber trees and 119 species of non-timber trees, including many species of climbers and palms (Appendix 1). Frugivorous mammals and birds A frugivore is defined as an animal whose diet is composed of 50% fleshy fruits (Terborgh 1986) and in this paper we use the term frugivore to loosely refer to those animals that are both obligatory and opportunistic fruit feeders. We focus on fruit eating bats, primates, squirrels and birds as these animals can be considered as seed dispersal agents. Journal of Tropical Forest Science 14(2): 234-263 (2002) 237 Fruit eating bats Bats make up about one third of the total number of mammalian fauna in Peninsular Malaysia and are divided into two suborders, the Megachiroptera and Microchiroptera. The former mainly feed on fruits with a few species feeding on nectar and pollen while the latter is insectivorous and/or carnivorous. In Malaysia, the role of bats as seed dispersal agents is still poorly understood as few studies have been conducted. One species that has been the subject of several studies is Cynopterus brachyotis which feeds on fruits. This bat is considered a potential seed dispersal agent because it carries fruits to feeding roosts located 50-70 m away from the fruiting tree (Funakoshi & Zubaid 1997). Fruit remnants, intact seeds, chewed leaves and flowers have been found underneath such roosts (Phua & Corlett 1989, Tan et al 1998). The role of bats as pollinators is, however, well known. From direct observations and fecal samples, bats are known to pollinate many cultivated and forest trees, including timber species (Appendix 1). Based on past studies, at least 80 plant species from 27 families of trees are known as sources of food (fruits, flowers, nectar) to bats (Appendix 1). Of these, 36 species from 17 families are timber trees and two, Madhuca selangorica and Payena maingayi (nyatoh durian),are endemic species. Both these species are from the family Sapotaceae and can be found in lowland forests. The latter can also be found in hill forests and is also a timber tree species. Since data on the food habits of bats in primary forests are scarce, we presently only know of a small number of economically important plant species that may be pollinated or dispersed by certain species of bats. Primates In Peninsular Malaysia, the primate fauna comprises one prosimian (Nycticebus coucang), three gibbons (Hylobates agilis, H. lor and H. syndactylus), three macaques (Macaco arctoides, M. fascicularis and M. nemestrina) and three langurs (Presbytis cristata, P. melalophos and P. obscura). Very little is known of the small, solitary and nocturnal slow loris (N. coucang) and it is not surprising that its role in seed dispersal has yet to be determined. Of the three gibbon species, the siamang (H. syndactylus) mainly feeds on leaves while the other two species feed on fruits. Figs were found to be an important source of food for all three species of gibbons (Chivers 1974, Gittins & Raemakers 1980). Macaques are reported to be omnivorous, feeding on fruits, leaves and insects (Harrison 1961, Lim 1968, Medway 1983). Of the three langurs, P. cristata is mainly folivorous (Kool cited in Laidlaw 1994). Half of the diet of P. obscura is made up of leaves while half of the diet of P. melalophos is made up of fruits and seeds (Curtin 1980). Primates obtain food from a total of 410 species of plants, of which 222 are timber species. Plant families which have a high number of species utilised by primates for food are Moraceae (33 species), Annonaceae (32 species), ^uphorbiaceae (29 species), Leguminosae (22 species), Meliaceae (19 species), Guttiferae (16 species), Anacardiaceae (15 species), Burseraceae and Ebenaceae 238 Journal of Tropical Forest Science 14(2): 234-263 (2002) (14 species each), and Sapindaceae (13 species). Of the plants consumed by primates, 23 are endemic species (Appendix 1). However, this data is probably still far from complete. Figs have been proposed as important keystone species for primates based on their characteristics and prominence in the diet of primates, i.e. large fruit crop and continuous availability due to the large number of species present (Raemakers et al 1980). Studies in the lowland forest at Kuala Lompat, Pahang, Peninsular Malaysia, showed that there are plants other than figs that contribute to the diet of primates during seasons of low food availability and that some of these species are available almost all year round (Curtin 1980, Bennet 1983). It would appear that figs are not the keystone species for most primates with the exception perhaps of the siamang (H. syndactylus) where our observations show that figs makeup the largest portion of their diet compared with other plant species. Yasuda (1998) reported that figs were not the keystone species for small mammals at Pasoh Forest Reserve, Negeri Sembilan, Peninsular Malaysia as their diversity and density were much lower than at Kuala Lompat. Similarly, Gautier-Horn and Michaloud (1989) found that figs were not the keystone species for mammals in Gabon due to their low densities, unpredictable fruiting patterns and low crown production. There is presently very little information on the role of primates in seed dispersal. Among them, gibbons are the most likely seed dispersal agents because of their feeding characteristics. They only eat the ripe pulp of fruits, swallowing seeds of most species and excreting them whole, and often at great distances from the source trees. They also often revisit food sources, making them reliable seed transporters. Another potential seed dispersal agent is the long-tailed macaque (M. fascicularis) which often drop and spit out larger seeds, and carry more fruit away from the fruiting tree (Corlett & Lucas 1990). In contrast, most of the other primates are highly opportunistic and tend to destroy seeds during feeding (Gittins & Raemakers 1980). Squirrels The importance of squirrels as seed dispersal agents is unclear as there is data showing both their roles as seed predators as well as seed dispersal agents. Some species of squirrels are also known to consume unripe fruits which would make them unlikely as effective dispersal agents. The dentition of squirrels also allow them to gnaw through hard tissue thereby destroying the fruit embryo (MacKinnon 1978). On the other hand, some species of squirrels are known to drop seeds and/ or consume the seeds far away from the fruiting tree and dropping some during transport. Some squirrels also bury seeds in the ground and unrecovered seeds could germinate and contribute to regeneration far away from the parent tree. More in-depth studies are needed to elucidate the role of squirrels in seed dispersal. A total of 235 species of trees from 49 families are known to provide sources of food for squirrels. Of these 160 are timber species and 14 are endemic species (Appendix 1). Important families are the Euphorbiaceae (21 species), Leguminosae and Annonaceae (ISspecies each), Meliaceae and Anacardiaceae (16 species each) Journal of Tropical Forest Science 14(2): 234-263 (2002) 239 and Burseraceae and Myristicaceae (10 species each). Squirrels generally eat the fruits and seeds of trees but leaves, bark and flowers are also sometimes consumed by some species. Birds Frugivorous birds are considered important mobile links by functioning as pollinators and seed dispersal agents. In Peninsular Malaysia, there are 19 species of obligate fruit eating birds, 13 of which are found in the lowlands and six in montane forests (Wells 1988). Of these 19 species, 10 species of pigeons and three species of parrots feed solely on fruits, seeds and flowers. Birds in the following families maybe considered partially orwhollyfrugivorous: Bucerotidae (hornbills), Campephagidae (minivets), Capitonidae (barbets), Chloropseidae (leafbirds), Columbidae (pigeons), Corvidae (magpies and crows), Dicaeidae (flowerpeckers), Irenidae (Asian fairy bluebird), Oriolidae (orioles), Phasianidae (pheasants), Psittacidae (parrots), Pycnonotidae (bulbuls), Sturnidae (hillmyna) andTimaliidae (babbler). Birds are highly dependent on trees not only for food but also for nesting and perching and some birds feed on insects found on the trees. In a study conducted in a hill dipterocarp forest at Sungai Tekam Forest Reserve, Pahang, Peninsular Malaysia, Johns (1983) found that 22 species of birds were intolerant of forest disturbance. These birds are found in unlogged forest but not in older logged forests. Another 20 species only occurred in logged forests and these were categorised as colonising species. Based on his study of fig-eating birds in a lowland forest at Kuala Lompat, Lambert (1989) proposed figs as keystone resources for birds because of their abundance, aseasonal fruiting, enormous crop size, distinctive intra-crown synchrony of fruit ripening, short intervals between fruiting by individual trees, and ease of harvesting by different groups of frugivores. Another important factor is that Ficus is the only known plant taxon to which frugivorous birds in Southeast Asia have specialised (Lambert & Marshall 1991). In comparison with the figs, detailed information on other tree species is lacking. Many trees on which birds depend for food have only been identified to genus level making it difficult tojudge the importance of a particular tree family in providing sources of food for the birds. Presently about 153 species of trees from 54 families are known to provide sources of food to various frugivorous birds (Appendix 1), with the Moraceae being the most important family by far. Other tree families which are important as food sources for birds are the Palmae, Burseraceae and Sapindaceae. From the above discussion it is clear that our understanding of pollination and seed dispersal of tropical rain forest trees is very limited. Discussion It has been increasingly recognised that within a tropical rain forest, an incredible number of linkages exist between plants and animals, from the food chain to dependence on pollination, dispersal, provision of roosts and nest sites, 240 Journal of Tropical Forest Science 14(2): 234-263 (2002) chemical defense and other functions. It is feared that if some of these links or interconnections are severed, whole ecosystems may collapse. For example, certain species of bats are the pollinators and dispersal agents for a large number of trees in the forest. If these bats become locally extirpated or severely decimated in numbers as a result of forest harvesting, a whole host of bat-dependent plants could become reproductively impotent. Such an event would threaten the successful regeneration of that group of plants. In addition, if that threatened plant group is economically useful, the fiscal value of the forest would also be considerably reduced. Therefore, for sustainable forest management, tree species that play important roles in animal-plant inter-relationships and that provide important sources of food or habitats for the mobile links, need to be recognised and retained or saved from damage. An effort to retain some important food source trees during forest operations has been made by the Forestry Department, Peninsular Malaysia. The department's ManualKerjaLuarSistemPengurusanMemilih (Anonymous 1997b) is a commendable operations manual containing guidelines, rules and procedures aimed at guiding the staff of the Forestry Department in performing their duties and responsibilities in an organised, uniform and effective manner. Among many operational guidelines, this manual contains a list (on page 261) of forest trees which are sources of food for animals. These trees have been reported elsewhere as sources of food for primates and other animals and are not permitted to be girdled except in circumstances where they are likely to have an important impact on the regeneration of the residual stand. We believe that with some modification and improvement, this list can be a very important and useful tool for forest conservation and management purposes. Firstly, the references from which the list were derived should be provided so that the accuracy of the data can be verified. The present list is rather general as the groups of primates, squirrels and birds referred to are not stated. It is known that not all primates, squirrels and birds act as dispersal agents; manyjust consume and destroy fruits and seeds. Many other important animal food source trees need to be included in the list. In addition it is important to include trees which are vital to specific groups of pollinators whose presence and activity ensure the success of the regeneration of our desired crop trees as well as the continued functioning of the forest ecosystem. Below we list some suggestions for improvement and usage of the aforementioned list. An example of an improved list is shown in Appendix 1. However, we would like to stress that our list is still far from complete or perfect and that review and modifications would be necessary as more comprehensive data become available. (1) (2) (3) Local names of trees and forest types where the trees are found should be included. The list can be made more specific to include only those species which are known to be pollinated and dispersed by the identified mobile link species. (The list shown here is rather extensive and may need to be trimmed down). Timber and endemic trees as well as non-timber trees should be noted/ highlighted. Journal of Tropical Forest Science 14(2): 234-263 (2002) (4) (5) (6) (7) 241 As the list is to be used in Peninsular Malaysia, Bornean species need not be included. Sources of references for the list of tree species and their mobile links should be cited for ease of reference and verification of data. The inclusion of phenological data on flowering and fruiting seasons should be considered, perhaps as an additional appendix, as such information is important to indicate which sources of food are available when. Data on the range of the mobile link species need to be collected as this is linked to the issue of forest fragmentation. By so doing, the list can be made (8) more concise with the inclusion of only a minimum set of important or critical food source trees. Training must be provided to field staff so that they are able to recognise these target trees. However, this is for the future. The more immediate need is to obtain detailed information on the mobile link species and keystone species in our forests so that a minimum set of species can be identified to ensure the continued regeneration and sustainable management of our forests. Conclusions Our understanding of the pollination and seed dispersal of many tropical rain forest trees is still very limited. Although we know that the Leguminosae, Moraceae, Annonaceae, Euphorbiaceae and Meliaceae are among the more important plant families that provide food for bats, primates, squirrels and birds, such data is insufficient to elucidate the effectiveness of these animals as efficient pollinators and/or seed dispersal agents. 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Ph.D. thesis, University of Tokyo, Japan. 90 pp. Appendix 1 Family Species Alangiaceae Anacardiaceae Annonaceae Tree species utilised by bats, primates, squirrels, birds and selected insect pollinators for food (timber (T) and endemic (E) tree species) Local name Forest type Alangium ebenaceum (T) Alangium ridleyi (T&E) Mentulang daun bujur Mentulang daun lebar Bouea inaquifolia Bouea macrophylla (T) Bouea oppositifolia (T) Buchanania arborescens (T) Buchanania sessifolia (T) Campnosperma auriculatum (T) Dracontomelon dao (T) Gluta elegans (T) Mangifera gracilipes (E) Mangifera griffithii (T) Mangifera indica (T) Mangifen laurina (T) Mangifera macropkytta (T) Mangifera magnifica (T&E) Mangifera quadrifida (T) Mangifera spp. Melanorrhoea fulvinervis (T) Melanorrhoea inappendiculata Melanorrhoea malayana (T) Pentaspadon velutinus (T) Semecarpus sp. Alphonsea elliptica (T) Annona squamosa Cananga odorata (T) Cyathocalyx carinatus Cyathocalyx pruniferus (T&E) Cyathocalyx scortechinii (E) Cyathocalyx sp. Deswxwsp. (liana) Fissistigma sp. Mezutia parviflora (T) Monocarpia marginalis (T) Afeo-UDoria/ortido Synonym D. mangiferum M. microphylla M. longipes Primates Squirrels LDF LDF + + Kundang Kundang hutan Kundang rumenia Otak udang daun tumpul Otak udang daun tajam Terentang daun besar Sengkuang Rengas Macang hutan daun halus Rawa LF LDF, HDF LDF, HDF CF, RV LDF, HDF LDF, MF LDF, RV LDF + + + + + + + + Mangga Cultivated Macang api Macang temuor Macang Macang LDF LDF, HDF LF Macang Rengas Rengas Pelong beludu Bats Birds Bees Thrips Beetles + + LDF + + + C/5 + + + + + $ I—» 5 + + + + + + + + + + + + + + + + + + + + OS + + LF •f + LDF, HDF NS o o NO + Mempisang, chaget Buah nona Kenanga Antoi daun kecil Antoi Antoi Antoi LDF Cultivated LDFfm LF LDF, HDF LF Akarlarak M. leptopoda Mempisang Mempisang LDF LDF + + + + + + + + + + + + + + + + + + (continued) Journal of Tropical Forest Science 14(2): 234-263 (2002) 246 •I a x. I o I^ 3 "2 a a s U, U." ST b," 1* U, 2 IK" 99999959 3 bo.C bo bo bo S bo bo S-S. bo gu aw aw .1 g- a E .1 .2 .. a. a a. be .a..- '5. a. ||§ M bo bo I sl,J n, u. u, u, 99 99 p ri tit (Sis 3 ;§. Lff 'in •& i OH »—^ M~J o B •a o u t 1a Journal of Tropical Forest Science 14(2): 234-263 (2002) 247 •o § " is a U. -o u. Q Q In LL, 33 aS a3 3333333 bo bo bo bo bo bo bo ill - S § c c c c 1 5 5 •o <u a •a 8 .§• I Journal of Tropical Forest Science 14(2): 234-263 (2002) 248 + + + + + . to, [* I II nil 332! u, 9S95 b99 u. 9 9 u. 9: 60 -r 8 1 .2 ill tl-«. JSJJ | S 5 O" II \ £ 2 11 tit in co in •a § 1 It 249 Journal of Tropical Forest Science 14(2): 234-263 (2002) t I IK Q It* Q IK IK Q Q 9 99 IK •r « Q -C IK Q IK 9 IK u. b< Si S X X IK 5 § Q Q IK IK IK IK IK IK 9 9 9 9 SSS c '3 1 •o 11 Si £.= i p s sa i •^ •a -a •-n 60 1 1 ii •3 3 1.3 S I* •3 § •" § S ?-B Jlli *• —— •-!•-( •a -a -a d* 1 •a 8 S 99 c" § Journal of Tropical Forest Science 14(2): 234-263 (2002) 250 JT § X 93 1 a Si III i r b o b c b o b o b o b o b o b o b o be bo be bo g 3 1 2 0 H H S 1 •a § u 2 M I S ^ ? •n U c Id §• U | i i 251 Journal of Tropical Forest Science 14(2): 234-263 (2002) + + + + u. to . to to sx sa ss K -e fc o t. t n U. b 99b"a&959959 h- 11 to to to" to" tolbto" 3 3 3 3 S139 939 1 i 2 to to 3 •a o u 369 I S ""S nil Kb bo So be c c c c 2 S S S 999 § -a it i SAS. = 11 S E E S ^ ^ 111 c '5 '5 s PS6 P •o v lb to 1 5 to- Journal of Tropical Forest Science 14(2): 234-263 (2002) 252 •c 03 JS '3 1 , . 939 H fe Q a si b." 9 sa 2 ft Q *•• , [i, tH ftQ - fcti UH fcU W 3C 99999SS = ii I §9 9 a ^s e bO'fl "O bo bo bo S. ss ISS 6 I o I •a o u 0 a .3 Appendix 1 (continued) Family Species Lauraceae Actinodaphne sp. Alseodaphne sp. Synonym Local name Forest type Bats Primates + Berambong Squirrels + Birds Bees Thrips Beedes + + •S | ^3" s£. daun satu Beilschmiedia spp. Cinnamomum men (T) Cinnamomum sp. (T) Cryptocarya sp. Dehaasia eUiptica (T) Dehaasia incrassata (T) Dehaasia polyneura (T) Zfeftoorio spp. Endtandra sp. Z-itwo spp. (T) Machilus sp. Nothaphoebe umbelliflora (T) Lecythidaceae Leguminosae Medang Medang teja Gamak Medang Medang Z). efli/rtico AreAtoVndnm clypearia Archidendron contortum Archidendron jiringa Bauhinia purpurea CaUerya atmpurpurea (T) Cassiafistula Cassia javanica ssp. nodosa (T) Casiia spectabilis Cynometra malaccensis (T) ZJemjsp. (liana) Dialium indum var. mdum (T) LF, HF LDFrv LDF Medang Medang Medang + + + + + + + + + + + + + + + Saga Kerdas LDFfm LF, cultivated LFb Gonderik buah merah LF + + Gonderik buah hitam LF + + Jiring LF, cultivated Tulang daing Cultivated Open areas + Milletia atmpurpurea Cultivated + C. nodosa Bebusok + C. inaeauifolia Kekatong D. laurinum. Keranji paya LDF "NO ^ + + LF, HF, RV LDF LDF Cultivated LDF, HDF Co h—' Putat Putat PitheceUobium bulbalinum PitheceUobium clypearia PitheceUobium contortum PitheceUobium jiringa K I LDF to MF A. pennata §] + + + Medang fiam'ngtanta spp. Planchonia valida (T) Acacia ameinna (liana) Adenanthera bicolor Archidendron bubalinum (T) + + LDF, HDF | w f NO O) OS + + NS O o + -IS + + + + + + + + + + + + + + + + + + D. patens Dialium kuntsleri (T) (Borneo) + to (continued) 254 Journal of Tropical Forest Science 14(2): 234-263 (2002) i .a. 1 s + + E + I 111? b ft Q (i. te. U. u. X XXX X X fcSSSa & 9 u." U« (b 3 33 1 ri |i I3 '3 £ aj aj flj >£££ u. Q Q § c 2 * q 8. 3 9o 3 03 I i p rt io If e i* _ o u Journal of Tropical Forest Science 14(2): 234-263 (2002) u. is 2 »B s." 3 3 313 255 ss s gBK * tt." uT 9 S333 9 •« bo ! •o I •a § ' 3 NS Appendix 1 (continued) Family Species Dysoxylum cauliflorum (Borneo) (T) Dysoxylum spp. Lansium domaticum (T) Metiasp. Keinwardtiodendron humile Sandoricum koetjapi (T) Sandaricum sp. Walsura pinnata (T) Menispermaceae Fibraureasp. (liana) Moraceae Antiaris toxicaria (T) Artocarpus fulvicortex (T) Artocarpus integer var. sihestris Artocarpus lanceifolius (T) Artocarpus loom (T) Artocarpus maingayi (T) Artocarpus rigidus (T) Artocarpus scortechinii (T) Artocarpus spp. Synonym Local name Forest type Mersindok Langsat Mostly LF LDF, cultivated Primates Squirrels Birds + + + + + Bees Thrips Beetles + + + Sentul W. neuroides Mersindok LDF, HDF LDF, HDF LF LDF, HDF + + + + + LDF LDF LDF to MF LDF, HDF LDF LDF LDF, HDF LDF + + + + + + + + + + + + + + + + + + + + Ipoh Keledang tampang gajah Cempedak Keledang keledang Miku Pudu Keledang tampang Terap hitam + + + fieuj annulata (T) Ftcus aurantiacea Ficus benjamina Ficus binnendykii Ficus bracteata (T) Ficus callophyUa Ficus caulocarpa Kuap Ara Ara Ara Ficus consociata Ficus CFasstramea Ficus cucurbitma Ficus delosyce FKUS drupacea Ficus dubia Ara Ara Ara Ara Ara Ara Ara Ara Ara Ara Ficus fstulosa (T) Ficus heteropkura Ficus kerkhovenii Ficus obscura\ar. bomeensis (T) Ficus parietalis Bats Ara Ara Ara F. obscura Ara LDF LDF LDF, cultivated LDF LDF + Wild and cultivated LF, RV, seashores LF, villages BF LDF LDF, HDF LDF SF, wayside LDFtoMF LDF to MF LDF LDF + + + + + + + + + + + + + + + + + + + + + + + + f (continued) Appendix 1 (continued) Family Species Synonym Ficus pdlucidopunctata Ficus pisocarpa Ficus religiosa Ficus roxburghii Ficus ruginervia Ficus sagittata Ficus stricta Ficus stupenda Ficus subcordata var. malayana Ficus subulata Ficus sumatrana Ficus sundaica (T) Ficus sundaica var. sundaica Ficus trichocarpa Ficus vasculosa (T) Ficus virms (T) Ficus vhmsvai. gfobeUa Paratocarpus bracteatus F. subcordata Local name Forest type Ara Ara Ara Ara Ara Ara Ara Ara Ara Ara LDF LDFrv Cultivated + Cultivated + Ara F. indica F.glabbeua Ara Ara Ara Ara Ara Ara Berteh Myricaceae Myricasp. Myristicaceae Gymnacranthera sp. HorsfieUia irya (T) HorsfieUia pofyspherula (T) Squirrels LDF LDF LDF HDF to MF HDF LDF, HDF LDF LDF LDF LDF, HDF LDF LDF LDF, HDF LDF, HDF M. malaccensis M. truncata Pisang Pisang Pisang LDF to LMF MF Cultivated Telur cicak CF Birds Bees Thrips Beetles + + + + + + + + + + + + + + + + + + + + + + + + + + + + Mostly LF Streblussp. Musa acuminata ssp. malaccensis Musa acuminata ssp. microcarpa Musasp. Primates + Tebakah Sloetia elongata (T) Musaceae Bats + + + + + + + Pianggu, penggu LDF, lowlying RV HorsfieUia sucosa (T&E) HorsfieUia supabam HorsfieUia spp. Penarahan Penarahan Penarahan Penarahan Knema ctnerfa Basong paya Knema fiafitracea (T) Penarahan Knema hookcriana (T) Knema lamina (T) Knema spp. Myristica gigantea (T) Penarahan Basong bulu LF LDF, HDF LDF Penarahan Penarahan arang bukit LDF LDF + + + + + + +• + + + + + + + + + + + + + + + + + + + + + + + + (continued) B? Journal of Tropical Fonst Science 14(2): 234-263 (2002) 258 8. Q II < aa 99 9 9 3 & 99 1 - ii o § fl JB , c •a o t I I u 259 Journal of Tropical Forest Science 14(2): 234-263 (2002) si 1 I HH O Q O2 Q Sac 2 as 11 II b" U." b U." 99 1 I C„ rt bo O. js ^ •a o u C g £ <3 •§ Jj 9 9 9 9999 Ills JjJ! i Jj till 11 a I. 3 -1S 32s 3 23J2 260 Journal of Tropical Forest Science 14(2): 234-263 (2002) .8. I + + + £ a 2 2* IK IK IK IK 9 3 39 S 1' 11 a" 9 9 IK lx" IK IK 3 •! il ! ll !b t I IE 6 ffO.ST3 I 1 .8- 8 5 s-is--I* -^ S » - sS -3f&aJ l 8-311 -1 i it) k| . IK IK 939 9 99 s- 261 Journal of Tropical Fanst Science 14(2): 234-263 (2002) c 1 IK Q 3 J! 1 >K T3 -O (K K 3 5 K Q ss I a 3 3 S 3 3 33 § ^|J g-§ I 33*3 X 2 3 ssss! §3 3 I II -S-.I S bo i gs sa ll a E E -S P I P s-: 3 •a 8 I §c ie*! e- NO Appendix 1 (continued) Family Species O) NO Synonym Sonneratia caseolaris (T) Sonneratia auata (T) Sterculiaceae ftrmiana malayana Pterocymbium linclorium (T) P. javanicum Pterospermumjavanicum(T) Pteiygota alata(T) Scaphium bomeensis (T) Scaphium linearicarpum (T) Scaphium langipetiolatum (T) (Borneo) Scaphium macropodum^T) Sterculia foetida (T) Sterculia panrifolia (T&E) Sterculia stipulata Local name Forest type Bats Berembang Gedabu Mangroves Mangroves + +• Bayur Kasah + + + (Borneo) Kembang semangkok bulat LDF, HDF + LDF, HDF CF HDF + + + Kelumpang jari Kelumpang Beetles + + + + + + + + Taccaceae Taccasp. (herb) Theaceae Admandra ilamponga (T) Adtnonttra sp. Tiup-tiup Aauilaria malaccenas (T) Gmystyhts cmfusus (T) Ramin pinang muda Tr^gimia5<ruffl hypoteucum (T) Thrips + Kelumpang Styrax benzoin (T) Trigonaceae Bees + Kembang semangkokjantong Styracaceae Microcosantidesmifolia(1) Microaa UattaefoKa (T) M icrocos crassifolia Microaa fUmcarpa (T) Microau laurifolia (T&E) Microaa tomentosa (T) Microcossp. (5 spp) Pmtaccjloribvnda Pentace triptrra (T) Schoutenia comeri (E) Birds + Alluvial LDF, HDF LDFrv Symptoms sp. TUiaceae Squirrels RV, open forests Melembu Symplocaceae Thymelaeaceae Primates Kemenyan + * + § ^ + Kelat paya + + Karas + LF + Chenderai LF + + + G.fitrocarpa G. laurifolia G. tomentosa Chenderai, damak bulu Chenderai Chenderai tanjung LF LDF, HDF LDF, HDF fm + + + + + 4- 4+ + + Marajali S. §• g + G. blattaefolia Ba'ang Melunak pusatbeludu LDFrv a ^g. 1 ^ + + HDF LDF + e* ^ 'Jo '7^' to V? f + + JS &> + bo + 1 (continued) Appendix 1 (continued) Family Species Ulmaceae Celtis rigescens (T) Celtissp. Gmmnieria hirta (T) Gmmnieria neruosa (T) Gmmnieria parvifolia (T) Gmmnieria subaequalis (T) Gmmnteria sp. Trema oruntalis Tremasp. Urticaceae Verbenaceae Synonym Local name Forest type Semantit LDF, HDF Bats Squirrels + + Birds Bees Thrips Beetles + Hampas tebu Hampas tebu Hampas tebu LDF, HDF + Hampas tebu LDF, HDF + + Menarong LFb + + + Laporteasp. Poikttospermum sp.(epiphyte) VUIebruntasp. CaUicarpa maingayi Lantana cammara Teijsmanniodendnm spp. Vitexpinnata (T) Vitec trifoliata Vitexspp. Primates + + + + + Cultivated V. pubescens Leban SF, wayside Mosdy LF + + + + + Violaceae Knoreasp. + Vitaceae Vitissp. (liana) + + + + rsy t I + LDF = Lowland dipterocarp forest; LDFs = Swampy lowland dipterocarp forest; LDFfm = Forest margin; LFb = Belukar HDF = Hill dipterocarp forest PSF = Peat swamp forest LMF = Lower montane forest RV = Riverbanks MF = Montane forest SF = Secondary forest CHF = Coastal hill forest CF = Coastal forest BF = Beach forest p, s = primary, secondary Sources: Mcdure (1966), Madge (1969), Chivers (1974, 1980), Start and Marshall (1976), Curtin and Chivers (1978), MacKinnon (1978), Payne (1979), Appanah (1981, 1987), Appanah and Chan (1982), Bennet (1983), Johns (1983), Appanah « aL (1986), Caldecott (1986), Kiew and Davison (1989), Lambert (1989), Momose et aL (1996, 1997), Nagamitsu et aL (1997), Sakai etaL (1997a, b), Tan etal (1998). hO O> oo