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Fruit-Frugivore Interactions in a Moist Evergreen Forest of Khao Yai National Park in Thailand

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Fruit-Frugivore Interactions in a Moist Evergreen Forest of Khao Yai National Park in Thailand TROPICS Vol. 14 (4) Issued July 31, 2005 Fruit-frugivore interactions in a moist evergreen forest of Khao Yai National Park in Thailand 1)＊ 2) 1) 1) 1) 3) Shumpei KITAMURA , Takakazu YUMOTO , Pilai POONSWAD , Phitaya CHUAILUA , Kamol PLONGMAI , Naohiko NOMA , 4) 5) Tamaki MARUHASHI and Prawat WOHANDEE 1) Thailand Hornbill Project, Department of Microbiology, Faculty of Science, Mahidol University, Rama 6 Rd, Bangkok 10400, Thailand 2) Research Institute of Humanity and Nature, Kyoto, 602-0878, Japan 3) School of Environmental Science, The University of Shiga Prefecture, Hikone, 522-8533, Japan 4) Department of Human and Culture, Musashi University, Nerima, Tokyo 176-8534, Japan 5) National Parks Division, Department of National Parks, Wildlife and Plant Conservation, Phaholyothin Rd, Chatuchak, Bangkok 10900, Thailand ＊ Corresponding author. Department of Microbiology, Faculty of Science, Mahidol University, Rama 6 Rd., Bangkok 10400, Thailand E-mail address: [email protected], Tel: 66-2-201-5532, Fax: 66-2-644-5411 ABSTRACT Seed dispersal by animals plays a crucial role in the tropics. Fruit-bearing plants serve not only as nutritional sources for frugivores, but also as seed sources for forest regeneration and as important foci for the re-establishment of other plant species by attracting seed-dispersing frugivores to their vicinity. However, opportunities for investigating the interactions between a diverse fruit flora and disperser fauna are rapidly disappearing in Southeast Asia. We observed the behaviors of 28 species of frugivorous visitors to 15 fruit-bearing plant species in a moist evergreen forest in Khao Yai National Park, Thailand, to determine their potential quality as seed dispersers. Behavioral observations included the frequency and duration of visits by each forager and their fruit-handling techniques. The highest numbers of frugivores were recorded at strangler figs, confirming their role as an important resource for frugivores at Khao Yai. Mammals and non-passerine birds spent significantly more time at food plants than did passerine birds. Our study provides a preliminary inventory of plant-frugivore interactions that is comparable to other study sites in Southeast Asia. Since fruit-frugivore interactions may differ among the forests, these kinds of studies need to be replicated at faunally intact forests, as well as depleted forests, in Southeast Asia. Key words: frugivore, frugivory, fruit characteristics, seed dispersal, seed predation INTRODUCTION In the tropics, the majority of woody plant species bear animal-dispersed fruits (Howe & Smallwood 1982; Howe 1986; Fleming et al. 1987; Jordano 2000). These fruits may serve not only as resources to fulfill the nutritional requirements of frugivores (McKey, 1975), but also as seed sources for forest regeneration (McDonnell & Stiles, 1983; Elliott et al., 2002) and as important foci for the re-establishment of other plant species by attracting seed-dispersing animals to their vicinity (Alcantara et al., 2000; Schupp et al., 2002; Clark et al., 2004; Russo & Augspurger, 2004). In Southeast Asia, too, the species diversity of animal-dispersed plants is high (Ng, 1983) and most forest birds and mammals consume various kinds of fruits (Corlett, 1998b). Despite the massive anthropogenic habitat modifications in Southeast Asia (Laurance, 1999; Sodhi et al., 2004), studies on fruit-frugivore interactions are very limited (Corlett, 1998b). Information on fruit species’ preferences by most frugivorous animals may be insufficient to distinguish seed dispersal roles among particular frugivore species within an assemblage in many tropical forests in Southeast Asia. As many dispersal agents are highly vulnerable to hunting and habitat loss (da Silva & Tabarelli, 2000; Peres, 2000; Pattanavibool & Dearden, 2002), opportunities for investigating the interactions between a diverse fruit flora and an intact disperser fauna are rapidly disappearing in Southeast Asia (Corlett, 2002; Sodhi et al., 2004). Khao Yai National Park is regarded as one of the best conservation areas in Southeast Asia (Elliott, 2001). It still contains sizable populations of large frugivorous animals that were mostly depopulated in other areas of Thailand, as well as Southeast Asia. Most of the plants in the moist evergreen forest of Khao Yai National Park bear fleshy fruits and are thus potentially dispersed by animals (Kitamura et al., 2005). As a result of our previous work (Kitamura et al., 2002), interactions between fruits and frugivores in Khao Yai National Park is one of the best-studied examples in Southeast Asia. However, our previous approach was predicated on the assumption that frugivores within a single functional group 346 Shumpei KITAMURA, Takakazu YUMOTO, Pilai POONSWAD, Phitaya CHUAILUA, Kamol PLONGMAI, Naohiko NOMA, Tamaki MARUHASHI and Prawat WOHANDEE consume similar plants and therefore disperse similar seeds. However, specific dietary information is required to assess the validity of this kind of approach (Moran et al., 2004). Furthermore, previous studies related to seed dispersal by animals in Khao Yai National Park concentrated on large frugivores such as gibbons (Whitington, 1990; Whitington, & Treesucon, 1991) and hornbills (Poonswad et al., 1998b; Kitamura et al., 2004a; Kitamura et al., 2004b; Kitamura et al., 2004c), without considering the seed-dispersing role of small frugivorous vertebrates that are diverse and abundant in Southeast Asia (Corlett, 1998b). The major aim of this study was to obtain data on fruit preferences of frugivores, including small frugivorous vertebrates that were mostly ignored in the previous studies conducted in Khao Yai National Park. To this end, the frequency and duration of frugivore visits to fruiting plants were recorded, and fruit-handling techniques are described. More specifically, we addressed the question of whether large-seeded plant species were visited by fewer disperser species than were plants with fruits or seeds that were more easily consumed in Khao Yai National Park. These trends are found mainly in Neotropical, African and temperate zone studies (Jordano, 1995), but there has been little work done on it in Southeast Asia. This study also provides a reference of fruit-frugivore interactions for ongoing ecological research and conservation in Khao Yai National Park, as well as Southeast Asia. MATERIALS AND METHODS Study site This study was conducted in Khao Yai National Park (hereafter KY) in northeastern Thailand from June 1998 to August 1999. The park lies between 14º 05’‒14º 15’ N and 101º 05’‒101º 50’ E in the Dongruk mountain range, and covers an area of 2,168 km2 (Fig. 1). The elevation of the park ranges from 250 to 1,351 m. The study site is located in the area adjacent ����� ������� ������� ���� �������� �������� ���������������������� Fig. 1. Location of Khao Yai National Park in relation to neighboring countries. to the park headquarters and covers approximately 70 km2. The study site ranges from 600 to 800 m in altitude and was dominated by moist evergreen forest, which covered approximately 64 % of the total park area, or 1,375 km2, between 400 and 1,000 m in elevation (Smitinand, 1977). In KY, at least, 72 mammal species (25 % of species recorded in Thailand) and 340 bird species (37 % of bird species recorded in Thailand) have been reported (MUCDC, 1989; Lekagul & Round, 1991; Srikosamatara & Hansel, 1996). The dominant plant families in the forest include the Lauraceae, Cornaceae, Euphorbiaceae, Meliaceae, Elaeocarpaceae, Dipterocarpaceae, and Annonaceae. Kitamura et al. (2005) provided a detailed botanical inventory. The mean annual rainfall from 1993‒2002 was 2,360 mm, and the wet season usually occurs Fruit and frugivore assemblages in Thailand 347 from April to October. The dry season usually occurs from November to March (Kitamura et al., 2004a). The mean monthly temperature ranges from 21 ºC (December and January) to 32 ºC (April and May). Although some ripe fruits (Ficus spp.) are available year-round (Poonswad et al. 1998a), fruit diversity and abundance are relatively high during the rainy season and reach a low point at the beginning of the dry season (Bartlett, 2003). Plant observations We observed the feeding assemblages of frugivores during the peak fruiting period of 15 plant species (Table 1). These plant watches consisted of continuous 12-h observations (0600‒1800) of one individual plant per species. Plants were selected for study if frugivores had been observed to forage on them, or they had a ripe crop of fruit known, or suspected to be, attractive to arboreal frugivores. Since the most likely external variables to influence frugivore foraging Table 1. Fruit and seed characteristics of the fruit-bearing plants selected for study. FL = fruit length (mm); FD = fruit diameter (mm); CO = color of fruit; NO = number of seeds per fruit; SL = seed length (mm); SD = seed diameter (mm). Fruit type: FT: D = dehiscent; I = indehiscent fruit with thin husk; T = indehiscent fruit with thick husk. Life form: LI = liana; ST = small tree (7−15 m); MT = medium- sized tree (15−30 m); TT = tall tree (> 30 m). Family Species FL FD CO NO SL SD FT LF Season Code Annonaceae Platymitra macrocarpa Boerl. 80.6 63.6 Brown 7 28.2 16.7 T TT Feb.99 1 Celastraceae Bhesa robusta (Roxb.) Hou 18.6 11.2 Yellow 1 15.4 8.1 D TT Dec.98 2 Elaeagnaceae Elaeagnus conferta Roxb. 41.1 25.4 Red 1 33.0 9.9 I LI Jan.99 3 Elaeocarpacae Sloanea sigun (Blume) K. Schum. 10.3 6.3 Orange 1 8.8 5.9 I MT Dec.98 4 Euphorbiaceae Glochidion sphaerogynum (Mull.Arg.) Kurz 5.7 4.4 Red 9 4.9 3.4 D ST Jan.99 5 Euphorbiaceae Macaranga tanarius (L.) Mull. Arg. 5.9 5.4 Black 1 5.2 4.2 D ST Sep.98 6 Gnetaceae Gnetum montanum Markgr. 19.4 15.8 Red 1 20.7 12.7 I LI Jul.98 7 Lauraceae Cinnamomum subavenium Miq. 16.7 10.5 Black 1 14.2 8.1 I MT Feb.99 8 Magnoliaceae Michelia baillonii Finet & Gagnep. 9.8 7.4 Red 1 7.6 5.8 D TT Jul.98 9 Meliaceae Aglaia lawii (Wight) C.
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