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Authors requiring further information regarding Elsevier’s archiving and manuscript policies are encouraged to visit: http://www.elsevier.com/copyright Author's personal copy Forest Ecology and Management 261 (2011) 531–544 Contents lists available at ScienceDirect Forest Ecology and Management journal homepage: www.elsevier.com/locate/foreco Bird community assembly in Bornean industrial tree plantations: Effects of forest age and structure Alison R. Styring a,1, Roslina Ragai b,2, Joanes Unggang b,2, Robert Stuebing b,3, Peter A. Hosner c,4, Frederick H. Sheldon c,∗ a The Evergreen State College, Olympia, WA 98505, United States b Grand Perfect Sdn. Bhd., ParkCity Commerce Square, 97000 Bintulu, Sarawak, Malaysia c Museum of Natural Science, Department of Biological Sciences, Louisiana State University, Baton Rouge, LA 70803, United States article info abstract Article history: Plantations of exotic trees for industrial and agricultural purposes are burgeoning in the tropics, and Received 24 June 2010 some of them offer the opportunity to study community ecology of animals in a simplified forest set- Received in revised form 1 November 2010 ting. We examined bird community assembly in different aged groves of the industrial tree mangium Accepted 2 November 2010 (Acacia mangium) at two plantations in Malaysian Borneo: Sabah Softwoods near Tawau, Sabah, and Available online 30 November 2010 the Planted Forest Project, near Bintulu, Sarawak. Bird communities were compared among three age- groups of mangium (2-, 5-, and 7-years old) and logged native forest. Mangium rapidly developed into Key words: a secondary forest consisting of a wide diversity of understory trees and shrubs. The bird community Acacia mangium Chronosequence correspondingly increased in species richness and diversity, and we were able to relate these increases Logged forest specifically to canopy height, secondary canopy development, and shrub cover. Species of common, small Mangium bodied frugivores, nectarivores, and insectivores were diverse in older plantation groves, as were com- Sabah mon mid-sized insectivores. However, large, specialized, and normally uncommon taxa (e.g., galliforms, Sarawak pigeons, hornbills, barbets, midsized woodpeckers, muscicapine flycatchers, and wren babblers) were Succession rare or nonexistent in the plantations. Because we lacked species-specific data on foraging, nesting, and other behaviors of most groups of birds, it was difficult to explain the precise causes of seral diversifica- tion in any group except woodpeckers, which have been well studied in Southeast Asia. Thus, in future, particular emphasis needs to be placed on obtaining such data. Industrial plantations, by virtue of their simple structure, variably aged groves, and bird community richness, are good places to gather such data. © 2010 Elsevier B.V. All rights reserved. 1. Introduction tions is also burgeoning (Barlow et al., 2007; Rotenberg, 2007; Koh, 2008; Sheldon et al., 2010). However, most of these studies have “Industrial” plantations that produce fast growing trees for pulp, focused on determining which kinds of animals occur in planta- composite board, and solid wood products are burgeoning in the tions. Relatively few have taken advantage of plantation structure tropics worldwide (Cossalter and Pye-Smith, 2003; Dvorak, 2004; to study animal ecology, especially community succession. This is Evans, 2009). Because of concern that monocultures of exotic trees surprising because some types of plantations offer a natural exper- will have an adverse effect on biodiversity (Fitzherbert et al., 2008; iment in community assembly. The occurrence of different aged Sodhi et al., 2008), research on native animals in tropical planta- groves of trees at a single location allows the examination of a variety of seral stages at a single point in time and space (e.g., Atkeson and Johnson, 1979; Mitra and Sheldon, 1993; Hanowski et al., 1997; Koh, 2008). This “space-for-time” approach, and the ∗ Corresponding author at: Louisiana State University, Museum of Natural Science, resulting “chronosequence” of observations (Pickett, 1989), allows 119 Foster Hall, Baton Rouge, LA 70808, United States. Tel.: +1 225 578 2887; biologists to compare habitat and community characteristics of col- fax: +1 225 578 3075. E-mail addresses: [email protected] (A.R. Styring), [email protected] onizing species, as long as a plantation’s groves develop adequate (R. Ragai), junis [email protected] (J. Unggang), [email protected] (R. Stuebing), botanical complexity during their relatively short existence. Indus- [email protected] (P.A. Hosner), [email protected] (F.H. Sheldon). trial tree plantations in the tropics are often well suited for such 1 Tel.: +1 360 867 6837; fax: +1 360 867 5430. studies because they comprise extremely fast growing trees that 2 Tel.: +62 086 335880; fax: +62 086 335890. can develop rich secondary understories (Mitra and Sheldon, 1993). 3 Tel.: +62 541 732898; fax: +62 541 732537. 4 Tel.: +1 785 864 3657; fax: +1 785 864 5335. Agricultural plantations, such as oil palm (Elaeis guineensis), how- 0378-1127/$ – see front matter © 2010 Elsevier B.V. All rights reserved. doi:10.1016/j.foreco.2010.11.003 Author's personal copy 532 A.R. Styring et al. / Forest Ecology and Management 261 (2011) 531–544 Fig. 1. Sarawak Planted Forest Project and Sabah Softwoods plantations. Numbers refer to transect sites listed in Appendix A. ever, tend not to be as useful because their undergrowth is more We studied bird community development at two industrial tree intensively managed and, thus, unusually depauperate in botanical plantations in Malaysian Borneo. The first was Sabah Softwoods and animal community structure (Koh, 2008; Sheldon et al., 2010). Sdn. Bhd. (hereafter SS). This plantation is located ca. 50 km NNW As long as investigators recognize that seral studies in plantations of Tawau in the Tawau District of southeastern Sabah (Fig. 1) and offer a simplified view of complicated processes, such studies have is administered from Brumas Camp (4◦30N, 117◦E; ca. 300 m ele- the potential to provide insight into important ecological questions, vation). SS was established in 1974 and covers about 60,000 ha, of especially how so many species are able to coexist in tropical rain- which some 35,000–40,000 ha are planted with exotic trees includ- forest (Klopfer and MacArthur, 1961; Karr, 1971). Moreover, such ing mangium (Acacia mangium), Albizia (Paraserianthes falcataria), studies should benefit conservation efforts by providing develop- white teak (Gmelina arborea), and oil palm (E. guineensis)(Pinso ers with information on community ecology that can be translated and Vun, 2000). Several faunal studies have been conducted at SS into plantation design and management to encourage biodiversity (e.g., Duff et al., 1984; Stuebing and Gasis, 1989; Mitra and Sheldon, (Hanowski et al., 1997; Stuebing, 2007; Nasi et al., 2008). 1993; Sheldon et al., 2010), and lists of birds found in native forest at In ornithology, most efforts to understand rainforest bird com- that site have been compiled periodically since 1977 (Sheldon et al., munity succession have focused on chronosequences in natural 2001). The second plantation is the Sarawak Planted Forest Project forest (Terborgh, 1985), logged or burned forests of different ages (hereafter PFP), located ca. 30 km S of Bintulu in the Tatau District of (e.g., Lambert, 1992; Johns, 1996; Barlow and Peres, 2004; Styring central Sarawak (Fig. 1). Its administrative center is the Samarakan and Zakaria, 2004a), or forest recovering from slash-and-burn agri- Nursery (2◦56N, 113◦07E; ca. 50 m elevation). The PFP was estab- culture (e.g., Bowman et al., 1990; Blankespoor, 1991; Raman lished in the mid-1990s, when the Sarawak government set aside et al., 1998; Borges, 2007). The potential for insight from these some 500,000 ha for forest development projects. About 200,000 ha approaches is substantial because each examines change or recov- have been planted with mangium (Stuebing, 2005). Like SS, the PFP ery of native forest, and native forest is richer than exotic forest in has been the focus of several faunal studies (e.g., Stuebing et al., compositional and structural information. This is particularly true 2007; Shadbolt and Ragai, 2010), and lists of birds in the PFP have of studies of natural forest and slash-and-burn succession because been compiled continuously since January 2005 (Stuebing, 2007). chronosequences may span hundreds of years. However, working The groves we examined for this study comprised logged with native systems is difficult. There may be uncertainty about native forest and three age-groups of mangium. Our surveys were the age and sequence of seral stages, or a lack of replicate plots designed to estimate bird species richness (number of species), of the same age, or unclear borders between age groups. Research diversity (number of species adjusted for abundance of individuals in natural forest is particularly difficult because it requires a huge in each), and density (individuals/hectare) occurring in each grove investment of time to understand the terrain and birds (Terborgh, type. Because we expected bird and plant community complex- 1985; Terborgh et al., 1990). Forest recovering from human pertur- ity to be correlated (MacArthur and MacArthur, 1961; Roth, 1976; bation has the additional problem that plots may have experienced Hanowski et al., 1997; Rotenberg, 2007), we related bird occur- different forms and intensity of disturbance (Johns, 1997; Raman rence in each grove to the physical structure of the grove, including et al., 1998).