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The Ecological Basis of Sensitivity of Brown Treecreepers to Habitat Fragmentation: a Preliminary Assessment

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The Ecological Basis of Sensitivity of Brown Treecreepers to Habitat Fragmentation: a Preliminary Assessment Biological Conservation 90 (1999) 13±20 The ecological basis of sensitivity of brown treecreepers to habitat fragmentation: a preliminary assessment Jerey R. Walters a,*, Hugh A. Ford b, Caren B. Cooper c aDepartment of Zoology, North Carolina State University, Raleigh, NC 27695-7617, USA bDepartment of Zoology, University of New England, Armidale, NSW 2351, Australia cDepartment of Biology, Virginia Polytechnic Institute and State University, Blacksburg, VA 24061-0406, USA Received 4 April 1998; received in revised form 22 October 1998; accepted 5 January 1999 Abstract We attempted to identify the mechanisms responsible for adverse eects of habitat fragmentation on brown treecreepers (Cli- macteris picumnus) inhabiting eucalyptus woodland in northeastern New South Wales, Australia by comparing demography and foraging ecology of birds in highly fragmented and relatively unfragmented landscapes. In particular, we investigated three possi- bilities, disrupted dispersal due to patch isolation, reduced fecundity due to elevated nest predation, and reduced food availability due to habitat degradation. Nesting success was high in both highly fragmented and less fragmented habitat. Of ®rst nests, 88% were successful, and 60% of successful groups attempted a second brood. However, there were many more groups in the more fragmented habitat than in the less fragmented habitat that lacked a female for most or all of the breeding season, and thus did not attempt nesting (64% vs 13%). In both the more fragmented and the less fragmented habitat, both males and females spent about 70% of their time foraging and 65% of their foraging time on the ground. We reject reduced fecundity in fragmented habitat as an explanation of adverse eects of habitat fragmentation on brown treecreepers. Thus sensitivity to habitat fragmentation has a dif- ferent basis for this species in this landscape than that suggested for Nearctic-Neotropical migrants in eastern North America. We also reject the possibility of reduced food availability in fragmented habitat. Our data support disrupted dispersal as a likely explanation for the decline of brown treecreepers in fragmented habitat. However, we can not rule out forms of habitat degradation other than reduced food availability. # 1999 Elsevier Science Ltd. All rights reserved. Keywords: Habitat fragmentation; Eucalyptus woodland; Demography; Foraging; Dispersal 1. Introduction are sensitive to fragmentation whereas others are not. For example, in eastern North America Nearctic-Neo- Adverse eects of habitat fragmentation on plant and tropical migrants are more sensitive than permanent animal species are one of the most pervasive conserva- residents (Whitcomb et al., 1981; Lynch and Whigham, tion problems (Temple and Wilcox, 1986). A species is 1984; Askins et al., 1990; Holmes and Sherry, 1992), considered sensitive to fragmentation if density or ®t- while in Amazonian forests sensitivity diers among ness of individuals within remaining patches of its foraging guilds (Lovejoy et al., 1984; Kattan et al., 1994; habitat is reduced as fragmentation of its habitat in the Stouer and Bierregaard, 1995). landscape increases (Walters, 1998). As its habitat It is dicult to detect the mechanisms by which habi- becomes reduced in extent and increasingly patchy tat fragmentation aects birds. It is necessary to deter- through the fragmentation process, such a species suf- mine not only if the decline of a species in a fragmented fers losses out of proportion to reduction of its habitat landscape exceeds that expected from habitat loss alone, because of its poor performance in remaining patches. but also whether excessive losses are due to the habitat Within any avian community, it seems that some species degradation that often accompanies fragmentation, rather than fragmentation itself. These possibilities have yet to be distinguished as explanations of population * Corresponding author at present address: Department of Biology, Virginia Polytechnic Institute and State University, Blacksburg, VA declines of avian species as their habitats are reduced and 24061-0406, USA. Tel.: +1-540-231-3847; fax: +1-540-231-9307; fragmented. We oer three speci®c hypotheses to explain e-mail: [email protected] the declines of many woodland birds in Australia 0006-3207/99/$ - see front matter # 1999 Elsevier Science Ltd. All rights reserved. PII: S0006-3207(99)00016-6 14 J.R. Walters et al. / Biological Conservation 90 (1999) 13±20 (Robinson and Traill, 1996) that illustrate the necessary leucophaea, remains common in this landscape and distinctions. First, the declines may be no more than apparently is unaected by fragmentation, whereas expected from (1) habitat loss in accordance with spe- another, the red-browed treecreeper Climacteris ery- cies±area relationships. Second, (2) habitat fragmenta- throps, appears to be highly sensitive, having dis- tion may be causing disproportionate declines as a appeared from all but the largest remaining woodland result of (a) disrupted dispersal due to increased isola- patches (Barrett et al., 1994). The rufous treecreeper C. tion of remnant habitat patches by alien habitat; (b) rufus, an allopatric form closely related and ecologically adverse eects of increased amounts of edge mediated similar to the brown treecreeper, has been similarly by changes in microclimate, microhabitat or food; or (c) aected by fragmentation in the Western Australian elevated levels of nest predation and/or nest parasitism wheatbelt (Saunders, 1989; Saunders and Ingram, due to increased amounts of edge. Third, (3) habitat 1995). The presence of these comparisons facilitates degradation may be causing disproportionate declines identifying mechanisms that might be responsible for through (a) increases in aggressively competitive species adverse eects of fragmentation. that exclude many other bird species; (b) increases in In this paper we focus on four of the possibilities nest predators and/or nest parasites mediated by chan- suggested above. (1) The decline of brown treecreepers ges in landscape composition; (c) introduced predators may be due to disrupted dispersal behavior resulting that reduce reproductive success or adult survival; (d) from habitat fragmentation, and thus population direct mortality resulting from collisions with vehicles performance may be related to degree of isolation of and power lines; (e) reduced food availability due to remnant patches (disrupted dispersal hypothesis, 2a eects of grazing; or (f) reduced availability of nesting above). The decline might be due to elevated nest pre- sites due to removal of mature trees for timber or ®re- dation associated with (2) increased edge or (3) wood. Adverse eects of fragmentation on birds inha- increased numbers of predators in fragmented land- biting forests of eastern North America, which involve scapes, as occurs in eastern North America (Robinson interactions between landscape structure and rates of et al., 1995; Faaborg et al., 1998) (nest predation nest predation and nest parasitism, most likely are hypotheses, 2c and 3b above, respectively). (4) The explained by hypothesis (2c) and/or hypothesis (3b) decline might result from reduced food availability due (Robinson et al., 1995; Faaborg et al., 1998). to habitat degradation, and thus population perfor- Studies of declining species are needed to distinguish mance may be related to the extent of degradation of between these and other possibilities, to identify the remnant woodland patches (food availability hypoth- range of mechanisms responsible for adverse eects of esis, 3e above). The disrupted dispersal hypothesis is an fragmentation, and to determine how these mechanisms especially intriguing possibility because the brown tree- vary among systems. The objective of this study is to creeper is a cooperative breeder. Generally in coopera- identify the mechanisms responsible for the decline of tive breeders fewer individuals disperse long distances the brown treecreeper Climacteris picumnus in frag- compared to other species (Emlen, 1991). In brown mented eucalypt woodland in the New England Table- treecreepers, many males, after spending a period as lands of northeastern New South Wales, Australia. The non-breeding helpers on their natal territory, acquire woodland in this region has been lost and fragmented breeding sites by claiming a portion of the natal terri- by clearing for livestock grazing. Remaining habitat has tory, a process known as budding (Noske, 1982, 1991). been degraded by eucalypt dieback (Jones et al., 1990; Thus, only a fraction of the males disperse beyond the Landsberg et al., 1990; Lowman and Heatwole, 1992). immediate vicinity of their natal territory. Females are The result is perhaps best described as a variegated not so sedentary, and may in fact have to disperse landscape (McIntyre and Barrett, 1992), consisting of beyond their natal neighborhood in order to encounter some open grassland and good woodland patches (i.e. unrelated males. healthy trees at normal densities), and much woodland We attempt to distinguish between the four hypoth- in various stages of degradation (i.e., some unhealthy eses outlined above by comparing pairing success, trees, reduced tree density) (Barrett et al., 1994). Barrett breeding success and time spent foraging between an et al. (1994) identi®ed a number of species, including the area of relatively unfragmented woodland in which brown treecreeper, that were once common but have brown treecreepers remain common and a highly frag- become uncommon
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