Animal Conservation (2003) 6, 299306 2003 The Zoological Society of London DOI:10.

1017/S1367943003003366 Printed in the United Kingdom

Effects of surface res on understorey insectivorous birds and terrestrial arthropods in central Brazilian Amazonia
Torbjrn Haugaasen, Jos Barlow and Carlos A. Peres
Centre for Ecology, Evolution and Conservation, School of Environmental Sciences, University of East Anglia, Norwich NR4 7TJ, UK (Received 3 October 2002; accepted 2 April 2003)

Abstract Understorey insectivorous birds are highly vulnerable to different forms of habitat disturbance in tropical forests. Here we examine the effects of an unprecedented surface re on understorey insectivores and forest litter arthropods in a terra rme forest of central Brazilian Amazonia, and compare species and guild abundance both close to and far from the clearly distinguishable fireline separating burnt and unburnt forest. All six of the most abundant insectivorous foraging guilds examined were detrimentally affected by re, with dead-leaf gleaners and professional ant-followers being the most heavily affected. While army ants were apparently absent from the burnt forest, results from pit-fall traps showed that wildres had only a limited effect on the abundance of most leaf-litter invertebrates. The reline appeared to be an abrupt barrier to many understorey insectivorous birds, and there was little evidence to suggest that primary forest species were recolonizing burnt forest up to 1 year after the res. Factors affecting the capture frequency of insectivorous birds are discussed.

INTRODUCTION The avifauna of neotropical forests is the most speciesrich in the world at both local and regional scales (Pearson, 1977; Terborgh, Fitzpatrick & Emmons, 1984). However, tropical forest ecosystems are under mounting pressure from human populations and are deteriorating rapidly because of anthropogenic disturbance. This is aggravated in vast tracts of seasonally dry forests occurring throughout the humid tropics, many of which temporarily breached their resistance to surface fires during severe droughts mediated by the 1997/98 El Nio/Southern Oscillation (ENSO) event (Goldammer, 1999; Nepstad et al., 1999). Forest understorey birds are among the most vulnerable vertebrate taxa to the increasing frequency and intensity of habitat disturbance (Wong, 1985; Johns, 1991; Thiollay, 1992, 1999; Canaday, 1996; Mason, 1996; Aleixo, 1999). Of these, avian insectivores seem to be particularly sensitive to different forms of disturbance, including selective logging (Wong, 1985; Johns, 1989; Thiollay, 1992; Mason, 1996), forest degradation and fragmentation (Bierregaard & Lovejoy, 1989; Kattan, Alvarez-Lopes & Giraldo, 1994; Stouffer & Bierregaard, 1995; Sekercioglu et al., 2002), slash-and-burn agriculture (Borges & Stouffer, 1999) and surface res (Barlow, Haugaasen & Peres, 2002).
All correspondence to: T. Haugaasen. Tel. (+44) 1603 5911299; Fax: (+44) 1603 250588; E-mail: T.Haugaasen@uea.ac.uk

This high vulnerability has been associated with a variety of life-history and ecological traits. Neotropical forest birds seem to have very specialized foraging techniques in order to nd and capture cryptic prey and have evolved a high degree of specicity in search images for prey detection (Pearson, 1977; Remsen & Parker, 1984). They also depend upon the use of narrow habitats and microhabitats, and are often sedentary and restricted to the shaded understorey of the forest interior (Orians, 1969; Remsen & Parker, 1984; Karr et al., 1990; Terborgh et al., 1990). Several hypotheses relating to these traits have been proposed to explain the disappearance of neotropical birds following disturbance (Canaday, 1996). Some studies suggest that population declines in neotropical forest understorey birds may be due to their poor dispersal abilities and their reluctance to cross open areas (Willis & Oniki, 1978; Thiollay, 1999; Develey & Stouffer, 2001; Sekercioglu et al., 2002) while others suggest that declines could be attributed to physiological constraints that prevent them from adapting to changes in understorey microhabitat structure and microclimate associated with forest disturbance (Pearson, 1977; Canaday, 1996). In our study area, these low-intensity surface res killed 36% of all trees 10 cm DBH (Diameter at Breast Height, or above the tallest buttress) 1 year after the res, dramatically altering the understorey structure and composition (Barlow et al., 2003; Haugaasen, Barlow & Peres, 2003). Although most of the understorey avifauna was affected, insectivorous species were frequently the

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T. HAUGAASEN ET AL. from the reline in burnt forest, eight were away from the fireline in unburnt forest, and four were placed perpendicular to the reline, with 12 nets in each forest type (hereafter, interface netlines). Netlines thus provided an equal sampling effort in burnt and unburnt forest. All new birds captured were rung with a plastic ring that was colour-coded for each site. At interface sites, different colour rings were used in the unburnt and burnt forest in order to track any bird movement across the fireline. Species were assigned to guilds based on diet and foraging modes following Terborgh et al. (1990). Although species in these guilds may differ in other aspects of their life histories (e.g., sociality and nesting behaviour), these guilds have been successfully used to group responses to disturbance (Barlow, 2003), and form ecologically meaningful groupings. Arthropod sampling Arthropods were collected using unbaited pit-fall traps consisting of plastic pots placed within the same 20 quarter-hectare forest plots where the understorey avifauna was sampled. At each plot, 20 traps were placed randomly along each netline 10 m off the netline transect, with no two traps closer than 10 m to one another. The traps had a diameter of 10 cm (area = 78.6 cm2) and a depth of 7 cm, and were buried so they were level with the soil surface. A 30:70 mixture of alcohol and water was added to each pot to a depth of 1 cm, thus killing and preserving any invertebrates captured. All traps were operated during the 2.5 days mist-netting period and examined after 72 hours, when all arthropods were collected, identied to order and family, and classied into two size classes (either <10 mm or >10 mm, with the exception of the Coleoptera, which were classified as either <10 mm, 1020 mm or >20 mm). All insect and arachnid specimens less than 2 mm were counted but not identied. The total biomass of arthropods collected at each site (wet weights) was measured in situ using precision Pesola scales accurate to 0.1 g, as no desiccation facilities were available. Sampling was restricted to a 5-month period (November 1998 to March 1999), and alternated between burnt and unburnt forest, in order to minimize any possible effects of seasonality. As with mist-netting, pit-fall traps measure levels of mobility and ease of capture as well as abundance (Topping & Sunderland, 1992). However, they have been shown to collect leaf litter arthropods efciently and to quantify their relative abundance (Mitchell, 1963), and are an effective way of measuring leaf-litter arthropod abundance in tropical forests. Data analysis Owing to the difference in sampling efforts between the fireline interfaces and the burnt and unburnt forest samples, bird captures were converted into number of captures per 1000 net hours. One-way ANOVAs with Tukeys post-hoc tests were used to compare guild

most vulnerable understorey birds to re disturbance and the resulting changes in forest structure (Barlow et al., 2002). We therefore consider the effects on insectivores in more detail, focusing on the six most abundant foraging and dietary guilds of understorey insectivores; the professional ant-followers (IAF), arboreal gleaning insectivores (IAG), arboreal sallying insectivores (IAS), bark-searching insectivores (IBS), dead-leaf searching insectivores (IDL) and terrestrial gleaning insectivores (ITG). Changes in the abundance of leaf-litter arthropods across the burnt/unburnt forest interface were also examined. We discuss whether changes in the abundance of insectivorous birds are best explained by changes in the post-burn leaf-litter arthropod fauna, or through changes in forest structure and foraging microhabitats associated with re disturbance. Finally we assess whether primary forest species were able to recolonize the burnt forest from adjacent areas of unburnt forest 1 year after the res, and examine how understorey insectivorous birds and leaf-litter arthropods responded to a soft edge within the forest. METHODS Study area This study was carried out between November 1998 and March 1999 in areas of both burnt and unburnt primary terra rme (non-ooded) forest of central Amazonia. The network of sampling sites was located in westernmost Par, Brazil (23408 S; 552215 W), along the Rio Mar (= upper Rio Arapiuns), a clear-water tributary of the lower Rio Tapajs. Burnt forest sites were affected by surface (or ground) res in NovemberDecember 1997, following a rainless period of at least 110 days linked to the 1997/98 ENSO event. These res scorched the forest understorey and killed a vast proportion of the saplings and small trees, as well as 3648% of the trees >10 cm in DBH (Peres, 1999; Barlow et al., 2002; Haugaasen et al., 2003). A map of the 20 quarter-hectare (10 250 m) forest plots sampled and detailed description of the study area can be found elsewhere (Barlow et al., 2002). Avian abundance and diversity The structure of the understorey avifauna in burnt and unburnt forest was examined using understorey mistnetting. Despite its limitations in sampling the forest avifauna (Terborgh, 1985; Thiollay, 1994; Remsen & Good, 1996) mist-netting was favoured as it is an effective method for deriving estimates of the relative density and ranging behaviour of birds in the tropical forest understorey (Karr, 1981). The data sampling protocol is described in detail elsewhere (Barlow et al., 2002) so only a brief summary is included here. A total of 2073 captures was obtained from 20 linear netlines operated during a period of 5 months. Each netline consisted of 24 12 m mist-nets, opened for 30 daylight hours (2.5 12-hour days), and all captures had their individual net number noted. Eight netlines were located at least 500 m away

Ecological effects of Amazonian surface res abundance in the four groupings of sampling sites (burnt forest away from netlines; unburnt forest away from netlines; burnt interfaces; and unburnt interfaces), whereas KruskallWallis tests were used to examine differences in the abundance of individual species. Data for three guilds (IAG, IBS and IDL) were log(x+1) transformed to increase normality for ANOVAs. Because of the high degree of variance in arthropod capture rates, all plots were grouped and 2 tests were used to examine differences between the numbers of arthropods caught in each group. RESULTS Avian abundance and diversity There were significant differences between the capture frequency of all six foraging guilds between the four disturbance treatments (Fig. 1). There was also a dramatic decrease in the total number of captures across the reline (Fig. 2). Species responses within individual guilds are dealt with in turn below. Fire negatively affected the four species classified as professional ant-followers (IAF, Fig. 1). Although direct observations and frequent captures away from ant-swarms suggest that the scale-backed antbird Hylophylax poecilonota and the white-chinned woodcreeper Dendrocincla merula did not behave as obligate antfollowers, their declines in burnt forest were highly significant (Table 1). The locally endemic harlequin antbird Rhegmatorhina berlepschi was the only species that was apparently extirpated from burnt forest sites far from the fireline, whereas the fivefold decline in the

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abundance of black-spotted bare-eyes Phlegopsis nigromaculata was not signicant. Arboreal gleaning insectivores (IAG) were signicantly less abundant in the burnt interface than in either burnt forest or unburnt forest plots far from the reline (Fig. 1). There were strong shifts in species abundance within this guild, and significant declines in some primary forest specialists (e.g., long-winged antwren Myrmotherula longipennis and plain-throated antwren M. hauxwelli) were compensated for by the signicant increase of other species such as the white-anked antwren M. axillaris and warbling antbird Hypocnemis cantator. Overall, half of the 22 species captured within this guild were less common in burnt than in unburnt forest. Arboreal sallying insectivores (IAS) in either burnt forest treatments were signicantly less abundant than in either group of netlines in unburnt forest. Species responses were mixed, and while the three most common species did show significant differences in abundance, others such as the royal flycatcher Onychorhynchus coronatus did not (Table 1). Of the 28 species caught in this guild, 16 declined, seven increased and five were captured in equal numbers in burnt and unburnt forest. However, all but three species were surprisingly least abundant in the burnt forest interfaces. Although there was no signicant difference between capture rates of the bark-searching insectivore guild (IBS) between burnt and unburnt forest, there was a signicant decline between the burnt and unburnt sides of interface netlines. The small guild-level difference between burnt and unburnt forest is in part a result of different responses by individual species, shown by the apparent increase of the olivaceous woodcreeper Sittasomus griseicapillus and

Table 1. Number of bird captures per 1000 net-hours in the four habitat treatments for the four most frequently caught insectivorous species in each foraging and dietary guild. UF = Unburnt forest, UFI = Unburnt forest interface, BFI = Burnt forest interface, BF = Burnt forest. Test statistics are for a KruskallWallis test, with 3 degrees of freedom. Signicant differences (at P < 0.05) are shown in bold. Tests were not performed for those species with empty cells in more than one treatment. Species Dendrocincla merula Hylophylax poecilonota Phlegopsis nigromaculata Rhegmatorhina berlepschi Myrmotherula longipennis Myrmotherula hauxwelli Hypocnemis cantator Myrmotherula axillaris Thamnomanes saturninus Schiffornis turdinus Dendrocincla fuliginosa Onychorhynchus coronatus Xiphorhynchus ocellatus Glyphorynchus spirurus Hylexetastes perrotii Sittasomus griseicapillus Myrmotherula leucophthalma Automolus ochrolaemus Philydor erythrocercus Automolus infuscatus Formicarius colma Myrmoborus myotherinus Sclerurus caudacutus Synallaxis rutilans Guild IAF IAF IAF IAF IAG IAG IAG IAG IAS IAS IAS IAS IBS IBS IBS IBS IDL IDL IDL IDL ITG ITG ITG ITG UF 176.4 65.3 19.4 15.3 90.3 47.2 11.1 9.7 81.9 66.7 20.8 19.4 30.6 51.4 15.3 4.2 27.8 11.1 6.9 1.4 25.0 5.6 18.1 0.0 UFI 50.0 19.4 8.3 16.7 11.1 13.9 16.7 8.3 33.3 16.7 33.3 25.0 44.4 5.6 22.2 2.8 13.9 0.0 13.9 2.8 13.9 5.6 5.6 0.0 BFI 8.3 0.0 5.6 2.8 0.0 0.0 13.9 5.6 0.0 0.0 2.8 2.8 11.1 0.0 2.8 13.9 0.0 0.0 0.0 0.0 2.8 2.8 0.0 0.0 BF 25.0 6.9 4.2 0.0 8.3 6.9 29.2 36.1 11.1 12.5 13.9 16.7 34.7 13.9 9.7 23.6 1.4 2.8 0.0 0.0 1.4 13.9 0.0 13.9 2 14.83 12.93 3.93 8.47 14.50 14.39 5.33 7.81 18.78 13.33 10.88 4.96 7.25 10.03 8.71 6.89 15.41 9.55 3.08 P 0.002 0.005 0.269 0.037 0.002 0.002 0.149 0.050 0.000 0.004 0.012 0.174 0.064 0.018 0.033 0.076 0.001 0.023 0.379

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0.04 0.03 0.02

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0.002 0.000 UF UFI BFI BF

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Forest sites
Fig. 1. Mean number of captures per net-hour in unburnt forest (UF), unburnt forest interface (UFI), burnt forest (BF) and burnt forest interface (BFI) for the following insectivorous guilds: IAF = professional ant-followers; IAG = arboreal gleaning insectivores; IAS = arboreal sallying insectivores; IBS = bark-searching insectivores; IDL = dead-leaf searching insectivores; ITG = terrestrial gleaning insectivores. One-way ANOVA: IAF F3,20 = 6.8, P = 0.002; IAG F3,20 = 5.3, P = 0.008; IAS F3,20 = 23.2, P < 0.001; IBS F3,20 = 3.7, P = 0.027; IDL F3,20 = 14.7, P < 0.001; ITG F3,20 = 7.0, P = 0.002 (data for IDL, IBS and IAG were log10 (x+1) transformed to increase normality). Numbers represent signicant subsets from Tukeys HSD test.

the disappearance of the wedge-billed woodcreeper Glyphorhynchus spirurus in burnt forest. The signicant difference between the burnt and unburnt interface netlines is likely to have been compounded by the responses of ocellated woodcreeper Xiphorhynchus ocellatus and the red-billed woodcreeper Hylexetastes perrotii which were most abundant in unburnt interfaces, and least abundant in burnt interfaces. Dead-leaf searching insectivores (IDL) appeared to be the most heavily affected, virtually disappearing in burnt forest. All species of dead-leaf gleaners exhibited lower abundances in the burnt forest, and the entire guild was

absent from the four burnt interface sites (Table 1). While unburnt forest netlines adjacent to or >500 m from the reline differed signicantly from burnt forest samples, they did not differ from one another (Fig. 1). Terrestrial gleaning insectivores (ITG) were significantly less abundant only in the burnt forest interface (Fig. 1), but nine of the 11 species captured were less frequently recorded in burnt forest compared to unburnt forest samples. While most species in this guild were too rare to permit statistical tests, the numeric decline in the rufous-capped antthrush Formicarius colma was significant (Table 1). Other species also appeared to be

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Table 2. Total numbers of arthropod captures in unburnt forest (UF), burnt forest (BF), unburnt forest interface (UFI) and burnt forest interface (BFI). Taxonomic group Coleoptera Blattaria Arachnidae Larvae Orthoptera Diptera Formicidae Other All taxa <2mm UF 783 34 79 19 77 51 170 50 180 BF 406 18 107 14 68 32 150 32 428 2 119.5 4.9 4.2 0.8 0.6 4.3 1.3 4.0 101.2 Sig *** * * * *** UFI 203 24 34 6 44 14 137 17 70 BFI 108 12 39 5 39 18 20 17 95 2 29.0 4.0 0.3 0.1 0.3 0.5 87.2 0.0 3.8 Sig *** *

***

*P < 0.05 **P < 0.01 ***P < 0.001

affected by re disturbance; for instance, the three species of Leaftossers (Sclerrurus spp.) were only captured in unburnt forest, as was the black-bellied gnateater Conopophaga melanogaster. However the decline of these relatively rare species was offset by the increased abundance of ruddy spinetails Synallaxis rutilans and black-faced antbirds Myrmoborus myotherinus in burnt forest (Table 1), accounting for the lack of significant changes in this guild between burnt and unburnt forest (Fig. 1). Understorey arthropods A total of 3600 invertebrates was collected, including a significantly different number of individuals in unburnt (1992) and burnt forest (1608; 2 = 40.96, d.f. = 1, P = <0.001). Total arthropod biomass obtained in unburnt forest (340.4 g) was also signicantly greater than that in burnt forest (197.4 g; paired-t = 2.34, d.f. = 11, P = 0.027). Aggregate pit-fall data from all 20 quarter-hectare forest plots indicate that beetles (Coleoptera) comprised the dominant invertebrate group captured (41.7%), followed by ants (Formicidae, Hymenoptera: 13.3%), spiders (Araneae: 6.9%) crickets, grasshoppers and katydids (Orthoptera: 6.1%), ies (Diptera: 3.2%) and cockroaches (Blattaria: 2.4%). The traps also captured small numbers of several other invertebrate taxa including termites (Isoptera), wasps (Hymenoptera), adult butterflies and moths (Lepidoptera), earwigs (Dermaptera), mites (Acarina) and various lepidopteran, dipteran and coleopteran larvae. In aggregate, however, these groups comprised only 3.2% of all arthropods sampled. Unidentified specimens smaller than 2 mm constituted 21.5% of the individuals counted. The six most abundant orders thus made up more than three-quarters of all invertebrates collected. Although most insect groups were less numerous in burnt forest, only four showed signicantly lower capture rates in burnt forest (Table 2). Numbers of Coleoptera and Blattaria were signicantly lower in both burnt interfaces and non-interface plots than in unburnt plots. Dipterans, however, were only signicantly less abundant in burnt forest plots farther than 500 m from the reline (Table 2). Ants only declined significantly in burnt forest close to the interface, although this was heavily inuenced by the

occurrence of a large army-ant (Eciton sp.) swarm that raided one of the unburnt interface plots. Six large armyant swarms were observed within unburnt forest during mist-netting, whereas no swarms were ever encountered within burnt forest during the 5-month duration of this study. The only arthropod group increasing signicantly in burnt forest were the arachnids, although their increase was only significant between burnt and unburnt forest away from the interfaces (Table 2). The numbers of unidentied insects <2mm also increased in burnt forest, but only away from the interfaces. DISCUSSION Avian insectivores typical of the shaded understorey of undisturbed primary forests were highly vulnerable to surface wildres, a pattern consistent with studies on other types of tropical forest disturbance (e.g., Wong, 1985; Bierregaard & Lovejoy, 1989; Johns, 1989; Thiollay, 1992; Stouffer & Bierregaard, 1995; Mason, 1996). In order to assess the possible causes of this vulnerability we relate our results to changes in the prey density, foraging substrate and habitat structure. We also examine the results in light of their relevance for recolonization from unburnt forest which could operate as source areas following re disturbance. Post-burn changes in leaf-litter arthropods In contrast to ndings following logging and re disturbance in dry tropical forests in Bolivia (Fredericksen & Fredericksen, 2002), the total number of individual invertebrates was generally higher in unburnt forest. However, there were signicant differences in the abundance of some key classes of prey items for insectivorous birds such as cockroaches and spiders, both of which increased signicantly in burnt forest. This also contrasts with a comparison of avian diets and invertebrate communities between small and large neotropical forest fragments (Sekercioglu et al., 2002). However, the wide range of resource dimensions and species richness of ecologically divergent insectivorous birds in neotropical forests make it very difcult to relate arthropod abundance to the availability of prey targeted by most insectivorous species. This is even the case for the terrestrial gleaning

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T. HAUGAASEN ET AL. Recolonization and edge effects If recolonization from unburnt forest had been occurring at the time of this study, then it should have been most apparent in those areas nearest the unburnt forest source populations (the burnt interface samples). However, this trend was not observed, either at small scales within the interface plots (Fig. 2) or in comparisons between bird abundance within burnt and unburnt plots far from the reline with the abundance levels at the interface (Fig. 1). In fact, the immediate reduction in capture rates across the reline suggests that this operated as an effective barrier to many species. As an unwillingness to enter open habitats is a distinct feature of many primary forest understorey insectivores (Stouffer & Bierregaard, 1995; Borges & Stouffer, 1999; Develey & Stouffer; 2001), it seems likely that the increased canopy openness following fire (Barlow et al., 2002; Haugaasen et al., 2003) was sufficient to limit the movement and dispersal of most insectivorous understorey birds associated with primary forest. Although edge effects on tree mortality can be influenced by the surrounding vegetation structure (Mesquita, Delamonica & Laurance, 1999), it appears that even relatively soft edges such as these can have marked effects and act as effective barriers for understorey birds. Furthermore, many forest species were less abundant in the burnt forest interface than in the burnt forest plots >500 m from the fireline (Table 1), a trend opposite to what would be expected if recolonization from primary forest had been occurring. As there were no significant differences between the forest structure at the burnt forest interfaces and the burnt forest >500 m from the reline (Haugaasen, 2000), the most parsimonious explanation for this relates to local shifts in home-range use, whereby birds with territories close to or overlapping the reline emigrated into adjacent areas of unburnt forest in the aftermath of the res.

insectivores, as changes in pit-fall captures in traps may not necessarily reect availability for these birds. For example, while the declines in many species in this guild may be related to changes in arthropod abundance in the leaf-litter, the increase in the abundance of some members of this guild (e.g., Myrmoborus myotherinus, see Table 1) suggests that their post-burn abundance was determined by their ability to forage in burnt forest rather than the prey base (cf. Sekercioglu et al., 2002). We are therefore unable to draw any unambiguous conclusions about food availability for insectivores. However, the results suggest that there may have been several shifts in the availability of the food supply and that the absence of some key invertebrate taxa targeted as food items (and army-ant swarms which dislodge canopy arthropod prey into the understorey) may have strongly inuenced the post-burn avifauna. Despite these difficulties, it is tempting to draw a comparison between the decline of the ant-following insectivores and the apparent absence of army-ant swarms from burnt forest. Here the effects of surface wildfires contrast markedly with those of other types of forest disturbance such as logging and fragmentation where army-ants appear to persist (Bierregaard & Lovejoy, 1989; Mason, 1996). In this respect, the short-term consequences of surface fires appear to be similar to those found following the isolation of small forest fragments, where the decline of wide-ranging ant swarms is also associated with the local extinction of professional army-ant followers (Willis, 1974; Bierregaard & Lovejoy, 1989). Structural changes It may be that it is the structural characteristics of burnt forest that dene which species were resilient to the res. Karr & Freemark (1983) observed that individual species showed some preference among the range of moisture conditions present in primary forest, and even altered their primary habitat association over time as they tracked suitable microclimatic conditions. Their hypothesis was that physiological factors may control the ability of these species to utilize locally abundant resources at specific sites. This is supported by Barlow et al. (2002), who found that changes in canopy openness and understorey vegetation cover were strongly correlated with changes in bird community structure following surface res. However, changes in habitat structure could also cause some birds to adjust their foraging heights and capture rates according to different environmental conditions (Pearson, 1977; Remsen & Good, 1996). Although changes in foraging height remain a possibility, the loss of most midstorey foliage because of the high sapling mortality (Haugaasen et al., 2003) makes it highly unlikely that specialised understorey species had shifted their foraging height to higher forest strata, while most of the observed bird species were foraging on regenerating plants, well within the capture height of our 2.5 m tall mist-nets. Furthermore, most conspicuous birds were most frequently heard or seen in the netlines where they were also most frequently captured (T. Haugaasen & J. Barlow, pers. obs.).

Mean ( SE) no. of captures

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30 20 10 0

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10 12 14 16 18 20 22 24

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Fig. 2. Mean number of captures (six insectivorous guilds combined) at each net placed in all interface netlines perpendicular to the fireline, where half of the nets were in unburnt forest and half were in burnt forest.

Ecological effects of Amazonian surface res CONCLUSION While these understorey res had dramatic effects on tree mortality and forest structure (Peres, 1999; Barlow et al., 2003; Haugaasen et al., 2003), they appear to have had only a limited effect on the numeric abundance of leaflitter invertebrates preyed upon by insectivorous birds. Nevertheless, all of the insectivorous guilds examined here declined in abundance following disturbance from low-intensity surface res. Furthermore, these differences were often particularly dramatic across the reline, and even 1 year after the re, there was little or no evidence of recolonization from adjacent unburnt forest of species sensitive to re disturbance into burnt forest. In fact, the spatial distribution pattern of many species suggests that they may have emigrated from burnt forest into unburnt forest. These results support hypotheses that foraging substrates and microhabitat conditions are important determinants of the community structure of understorey insectivores, and highlight the extreme susceptibility of primary forest species to wildfires. This has important implications for wildlife conservation in neotropical forests (Peres, Barlow & Haugaasen, 2003) which are becoming increasingly fire-prone because of logging, deforestation and climate change including more frequent extreme El Nio events, and suggests that wildre hazards must be taken into account when planning forest reserves under any degree of protection. This includes the expanding matrix of National Forests earmarked for lowimpact logging in seasonally dry regions of Amazonia (Verissimo, Cochrane & Sauza, 2002), which are often proposed as the best compromise for biodiversity conservation in the long term. Acknowledgements This study was funded by the Center for Applied Biodiversity Science (CABS) of Conservation International and the Josephine Bay and Michael Paul Foundation. Jos Barlows eldwork was funded by a NERC PhD studentship at the University of East Anglia. We are especially thankful to the villagers and political leadership of the Reserva Extrativista do Tapajs-Arapiuns for allowing us to conduct this study, and to the local assistance of, among others, Nan, Arnei and Torzinho. Rionaldo and Edith de Santos provided enormous logistical assistance during all stages of the project. REFERENCES
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