Journal of Tropical Ecology (2002) 18:311?325. With 4 ?gures. Copyright ? 2002 Cambridge University Press DOI:10.1017/S0266467402002237 Printed in the United Kingdom Fire as a large-scale edge effect in Amazonian forests MARK A. COCHRANE* and WILLIAM F. LAURANCE??1 *Basic Science and Remote Sensing Initiative, Michigan State University, East Lansing, MI 48823, USA (Email: cochrane@bsrsi.msu.edu) ?Smithsonian Tropical Research Institute, Apartado 2072, Balboa, Republic of Panama? ?Biological Dynamics of Forest Fragments Project, National Institute for Amazonian Research (INPA), C.P. 478, Manaus, AM 69011-970, Brazil (Accepted 14th July 2001) ABSTRACT. Amazonian forests are being rapidly cleared, and the remaining forest fragments appear unusually vulnerable to ?re. This occurs because forest remnants have dry, ?re-prone edges, are juxtaposed with frequently burned pas- tures, and are often degraded by selective logging, which increases forest desicca- tion and fuel loading. Here we demonstrate that in eastern Amazonia, ?res are operating as a large-scale edge effect in the sense that most ?res originate outside fragments and penetrate considerable distances into forest interiors. Multi- temporal analyses of satellite imagery from two frontier areas reveal that ?re frequency over 12?14-y periods was substantially elevated within at least 2400 m of forest margins. Application of these data with a mathematical core-area model suggests that even large forest remnants (up to several hundred thousand ha in area) could be vulnerable to edge-related ?res. The synergistic interactions of forest fragmentation, logging and human-ignited ?res pose critical threats to Ama- zonian forests, particularly in more seasonal areas of the basin. KEY WORDS: Amazon, Brazil, conservation, carbon emissions, deforestation, drought, ?re, habitat fragmentation, logging, rain forest, remote sensing, tropical forest INTRODUCTION The Amazon basin contains nearly 60% of the world?s remaining tropical rain forest, and plays critical roles in maintaining biodiversity, regional hydrology and climate, and carbon storage (Fearnside 1999). It also has the world?s high- est absolute rate of forest destruction, averaging roughly 3?4 million ha y-1 (INPE 2001, Whitmore 1997). 1 Corresponding author. Smithsonian Tropical Research Institute, Apartado 2072, Balboa, Republic of Panama?. Email: laurancew@tivoli.si.edu 311 MARK A . COCHRANE AND WILL IAM F . LAURANCE312 The rapid pace of Amazon deforestation is causing widespread habitat frag- mentation. By 1988, the area of forest in the Brazilian Amazon that was frag- mented (< 100 km2 in area) or vulnerable to edge effects (< 1 km from clearings) was over 150% larger than the total area deforested (Skole & Tucker 1993). Because over 15% of the Brazilian Amazon has now been cleared (INPE 2001), the total area altered by deforestation, fragmentation and edge effects may comprise a third or more of the region today (Laurance 1998). Habitat fragmentation affects the ecology of Amazonian forests in many ways, such as altering the diversity and composition of fragment biotas (Bierregaard et al. 1992, Lovejoy et al. 1986) and changing ecological processes such as pollination and nutrient cycling (Didham et al. 1996, Klein 1989). Recent evidence indicates that fragmentation also alters rain-forest dynamics, causing markedly elevated rates of tree mortality, damage and canopy-gap formation (Laurance et al. 1998a), apparently as a result of microclimatic changes (Kapos 1989), increased wind turbulence and proliferating lianas near fragment edges (Laurance et al. 2001b). These changes lead to a substantial loss of living biomass in fragments that may be a signi?cant source of green- house gas emissions (Laurance et al. 1997, 1998b). In addition to fragmentation, Amazonian forests are being degraded by vari- ous other activities, such as human-ignited ?res, legal and illegal logging, gold mining and over-hunting (Fearnside 1990, Laurance et al. 2001a). These addi- tional threats may interact additively or synergistically with fragmentation. An increasing body of evidence, for example, reveals that fragmentation increases the vulnerability of Amazonian forests to ?re (Cochrane & Schulze 1999, Cochrane et al. 1999, Gascon et al. 2000, Kauffman & Uhl 1990, Nepstad et al. 1999). This occurs because fragments have relatively dry, ?re-prone edges (Kapos 1989), are juxtaposed with frequently burned pastures and regrowth forests (Nepstad et al. 1996, Uhl & Kauffman 1990), and are often degraded by selective logging, which increases forest desiccation and fuel loading (Holdsworth & Uhl 1997, Uhl & Kauffman 1990). While it is apparent that fragmented forests are vulnerable to ?res, the spatial scale of this phenomenon has not yet been quanti?ed. Most ?res origin- ate in surrounding pastures or regrowth and then burn into fragment interiors, and thus appear to be operating as a kind of edge effect. To date, edge effects in Amazonian forests, such as alterations in microclimate, forest dynamics and faunal communities, have been shown to penetrate from 10?400 m into frag- ment interiors (Bierregaard et al. 1992, Laurance et al., in press, Lovejoy et al. 1986). Preliminary evidence, however, suggests that some edge-related changes in tropical forests could penetrate much further than this, perhaps as far as several km into fragment interiors (Curran et al. 1999, Laurance 2000). Here we provide evidence that, in two relatively seasonal landscapes of the eastern Amazon, ?re is operating as such a large-scale edge effect. Using a simple mathematical model, we demonstrate that even large forest fragments Fire and forest fragmentation 313 and isolated nature reserves could be vulnerable to edge-related ?res. We argue that, if fragmented, large expanses of Amazonian forests are likely to be destroyed or severely degraded by ?re. METHODS Dynamics of rain-forest ?res Natural ?res occur only rarely in undisturbed tropical rain forests (Goldammer 1990). However, colonists and ranchers use ?res both to clear forests and maintain pastures, which often burn into adjoining forest blocks. These surface ?res are deceptively unimpressive, creeping along the ground as a thin ribbon of ?ames burning through the leaf litter. Except for areas with unusual fuel structure, the ?res reach only 10 cm in height (Cochrane et al. 1999). Even during extreme droughts, rain forests maintain high humidity that makes combustion dif?cult. Many surface ?res burn during the day, only to be extinguished when relative humidity increases in the evening. Such ?res, how- ever, can smoulder in old treefalls and standing dead trees for up to several weeks, until conditions are right for the ?re to continue propagating through the leaf litter. Surface ?res may cover as little as 150 m distance in a day (Cochrane & Schulze 1998) but are deadly to trees, because the slow propaga- tion results in long residence times of the ?ames at the base of encountered trees. Most rain-forest trees have thin bark (typically < 1 cm thick; Uhl & Kauffman 1990) and the heat of even small ?res girdles many trees, killing around 40% of all standing stems ( 10 cm diameter at breast height (dbh); Cochrane & Schulze 1999). An initial surface ?re kills mostly smaller trees (< 40 cm dbh), with trees continuing to die for 2 y or more. The forest canopy becomes fragmented and the quantity of dead fuels increases as dead leaves and trees begin to fall. Soon, the forest is far more prone to subsequent ?res, because the diminished canopy allows rapid drying and the dying trees provide large amounts of combustible fuel. Burned forests commonly adjoin ?re-maintained pastures and agricultural lands. While initial rain-forest ?res may require an extensive drought, sub- sequent ?res can occur after just a few weeks without rain (Cochrane & Schulze 1999). Forests that burn a second time fare much worse because the ?res are far more intensive, overwhelming the defences of even larger, thicker-barked trees. Typically, second ?res kill 40% of the standing trees in all size classes, comprising around 40% of the standing biomass (Cochrane et al. 1999). Subsequent ?res are even more likely and severe. During the ?rst several ?res, more fuels are created than destroyed, and a positive feedback results in which each ?re becomes more probable and intensive. This process can eradic- ate trees from an area and result in extensive grass invasion. In the absence of seed sources for grassland or open-woodland species, anthropogenic savanna MARK A . COCHRANE AND WILL IAM F . LAURANCE314 (sometimes with certain palm species) will result. For regions with a pro- nounced dry season, this conversion of rain forest to savanna is likely to be irreversible (Cochrane et al. 1999). Study areas The two study areas, Taila?ndia (2470 km2) and Paragominas (1280 km2), are in the eastern Brazilian Amazon and have extensively fragmented forest cover. Both areas have similar evergreen terra-?rme forests and pronounced dry sea- sons, with mean annual rainfall ranging between 1500?2000 mm (Cochrane et al. 1999, Kauffman et al. 1995). The terrain in both areas is quite ?at and uniform, but is dissected by a number of streams and small rivers. Taila?ndia is a new frontier where large-scale deforestation began in the early 1990s when a government-sponsored colonization project was initiated. As is typical, a ??sh-bone? pattern of deforestation quickly developed as colonists situated along parallel roads employed slash-and-burn farming (Figure 1). By 1997, 44% of the area?s forests had been destroyed. Paragominas is an older frontier where logging and cattle ranching predominate (Figure 1). Large-scale clearing began in the early 1960s with construction of the nearby Bele?m?Brasi- lia Highway. By 1995, 64% of the area?s forests had been destroyed. Further information on these study areas is provided in Cochrane (2000). Core-area model We used a mathematical ?core-area model? to predict the areal extent of edge-related ?res in forest fragments of varying sizes and shapes (Laurance & Yensen 1991). This model provides accurate (> 99%) predictions of the size of unaffected core-area for any fragment based on three parameters: fragment area, a fragment-shape index (SI), and the average distance (d) to which edge effects penetrate into fragment interiors. SI is derived by dividing the frag- ment?s perimeter-length by that of an equal-sized circle, and varies from 1.0 for perfect circles to 8 or higher for very irregularly shaped fragments. SI is calculated as SI = P/(2[TA0.5]), where P is the perimeter-length of the fragment and TA is fragment area. The core area (CA) is given by CA = TA ? affected area (AA), where AA = 3.55 d SI (TA/104)0.5. AA is overestimated for more- circular fragments and is therefore adjusted downward by AAadj = (0.265 AA)/ SI0.5 (see Laurance & Yensen 1991 for a detailed explanation of the model). For both of our study areas, the total area and perimeter-length of all forest fragments of at least 0.1 ha were determined using georeferenced imagery from the Landsat Thematic Mapper (TM) satellite, at a 30-m spatial scale. Calculations were conducted using ArcInfo 8.0.1. Fire rotation-times The incidence of subcanopy-surface ?res in the two study areas was assessed for periods of 12?14 y, from 1984 to the mid-late 1990s, using multi-temporal analyses of Landsat TM imagery, augmented with extensive ground-truthing Fire and forest fragmentation 315 Figure 1. Map showing location of study areas. Below, portions of each study area in eastern Amazonia, showing the complex ??sh-bone? pattern of fragmentation arising from a forest-colonization project (Taila?ndia) and the somewhat less irregular fragmentation pattern caused by cattle ranching (Paragominas). Black areas are forest and grey areas are deforested. Each scene shows an area of about 600 km2. MARK A . COCHRANE AND WILL IAM F . LAURANCE316 and interviews of local residents (Cochrane et al. 1999). As is typical in the eastern Amazon, our study areas were affected by 2?3 droughts or rainfall de?cits associated with periodic El Nin?o events (occurring in 1986 and 1991 at both sites, and in 1997 at Taila?ndia), during which forest burning increased markedly. A sub-pixel linear spectral mixture-modelling technique (Cochrane & Souza 1998) was used to detect and classify forests that had been impacted by subcan- opy surface ?res in the Taila?ndia (1984, 1991, 1993, 1995, 1997) and Paragom- inas (1984, 1991, 1993, 1995) study areas. Speci?cally, ?re-induced tree mortal- ity opens the forest canopy, and even though the resulting gaps may be small they are suf?cient to allow sunlight from more of the forest ?oor and woody vegetation to be re?ected. These non-photosynthetic vegetation (NPV) re?ections are a small fraction of the whole scene but the spectral signature of these substances is signi?cantly higher in ?re-damaged forests than in undamaged forests. Separation of satellite-imagery information into its frac- tional components (e.g. vegetation, shade, NPV) allows ?re-damaged forests to be clearly distinguished for up to 2 y after a ?re (in an earlier study, ground- truthing revealed that 93% of forests burned < 1 y previously and 71% of areas burned 1?2 y previously were correctly classi?ed; Cochrane et al. 1999). A ?ltering technique was used to reduce the likelihood that logged forests could be misclassi?ed as burned forests (Cochrane & Souza 1998). In both study areas, ?re rotation-times were calculated as a function of dis- tance from forest edge (the ?re rotation is the average number of years required to burn an area under consideration, with the understanding that some areas may not burn while others may burn more than once during a cycle; Van Wagner 1978). For each year of the study, all forest was mapped using classi?ed images at a spatial scale of 30-m pixels. The area of forest in each edge-distance category (e.g. 0?30 m, 30?60 m, 60?90 m, etc.) was then calculated, using ArcInfo 8.0.1. Using the mixture-modelling technique described above, the distribution of surface ?res was also mapped, and the proportion of pixels that burned in each edge-distance category was calculated. Data for all years of the study were then combined to yield an average propor- tion of burned pixels for each edge-distance category. The ?re rotation-time was simply the inverse of this proportion (for example, if an average of 5% of all pixels at 0?30 m from the edge burned each year, then the ?re rotation-time would be 1/0.05 = 20 y). The distance (d) to which ?re rotation-times near forest edges deviated from those in forest interiors was estimated as follows: the 95% data distribution (mean ? 1.96 standard deviations) was calculated for ?re rotation-times in forest interiors (de?ned by visual inspection as being over 2.5 km from the nearest edge); and the point at which the observed curve of ?re rotation-times fell below this range was de?ned as d. The 95% data distribution is an appropri- ate measure of variability because ?re rotation-times in forest interiors (at Fire and forest fragmentation 317 Taila?ndia only; see below) did not deviate signi?cantly from normality (Wilk? Shapiro test, P > 0.50). RESULTS Fragment characteristics We identi?ed a total of 722 forest fragments in our two study areas, with individual fragments ranging from 0.1 ha to over 57 000 ha in area (Table 1). Small (< 100 ha) forest patches were abundant, accounting for 91?93% of all fragments in each study area, but supported only a small proportion (< 4.5%) of the remaining forest cover at each site. When shape index (SI) values were plotted against fragment area, two trends emerged. First, there was a positive relationship between SI and area (Figure 2), because larger fragments tended to be more irregularly shaped than smaller fragments (rs > 0.74, P < 0.00001 in both areas; Spearman rank correlations). This pattern resulted in part from fractal effects (because irregu- larities in the margins of small fragments tended to be smoothed over using 30-m pixels, thereby causing SI values of smaller fragments to be underestim- ated relative to those of larger fragments; cf. Li & Reynolds 1994). It does, nevertheless, re?ect a tendency for most large fragments to be very irregularly shaped at a spatial scale that is relevant to edge effects in this system. Second, fragments in Taila?ndia were often more irregularly shaped than those in Para- gominas (Table 1, Figure 2), re?ecting the complex patterns of forest frag- mentation caused by government-sponsored colonization projects in the Amazon. Core-area model For Taila?ndia, ?re rotation-times declined sharply near edges, and were found to deviate from those in forest interiors at a distance of up to 2.35 km from the forest edge (Figure 3). The core-area model for Taila?ndia was gener- ated for fragments using shape indices of 2.0, 4.0 and 6.0. This range of values is realistic but slightly conservative; mean SI values for Taila?ndia ranged from 2.9 to 7.1 (Table 1), indicating that many fragments there were very irregularly Table 1. Description of forest fragments in two study areas in the eastern Amazon. Taila?ndia (2470 km2) and Paragominas (1280 km2) had 419 and 303 fragments, respectively. Fragment size- Per cent of fragments Per cent of forest area Mean fragment shapecategory (ha) Taila?ndia Paragominas Taila?ndia Paragominas Taila?ndia Paragominas <1 23.1 27.0 0.1 0.1 1.30 1.29 1?10 50.2 44.4 1.1 0.4 1.42 1.42 10?100 17.5 22.0 3.2 1.6 1.79 1.79 100?1000 6.9 5.5 14.8 5.6 2.33 2.94 1000?10 000 1.7 0.2 35.2 0.9 3.97 3.88 10 000?100 000 0.7 1.0 45.7 91.5 4.16 7.14 MARK A . COCHRANE AND WILL IAM F . LAURANCE318 Figure 2. Relationship between fragment area and shape index (SI) for study areas at Paragominas and Taila?ndia in the eastern Amazon. Although SI values were somewhat underestimated in small fragments (because of fractal effects), this analysis reveals that most large fragments were very irregularly shaped at a spatial scale that is relevant to edge effects in this system. Fire and forest fragmentation 319 Figure 3. Fire rotation-times as a function of distance from forest edge for Taila?ndia (diamonds) and Paragominas (triangles). The curves were ?tted with a smoothing function. Dotted lines show the 95% range of variation (mean ? 1.96 standard deviations) for forest interiors at Taila?ndia (more than 2500 m from edge). shaped. The core-area model (Figure 4) suggests that ?re regimes in even large fragments are being substantially altered. A fragment with an SI of 2.0, for example, would need to be about 90 000 ha in area to ensure that half of its total area will be unaffected by edge-related ?res. If the fragment were more irregularly shaped (SI = 6.0), it would need to be about ten times larger (890 000 ha) to retain half of its total area in natural condition. Fragments in Paragominas were generally less irregularly shaped than those at Taila?ndia, with mean SI values ranging from 2.3 to 4.3 (Table 1). The fragments in this older frontier, however, were often smaller, and the effects of ?res even more devastating, than was observed at Taila?ndia. No forest at Paragominas was further than 2.7 km from the nearest edge, and even at this distance, predicted ?re rotations were much shorter than those observed in forest interiors in Taila?ndia (Figure 3). Intact forest interiors have apparently MARK A . COCHRANE AND WILL IAM F . LAURANCE320 Figure 4. A core-area model showing the predicted impacts of edge-related ?res on fragmented rain forests in Taila?ndia, Brazil. vanished from the Paragominas area, and as a result it was not possible to estimate d and generate a core-area model. DISCUSSION Generality and limitations of ?ndings Our modelling results suggest that habitat fragmentation dramatically increases the vulnerability of rain forests in eastern Amazonia to anthropogenic ?res, and that such ?res are operating as an edge effect over surprisingly large spatial scales. Are our ?ndings typical? Both of our study areas are experiencing intensive exploitation; the Paragominas forests, in particular, have been severely degraded over the past several decades. Nevertheless, the spatial pat- terns of forest fragmentation at our two sites are representative of those caused by ranching and large-scale forest-colonization projects, the two most prevalent land uses in the Amazon today. In addition, both of our study areas occur in relatively seasonal areas of the basin. However, about half of the closed-canopy forests in Brazilian Amazonia experience comparably strong dry seasons, espe- cially in the eastern, southern and north-central areas of the basin (Nepstad et Fire and forest fragmentation 321 al. 1994, 1999), and thus could be similarly susceptible to ?re when fragmented. Analyses of recent Landsat TM imagery suggest that at least 45 000 000 ha of forest in Brazilian Amazonia (over 13% of the total remaining forest area) are currently vulnerable to edge-related ?res (Cochrane 2001). Our study was only 12?14 y in duration ? the period over which satellite imagery of suf?cient resolution was available ? and our results might be biased if weather conditions during this interval were unusually dry. During the study there were moderate El Nin?o droughts in 1986 and 1991 at both sites, and a strong drought in 1997 at Taila?ndia, and most forest burning occurred during these drier years (Cochrane 2000, Cochrane et al. 1999). Nevertheless, over the last century there have been 23 El Nin?o events in the Amazon (Suplee 1999), and the 1997 drought was comparable to other strong droughts, such as that in 1983. Thus, droughts or rainfall de?cits appear to be relatively common occurrences in the eastern Amazon, and the overall weather patterns during our study did not seem unusual. Previous studies suggest that, under natural conditions, major ?res are rare in Amazonian terra-?rme forests, perhaps occurring only once or twice per millennium on average (Meggers 1994, Piperno & Becker 1996, Saldarriaga & West 1986, Sanford et al. 1985). At Taila?ndia, however, ?res in intact forest interiors appeared to occur more frequently than this, with observed burns implying ?re return-intervals of 100?150 y (Figure 3). Although this pattern could have resulted from the relatively seasonal nature of the Taila?ndia cli- mate, we believe a more plausible explanation is that loggers or hunters occa- sionally ignited ?res in forest interiors. Because our study was relatively short- term, even a few small ?res in forest interiors could decrease the inferred ?re rotation-time substantially. If so, then our analysis of edge-related ?re fre- quency might be conservative, tending to underestimate the distance to which altered ?re regimes penetrate into forest interiors. It must be emphasized that our core-area model (Figure 4) predicts only the effects of habitat fragmentation on ?re frequency, and does not consider the in?uence of other factors, such as temporal variability in weather and local topography. Clearly, forest ?res vary in severity and frequency among years; the estimated ?re rotation-times used in our model (Figure 3) are 12?14-y averages, and may not be typical of any single year. Topography also in?uences ?re behaviour; trees along rivers and swamps, for example, are less likely to burn than those in upland areas (e.g. Kellman & Meave 1997). Nevertheless, ?re frequency in our study areas was increased so radically near fragment edges that local topographic effects were largely obscured. The high incidence of logging in fragmented Amazonian landscapes (e.g. Nepstad et al. 1999) will further increase the vulnerability of forest remnants to ?re. Implications for forest management Until the past few decades, major ?res have been very rare in Amazonian forests. As a result, few rain-forest species are adapted for ?re (Goldammer MARK A . COCHRANE AND WILL IAM F . LAURANCE322 1990, Kauffman & Uhl 1990) and even low-intensity surface ?res that usually originate in adjoining burned pastures kill many rain-forest trees (Cochrane & Schulze 1999, Kauffman 1991). Such surface ?res increase ?ne litter, wood debris and canopy openness, making forests highly vulnerable to intensive wild- ?res in the future. In recurring ?res, up to 98% of all trees become susceptible to ?re-induced mortality (Cochrane et al. 1999). An earlier study in Australia suggested that ?re return-intervals of less than 90 y might eliminate rain-forest tree species, while intervals of less than 20 y could eradicate all but short-lived pioneer trees (Jackson 1968). Our analyses suggest that forests within 2.35 km of edges at Taila?ndia, and all remaining forests in Paragominas, had average ?re return-intervals less than that evid- ently required to maintain rain-forest trees. Of equal concern is that ?re return-intervals were less than 20 y ? the threshold at which non-pioneer trees may be largely eradicated ? within 600?700 m of edges at Taila?ndia and at least 2500 m of edges at Paragominas (Figure 3). These patterns suggest that rain forests in both study areas will be largely replaced over time by anthropo- genic savannas and scrubby regrowth. Fire-adapted trees (e.g. Byrsonima spp., Curatella spp., certain palms) are unlikely to colonize these areas because there are no natural savannas or open woodlands nearby. The predicted deforestation process may be reinforced not only by the tendency for once-burned forest to become far more vulnerable to subsequent ?res (Cochrane & Schulze 1999, Cochrane et al. 1999), but also by reductions in rainfall caused by increasing deforestation (Lean & Warrilow 1989, Shukla et al. 1990) and by the moisture- trapping effects of smoke caused by extensive forest and pasture ?res (Rosenfeld 1999). Because fragmentation drastically increased ?re incidence, it appears likely that the margins of many forest fragments will recede over time, leading to a progressive ?implosion? of the fragments. Gascon et al. (2000) suggested that Amazonian fragments of less than 5000 ha will be susceptible to such effects, but our results imply that much larger fragments (up to several hundred thou- sand ha) could also be vulnerable to edge-related ?res. An important factor in this regard is that prevailing land-uses in the Amazon, such as forest- colonization projects, produce forest remnants that are highly irregular in shape and thus vulnerable to edge-related ?res and other external disturbances (Bierregaard et al. 1992, Laurance et al., in press, Lovejoy et al. 1986). Because forest margins will tend to erode over time, the predictions of our static core- area model may be conservative, and a dynamic modelling approach would be useful for predicting longer-term interactions of fragmentation and ?re. A key implication of our study (see Figure 4) is that even the largest Amazo- nian nature reserves, if isolated by swaths of deforested land, may be seriously degraded by ?res. This is especially likely to occur in the extensive areas of the basin with pronounced dry seasons. In such areas, nature reserves are likely to require deep buffers (> 3 km wide) of intact forest to ensure they are not Fire and forest fragmentation 323 chronically degraded by ?res and other edge-related phenomena. Adequate staf?ng and protection of reserves is also critical; a recent analysis of 86 federal parks and protected areas in Brazil found that 43% were at high to extreme risk because of illegal logging, deforestation, colonization, hunting, isolation of the reserve from other forest areas, and additional forms of encroachment. More than half of all reserves (54.6%) were judged to have nearly non-existent management (Ferreira et al. 1999). Future changes in climate could make the Amazon even more vulnerable to ?re. Some leading global-circulation models suggest that global warming may increase the frequency of El Nin?os (Timmerman et al. 1999) and warm-weather events (Mahlman 1997). Such changes, in concert with rapid forest fragmenta- tion and logging, have the potential to accelerate losses of rain forest across large expanses of the Amazon basin. ACKNOWLEDGEMENTS We thank Francis Putz, Claude Gascon, Philip Fearnside, Jeff Chambers and three anonymous referees for useful comments, and Christopher Barber for assistance with data analysis. Support was provided by the NASA-LBA program, A. W. Mellon Foundation, Basic Science and Remote Sensing Initiative (BSRSI) of Michigan State University, and U.S. Agency for International Development. This is publication number 355 in the Biological Dynamics of Forest Fragments Project technical series and number 01?04 in the BSRSI technical series. LITERATURE CITED BIERREGAARD, R. O., LOVEJOY, T. E., KAPOS, V., DOS SANTOS, A. A. & HUTCHINGS, R. W. 1992. The biological dynamics of tropical rain forest fragments. Bioscience 42:859?866. COCHRANE, M. A. 2000. Forest ?re, deforestation and landcover change in the Brazilian Amazon. Pp. 170?176 in Neuenschwander, L. F., Ryan, K. C., Gollberg, G. E. & Greer, J. D. (eds). Crossing the millennium: integrating spatial technologies and ecological principles for a new age in ?re management. University of Idaho, Moscow. COCHRANE, M. A. 2001. In the line of ?re: Understanding the impacts of tropical forest ?res. Environment 43:28?38. COCHRANE, M. A. & SCHULZE, M. D. 1998. Forest ?res in the Brazilian Amazon. Conservation Biology 12:948?950. COCHRANE, M. A. & SCHULZE, M. D. 1999. Fire as a recurrent event in tropical forests of the eastern Amazon: effects on forest structure, biomass, and species composition. Biotropica 31:2?16. COCHRANE, M. A. & SOUZA, C. M. 1998. Linear mixture model classi?cation of burned forests in the eastern Amazon. International Journal of Remote Sensing 19:3433?3440. COCHRANE, M. A., ALENCAR, A., SCHULZE, M. D., SOUZA, C. M., NEPSTAD, D. C., LEFEBVRE, P. & DAVIDSON, E. 1999. Positive feedbacks in the ?re dynamics of closed canopy tropical forests. Science 284:1832?1835. CURRAN, L. M., CANIAGO, I., PAOLI, G., ASTIANTI, D., KUSNETI, M., LEIGHTON, M., NIRARITA, C. & HAERUMAN, H. 1999. Impact of El Nin?o and logging on canopy tree recruitment in Borneo. Science 286:2184?2188. DIDHAM, R. K., GHAZOUL, J., STORK, N. E. & DAVIS, A. J. 1996. Insects in fragmented forests: a functional approach. Trends in Ecology and Evolution 11:255?260. FEARNSIDE, P. M. 1990. Environmental destruction in the Amazon. Pp. 179?225 in Goodman, D. & Hall, A. (eds). The future of Amazonia: destruction or sustainable development? MacMillan, London. FEARNSIDE, P. M. 1999. Biodiversity as an environmental service in Brazil?s Amazonian forests: risks, value and conservation. Environmental Conservation 26:305?321. MARK A . COCHRANE AND WILL IAM F . LAURANCE324 FERREIRA, L. V., DE SA?, R. M. L., BUSCHBACHER, R., BATMANIAN, G., BENSURAN, B. R. & COSTA, K. L. 1999. A?reas protegidas ou espac?os ameac?ados? World Wide Fund for Nature, Bras??lia. GASCON, C., WILLIAMSON, G. B. & DA FONSECA, G. A. B. 2000. Receding edges and vanishing reserves. Science 288:1356?1358. GOLDAMMER, J. G. (ed). 1990. Fire in the tropical biota. Springer-Verlag, New York. HOLDSWORTH, A. R. & UHL, C. 1997. Fire in the eastern Amazonian logged rain forest and the potential for ?re reduction. Ecological Applications 7:713?725. INPE 2001. Deforestation estimates in the Brazilian Amazon, 2000. National Institute for Space Research (INPE), Sa?o Jose dos Campos, Brazil. JACKSON, W. D. 1968. Fire, air, water and earth: an elemental ecology of Tasmania. Proceedings of the Ecological Society of Australia 2:9?16. KAPOS, V. 1989. Effects of isolation on the water status of forest patches in the Brazilian Amazon. Journal of Tropical Ecology 5:173?185. KAUFFMAN, J. B. 1991. Survival by sprouting following ?re in the tropical forests of the eastern Amazon. Biotropica 23:219?224. KAUFFMAN, J. B. & UHL, C. 1990. Interactions of anthropogenic activities, ?re, and rain forests in the Amazon Basin. Pp. 117?134 in Goldammer, J. (ed). Fire in the tropical biota. Springer-Verlag, New York. KAUFFMAN, J. B., CUMMINGS, D. L., WARD, D. E. & BABBITT, R. 1995. Fire in the Brazilian Amazon: biomass, nutrient pools, and losses in slashed primary forests. Oecologia 104:397?408. KELLMAN, M. & MEAVE, J. 1997. Fire in the tropical gallery forests of Belize. Journal of Biogeography 24:23?34. KLEIN, B. C. 1989. Effects of forest fragmentation on dung and carrion beetle communities in central Amazonia. Ecology 70:1715?1725. LAURANCE, W. F. 1998. A crisis in the making: responses of Amazonian forests to land use and climate change. Trends in Ecology and Evolution 13:411?415. LAURANCE, W. F. 2000. Do edge effects occur over large spatial scales? Trends in Ecology and Evolution 15:134?135. LAURANCE, W. F. & YENSEN, E. 1991. Predicting the impacts of edge effects in fragmented habitats. Biological Conservation 55:77?92. LAURANCE, W. F., LAURANCE, S. G., FERREIRA, L. V., RANKIN-DE MERONA, J. M., GASCON, C. & LOVEJOY, T. E. 1997. Biomass collapse in Amazonian forest fragments. Science 278:1117?1118. LAURANCE, W. F., FERREIRA, L. V., RANKIN-DE MERONA, J. M. & LAURANCE, S. G. 1998a. Rain forest fragmentation and the dynamics of Amazonian tree communities. Ecology 79:2032?2040. LAURANCE, W. F., LAURANCE, S. G. & DELAMONICA, P. 1998b. Tropical forest fragmentation and greenhouse gas emissions. Forest Ecology and Management 110:173?180. LAURANCE, W. F., COCHRANE, M. A., BERGEN, S., FEARNSIDE, P. M., DELAMONICA, P., D?ANGELO, S., BARBER, C. & FERNANDES, T. 2001a. The future of the Brazilian Amazon. Science 291:438?439. LAURANCE, W. F., PEREZ-SALICRUP, D., DELAMONICA, P., FEARNSIDE, P. M., D?ANGELO, S., JEROZOLINSKI, A., POHL, L. & LOVEJOY, T. E. 2001b. Rain forest fragmentation and the structure of Amazonian liana communities. Ecology 82:105?116. LAURANCE, W. F., LOVEJOY, T. E., VASCONCELOS, H. L., BRUNA, E., DIDHAM, R. K., STOUFFER, P. C., GASCON, C., BIERREGAARD, R. O., LAURANCE, S. G. & SAMPIAO, E. In press. Ecosystem decay of Amazonian forest fragments, a 22-year investigation. Conservation Biology. LEAN, J. & WARRILOW, D. A. 1989. Simulation of the regional climatic impact of Amazon deforestation. Nature 342:411?413. LI, H. & REYNOLDS, J. F. 1994. A simulation experiment to quantify spatial heterogeneity in categorical maps. Ecology 75:2446?2455. LOVEJOY, T. E., BIERREGAARD, R. O., RYLANDS, A. B., MALCOLM, J. R., QUINTELA, C. E., HARPER, L. H., BROWN, K. S., POWELL, A. H., POWELL, G. V. N., SCHUBART, H. & HAYS, M. 1986. Edge and other effects of isolation on Amazon forest fragments. Pp. 257?285 in Soule?, M. E. (ed). Conservation biology: the science of scarcity and diversity. Sinauer, Sunderland. MAHLMAN, J. D. 1997. Uncertainties in projections of human-caused climate warming. Science 278:1416?1417. MEGGERS, B. J. 1994. Archeological evidence for the impact of mega-Nin?o events on Amazonian forests during the past two millennia. Climate Change 28:321?338. NEPSTAD, D. C., CARVALHO, C. R., DAVIDSON, E. A., JIPP, P. H., LEFEBVRE, P. A., NEGREIROS, P., SILVA, E. D., STONE, T. A., TRUMBORE, S. E. & VIEIRA, S. 1994. The role of deep roots in the hydrological cycles of Amazonian forests and pastures. Nature 372:666?669. NEPSTAD, D. C., MOUTINHO, P. R., UHL, C., VIEIRA, I. C. & DA SILVA, J. M. C. 1996. The ecological importance of forest remnants in an eastern Amazonian frontier landscape. Pp. 133?150 Fire and forest fragmentation 325 in Schelhas, J. & Greenberg, R. (eds). Forest patches in tropical landscapes. Island Press, Washington, DC. NEPSTAD, D. C., VERISSIMO, A., ALENCAR, A., NOBRE, C., LIMA, C., LEFEBVRE, P., SCHLESINGER, P., POTTER, C., MOUTINHO, P., MENDOZA, E., COCHRANE, M. & BROOKS, V. 1999. Large-scale impoverishment of Amazonian forests by logging and ?re. Nature 398:505?508. PIPERNO, D. R. & BECKER, P. 1996. Vegetational history of a site in the central Amazon Basin derived from phytolith and charcoal records from natural soils. Quaternary Research 45:202?209. ROSENFELD, D. 1999. TRMM observed ?rst direct evidence of smoke from forest ?res inhibiting rainfall. Geophysical Research Letters 26:3105?3108. SALDARRIAGA, J. & WEST, D. C. 1986. Holocene ?res in the northern Amazon basin. Quaternary Research 26:358?366. SANFORD, R. L., SALDARRIAGA, J., CLARK, K., UHL, C. & HERRERA, R. 1985. Amazon rain-forest ?res. Science 227:53?55. SHUKLA, J., NOBRE, C. A. & SELLERS, P. 1990. Amazon deforestation and climate change. Science 247:1322?1325. SKOLE, D. & TUCKER, C. J. 1993. Tropical deforestation and habitat fragmentation in the Amazon: satellite data from 1978 to 1988. Science 260:1905?1910. SUPLEE, C. 1999. El Nin?o/La Nin?a: nature?s vicious cycle. National Geographic Magazine, March, pp. 21?34. TIMMERMAN, A., OBERHUBER, J., BACHER, A., ESCH, M., LATIF, M. & ROECKNER, E. 1999. Increased El Nin?o frequency in a climate model forced by future greenhouse warming. Nature 398:694?697. UHL, C. & KAUFFMAN, J. B. 1990. Deforestation effects on ?re susceptibility and the potential response of the tree species to ?re in the rain forest of the eastern Amazon. Ecology 71:437?449. VAN WAGNER, C. E. 1978. Age-class distribution and the forest ?re cycle. Canadian Journal of Forest Research 8:220?227. WHITMORE, T. C. 1997. Tropical forest disturbance, disappearance, and species loss. Pp. 3?12 in Laurance, W. F. & Bierregaard, R. O. (eds). Tropical forest remnants: ecology, management, and conservation of fragmented communities. University of Chicago Press, Chicago.