Species loss, delayed numerical responses, and functional compensation in an antbird guild

When a community loses species through fragmentation, its total food consumption may drop. Compensatory responses of remaining species, whereby survivors assume roles of extinct competitors, may reduce the impact of species loss through numerical or functional responses. We measured compensatory responses in two remaining antbird species on Barro Colorado Island, Panama, four decades after the loss of their dominant competitor, the Ocellated Antbird, Phaenostictus mcleannani. We compared current abundances and behavior of these two species on Barro Colorado to those reported before the island lost Ocellated Antbirds, and to those in a nearby mainland population where all three species still exist as a space-for-time substitution. The smaller, more subordinate Spotted Antbird, Hylophylax naevioides, responded far more strongly than the larger Bicolored Antbird, Gymnopithys leucaspis, which is functionally more like the Ocellated Antbird. Islandwide density of Spotted Antbirds has more than doubled since the loss of Ocellated Antbirds. Moreover, Spotted Antbirds now spend so much more of their time following ant swarms that their metabolic biomass at these swarms has more than tripled since Ocellated Antbirds disappeared. These responses in Spotted Antbirds were apparently delayed by >20 years. Bicolored Antbirds have not increased substantially in islandwide density or metabolic biomass at ant swarms. We hypothesize that behavioral flexibility, as shown by Spotted Antbirds on Barro Colorado Island, is a major factor governing the extent to which fragmented ecosystems can buffer the impacts of species loss.     

Avian Survival Rates and the Extinction Process on Barro Colorado Island, Panama

Abstract: Selective extinction following isolation of habitat patches may be due to biogeographical (e.g., island size or isolation) and ecological (species natural histories, interspecifc interactions) factors, or their interactions. Among the demographic and life history attributes commonly associated with high extinction probability are small populations, large size of individuals, and population variability. Long-term capture-recapture data from forest habitat in central Panama permit an examination of the association between mainland survival rates and extinction on a nearby land-bridge island Species of birds that no longer occur on Barro Colorado Island (BCI), Panama, have, on average, lower survival rates on the adjacent mainland than species that have persisted on BCI. Moreover, of the species that no longer occur on BCI, those with lower mainland survival rates generally disappeared earlier from the island. My analysis provides little evidence of a relationship between extinction and population size. Recolonization of BCI from the adjacent mainland by the forest undergrowth species studied here is unlikely. Reduced reproductive success on BCI combined with naturally low adult survival rates seems to be responsible for these BCI extinctions. High nest predation and/or altered landscape dynamics are probable agents in the low reproductive success. The methods used here could be employed in other circumstances to identify fragmentation-sensitive species.     

Evolution of Nesting Height in an Endangered Hawaiian Forest Bird in Response to a Non‐Native Predator

Abstract: The majority of bird extinctions since 1800 have occurred on islands, and non‐native predators have been the greatest threat to the persistence of island birds. Island endemic species often lack life‐history traits and behaviors that reduce the probability of predation and they can become evolutionarily trapped if they are unable to adapt, but few studies have examined the ability of island species to respond to novel predators. The greatest threat to the persistence of the Oahu Elepaio (Chasiempis ibidis), an endangered Hawaiian forest bird, is nest predation by non‐native black rats (Rattus rattus). I examined whether Oahu Elepaio nest placement has changed at the individual and population levels in response to rat predation by measuring nest height and determining whether each nest produced offspring from 1996 to 2011. Average height of Oahu Elepaio nests increased 50% over this 16‐year period, from 7.9 m (SE 1.7) to 12.0 m (SE 1.1). There was no net change in height of sequential nests made by individual birds, which means individual elepaios have not learned to place nests higher. Nests ≤3 m off the ground produced offspring less often, and the proportion of such nests declined over time, which suggests that nest‐building behavior has evolved through natural selection by predation. Nest success increased over time, which may increase the probability of long‐term persistence of the species. Rat control may facilitate the evolution of nesting height by slowing the rate of population decline and providing time for this adaptive response to spread through the population.     

Effect of Slash-and-Burn Shifting Cultivation on Rainforest Birds in Mizoram, Northeast India

Abstract: Slash-and-burn shifting cultivation, or jhum , the predominant form of agriculture in the hill tracts of northeast India, is believed to have caused considerable loss of forest cover in the species-rich tropical rainforests of the region. In this study I sought to understand how rainforest bird communities are affected by shifting cultivation in Mizoram State. I studied bird occurrence and abundance patterns in secondary successional and mature tropical rainforests in a shifting-cultivation habitat mosaic in Dampa Tiger Reserve. To compare replicate sites in fallows aged 1, 5, 10, 25, and 100 years with undisturbed primary forest, I used systematic strip-transect sampling over the winter and early summer ( breeding) seasons during 1994–1995. Many forest bird species, especially those with ranges restricted to northeast India, declined in abundance or disappeared in successional fallows that had regenerated for ≤10 years. Birds that colonized or increased in abundance in regrowth habitats were mainly common and widespread species of open-country and secondary-forest habitats, and of low importance for conservation. Primary forest was the main habitat for specialized forest birds, intrinsically rare species, and elevational migrants. Although protection and conservation of relatively undisturbed mature forests is imperative in the core area of the reserve, management in the buffer zone for long-rotation shifting cultivation (>10-year cycles) instead of plantation of monocultures may be important until alternate means of livelihood are available to cultivators.     

Neutral theory and the evolution of ecological equivalence

Since the publication of the unified neutral theory in 2001, there has been much discussion of the theory, pro and con. The hypothesis of ecological equivalence is the fundamental yet controversial idea behind neutral theory. Assuming trophically similar species are demographically alike (symmetric) on a per capita basis is only an approximation, but it is equivalent to asking: How many of the patterns of ecological communities are the result of species similarities, rather than of species differences? The strategy behind neutral theory is to see how far one can get with the simplification of assuming ecological equivalence before introducing more complexity. In another paper, I review the empirical evidence that led me to hypothesize ecological equivalence among many of the tree species in the species-rich tropical forest on Barro Colorado Island (BCI). In this paper, I develop a simple model for the evolution of ecological equivalence or niche convergence, using as an example evolution of the suite of life history traits characteristic of shade tolerant tropical tree species. Although the model is simple, the conclusions from it seem likely to be robust. I conclude that ecological equivalence for resource use are likely to evolve easily and often, especially in species-rich communities that are dispersal and recruitment limited. In the case of the BCI forest, tree species are strongly dispersal- and recruitment-limited, not only because of restricted seed dispersal, but also because of low recruitment success due to heavy losses of the seedling stages to predators and pathogens and other abiotic stresses such as drought. These factors and the high species richness of the community strongly reduce the potential for competitive exclusion of functionally equivalent or nearly equivalent species.     

Directional changes in the species composition of a tropical forest

Long-term studies have revealed that the structure and dynamics of many tropical forests are changing, but the causes and consequences of these changes remain debated. To learn more about the forces driving changes within tropical forests, we investigated shifts in tree species composition over the past 25 years within the 50-ha Forest Dynamics Plot on Barro Colorado Island (BCI), Panama, and examined how observed patterns relate to predictions of (1) random population fluctuations, (2) carbon fertilization, (3) succession from past disturbance, (4) recovery from an extreme El Ni?o drought at the start of the study period, and (5) long-term climate change. We found that there have been consistent and directional changes in the tree species composition. These shifts have led to increased relative representations of drought-tolerant species as determined by the species' occurrence both across a gradient of soil moisture within BCI and across a wider precipitation gradient from a dry forest near the Pacific coast of Panama to a wet forest near its Caribbean coast. These nonrandom changes cannot be explained by stochastic fluctuations or carbon fertilization. They may be the legacy of the El Ni?o drought, or alternatively, potentially reflect increased aridity due to long-term climate change. By investigating compositional changes, we increased not only our understanding of the ecology of tropical forests and their responses to large-scale disturbances, but also our ability to predict how future global change will impact some of the critical services provided by these important ecosystems.     

Group size and growth rate in hatchling green iguanas (Iguana iguana)

Summary The post-hatching growth and sleeping group sizes of green iguanas (Iguana iguana) were followed on Barro Colorado Island, Panama, for 3 successive years. The laboratory clearing was subdivided and surveyed routinely for iguanas, most intensively the six weeks following onset of hatching. A similar pattern of influx of iguanas into the clearing took place each May. Over the early weeks the size of groups generally increased. Animals sleeping in larger groups had significantly faster growth rates. Animals that disappeared from the clearing prior to July had larger group sizes prior to disappearance than those that remained. Growth rates measured in animals that remained declined markedly after June. Possible reasons for the social grouping of young iguanas are discussed.     

Darwin's Fox: A Distinct Endangered Species in a Vanishing Habitat

The temperate rain forest of Chiloé Island, Chile, is inhabited by an endemic fox ( Dusicyon fulvipes ) first described by Charles Darwin and now designated Darwin's fox. Despite morphological differences, Darwin's fox has been considered only an insular subspecies of the mainland chilla fox ( D. griseus ). This follows the assumption that the island population, with an estimated population of less than 500, has been separated from the mainland chilla fox for only about 15,000 years and may have received occasional immigrants from the mainland. Consequently, this island population has not been protected as endangered or bred in captivity. Recently, a population of Darwin's fox was discovered on the Chilean mainland 600 km north of Chiloé Island. This population exists in sympatry with chilla and possibly culpeo ( D. culpaeus ) foxes, which suggests that Darwin's fox may be reproductively isolated. To clarify the phylogenetic position of Darwin's fox, we analyzed 344 bp of mitochondrial DNA control-region sequence of the three species of Chilean foxes. Darwin's foxes from the island and mainland populations compose a monophyletic group distinct from the two other Chilean fox species. This indicates that Darwin's fox was probably an early inhabitant of central Chile, and that its present distribution on the mainland may be a relict of a once much wider distribution. Our results highlight the ability of molecular genetic techniques to uncover historical relationships masked by recent events, such as local extinctions. The "rediscovery" of Darwin's fox as a distinct species implies that greater significance should be given to the protection of this species and its unique habitat and to documenting the extent of its mainland distribution.     

Application of Extinction and Conservation Theories for Forest Birds in Nicaragua

Abstract: An increasing number of empirical studies have been done on the effects of tropical forest fragmentation on avian communities, but few researchers have applied these theories to assess the vulnerability of birds in poorly researched countries such as Nicaragua. I used a logistic regression to determine which natural-history characteristics were most important in predicting a list of threatened birds known to occur in Nicaragua. The best model included five macroecological variables ( body weight, habitat specificity, trophic group, forest preference, and biogeography within Nicaragua). I used this model to generate predicted probabilities of extinction for all forest birds in Nicaragua. The predicted probability of extinction from the best model ranked 63% of the extinction-prone birds from La Selva, Costa Rica, and 59% of the extinction-prone birds from Barro Colorado, Panama, in the first quartile of all forest birds recorded in Nicaragua. This method provides a first-order approximation of which species deserve global and national priorities for conservation. The central and Atlantic regions of Nicaragua deserve high priority for conservation at a global scale, whereas the Atlantic region deserves the highest priority for conservation at a national scale. The Nicaraguan Ministry of Natural Resources and the Environment has done an adequate job of identifying areas for conservation based on the proportion of decreed nature reserves in each biogeographic region and the distribution of forest birds with a high predicted probability of extinction. Forest birds in central Nicaragua, however, may currently be the most vulnerable to local extinction because of low forest cover within decreed reserves.     

Life history trade-offs in tropical trees and lianas

It has been hypothesized that tropical trees partition forest light environments through a life history trade-off between juvenile growth and survival; however, the generality of this trade-off across life stages and functional groups has been questioned. We quantified trade-offs between growth and survival for trees and lianas on Barro Colorado Island (BCI), Panama using first-year seedlings of 22 liana and 31 tree species and saplings (10 mm < dbh < 39 mm) of 30 tree species. Lianas showed trade-offs similar to those of trees, with both groups exhibiting broadly overlapping ranges in survival and relative growth rates as seedlings. Life history strategies at the seedling stage were highly correlated with those at the sapling stage among tree species, with all species showing an increase in survival with size. Only one of 30 tree species demonstrated a statistically significant ontogenetic shift, having a relatively lower survival rate at the sapling stage than expected. Our results indicate that similar life history trade-offs apply across two functional groups (lianas and trees), and that life history strategies are largely conserved across seedling and sapling life-stages for most tropical tree species.     

Understanding Trait‐Dependent Community Disassembly: Dung Beetles,Density Functions,and Forest Fragmentation

Abstract: Anthropogenic disturbances such as fragmentation are rapidly altering biodiversity, yet a lack of attention to species traits and abundance patterns has made the results of most studies difficult to generalize. We determined traits of extinction‐prone species and present a novel strategy for classifying species according to their population‐level response to a gradient of disturbance intensity. We examined the effects of forest fragmentation on dung beetle communities in an archipelago of 33 islands recently created by flooding in Venezuela. Species richness, density, and biomass all declined sharply with decreasing island area and increasing island isolation. Species richness was highly nested, indicating that local extinctions occurred nonrandomly. The most sensitive dung beetle species appeared to require at least 85 ha of forest, more than many large vertebrates. Extinction‐prone species were either large‐bodied, forest specialists, or uncommon. These explanatory variables were unrelated, suggesting at least 3 underlying causes of extirpation. Large species showed high wing loading (body mass/wing area) and a distinct flight strategy that may increase their area requirements. Although forest specificity made most species sensitive to fragmentation, a few persistent habitat generalists dispersed across the matrix. Density functions classified species into 4 response groups on the basis of their change in density with decreasing species richness. Sensitive and persistent species both declined with increasing fragmentation intensity, but persistent species occurred on more islands, which may be due to their higher baseline densities. Compensatory species increased in abundance following the initial loss of sensitive species, but rapidly declined with increasing fragmentation. Supertramp species (widespread habitat generalists) may be poor competitors but strong dispersers; their abundance peaked following the decline of the other 3 groups. Nevertheless, even the least sensitive species were extirpated or rare on the smallest and most isolated islands.     

The Early Development of Forest Fragmentation Effects on Birds

The early development of forest fragmentation effects on forest organisms is poorly understood partly because most fragmentation studies have been done in agricultural or suburban landscapes, long after the onset of fragmentation. We develop a temporal model of forest fragmentation effects on densities of forest-breeding birds and provide data from an active industrial forest landscape to test the model. The model and our empirical data indicate that densities of several forest-dwelling bird species can increase within a forest stand soon after the onset of fragmentation as a result of displaced individuals packing into remaining habitat. Along with higher densities in the newly formed fragments, pairing success in one species, the Ovenbird ( Seiurus aurocapillus ), was lower in fragments than nonfragments, possibly due to behavioral dysfunction resulting from high densities. Thus, density was inversely related to productivity. The duration and extent of increased densities following onset of fragmentation depends on many factors, including the sensitivity of a species to edge and area effects, the duration and rate of habitat loss and fragmentation, and the proximity of a forest stand to the disturbance. Incipient forest fragmentation may affect populations differently from later stages of fragmentation when the geometry of the landscape has reached a more stable configuration. Our model and data indicate, for reasons unrelated to traditional edge effects, that large tracts of forest can be important because they are relatively free from the variety of plant and animal population dynamics that might take place near new edges, including the encroachment of individuals displaced by habitat loss.     

Adaptive Units for Conservation: Population Distinction and Historic Extinctions in the Island Scrub-Jay

Abstract:  The Island Scrub-Jay ( Aphelocoma insularis ) is found on Santa Cruz Island, California, and is the only insular bird species in the continental United States. We typed seven microsatellite loci and sequenced a portion of the mitochondrial DNA control region of Island Scrub-Jays and their closest mainland relative, the Western Scrub-Jay ( Aphelocoma californica ), to assess levels of variability and effective population size and to examine the evolutionary relationship between the two species. The estimated female effective population size, N _ef, of the Island Scrub-Jay was 1603 (90% confidence interval: 1481–1738) and was about 7.5% of the size of the mainland species. Island and Western Scrub-Jays have highly divergent control-region sequences, and the value of 3.14 ± 0.09% sequence divergence between the two species suggests a divergence time of approximately 151,000 years ago. Because the four northern Channel Islands were joined as one large island as recently as 11,000 years ago, extinctions must have occurred on the three other northern Channel islands, Santa Rosa, San Miguel, and Anacapa, highlighting the vulnerability of the remaining population. We assessed the evolutionary significance of four island endemics, including the Island Scrub-Jay, based on both genetic and adaptive divergence. Our results show that the Island Scrub-Jay is a distinct species of high conservation value whose history and adaptive potential is not well predicted by study of other island vertebrates.     

Heavy Extinctions of Forest Avifauna in Singapore: Lessons for Biodiversity Conservation in Southeast Asia

Abstract: The consequences of rapid rainforest clearance on native avifauna are poorly understood. In Southeast Asia, Singapore, a newly developing country, has had 95% of its native lowland rainforest cleared. Most of the rainforest was lost in the mid- to late-nineteenth century. We compared avifauna checklists from 1923, 1949, and 1998 to determine the extent of extinctions between 1923 and 1998 in Singapore. Of 203 diurnal bird species, 65 were extirpated in Singapore in the past 75 years. Four of these species were nonforest- dependent species, whereas 61 (94%) were forest bird species dependent on the primary or old secondary forest to survive. Twenty-six forest bird species became extinct between 1923 and 1949, whereas 35 forest species disappeared after 1949. We compared the body lengths, feeding guilds, and vertical feeding zones between extinct and extant forest bird species to determine whether extinction patterns were dependent on these characteristics. Larger forest bird species went extinct between 1923 and 1949. Body sizes, however, did not affect the loss of forest bird species between 1949 and 1998. We observed high losses of insectivorous birds; the insectivore-carnivore and insectivore-granivore guilds lost> 80% of the species present in 1923. The highest losses were among birds that fed in the canopy. None of the forest bird species are currently common (>100 individuals/species) within Singapore. Our study shows that more than half the forest avifauna became locally extinct after extensive deforestation. Based on this fact, the countries within Southeast Asia should reconsider their heavy deforestation practices.     

Quantifying the global threat to native birds from predation by non-native birds on small islands

Although invasive non-native species can adversely affect biodiversity in many ways, predation of native species by non-native species on islands can be severely damaging. Results of numerous studies document non-native birds preying on birds on islands, but our understanding of the number and type of species affected has been limited by the lack of a global review of these impacts. I identified the non-native bird species that have been recorded preying on birds, the locations where this predation occurred, and the bird species affected. Because the impacts of non-native birds can be particularly severe on small islands, I then identified the islands <500 km² around the world that are occupied by predatory non-native birds. By taking into account their life-history traits and predation history, I also identified the near-threatened and threatened bird species on these islands that they may prey on. The results indicated that predation by non-native birds was primarily a concern for threatened bird conservation on small islands; almost all predation impacts (91%) on near-threatened and threatened birds were recorded on islands, and median island size was 106 km². I also found non-native bird predation was a poorly known and widespread potential threat to avian biodiversity; worldwide, 194 islands of <500 km² were occupied by predatory non-native birds, but information on their impacts was unavailable for most of these islands. On them, where the impacts of non-native species can be severe, non-native birds may be preying on approximately 6% of the world's near-threatened and threatened bird species. Four non-native bird species I identified have been successfully eradicated from islands. If they were eradicated from the small islands they occupy, 70% of the near-threatened and threatened bird species I identified would no longer be affected by nest predation by non-native birds on small islands.     

Soil drying in a tropical forest: Three distinct environments controlled by gap size

Soil water and temperature regimes in the tropical moist forest on Barro Colorado Island, Panama, were simulated directly from meteorological data using the model SWEAT. Separate field observations from root-exclusion, litter-removal and control treatments in one small and one large forest gap were used for calibration and validation. After irrigating all treatments to field capacity, soil matric potential and temperature were measured over 17 days at four depths ≤50 mm using the filter-paper technique and bead thermistors. Understorey environments were also simulated under the same initial conditions. The results suggest that three distinct scenarios, controlled by gap size, describe how the above- and below-ground processes controlling soil drying are coupled: (1) in the large gap, root water extraction by surrounding trees is negligible so soil drying is dominated by evaporation from the soil surface. Soil temperature is dominated by direct solar heating and cooling due to evaporation. (2) In the small gap, root water extraction dominates soil drying with soil evaporation playing a minor role. Soil temperature is still dominated by direct sunlight with some cooling due to evaporation. (3) In the understorey, root water extraction dominates soil drying. Soil temperature is dominated by heat conduction from deep soil layers with some evaporation and sensible heat transfer. The contrasting soil drying regimes imposed by variation in canopy structure enhance micro-environmental heterogeneity and the scope for differential germination and seedling establishment in coexisting tropical tree species.     

Deforestation and Avian Extinction on Tropical Landbridge Islands

Abstract: There are few empirical data, particularly collected simultaneously from multiple sites, on extinctions resulting from human‐driven land‐use change. Southeast Asia has the highest deforestation rate in the world, but the resulting losses of biological diversity remain poorly documented. Between November 2006 and March 2008, we conducted bird surveys on six landbridge islands in Malaysia and Indonesia. These islands were surveyed previously for birds in the early 1900s, when they were extensively forested. Our bird inventories of the islands were nearly complete, as indicated by sampling saturation curves and nonparametric true richness estimators. From zero (Pulau Malawali and Pulau Mantanani) to 15 (Pulau Bintan) diurnal resident landbird species were apparently extirpated since the early 1900s. Adding comparable but published extinction data from Singapore to our regression analyses, we found there were proportionally fewer forest bird extinctions in areas with greater remaining forest cover. Nevertheless, the statistical evidence to support this relationship was weak, owing to our unavoidably small sample size. Bird species that are restricted to the Indomalayan region, lay few eggs, are heavier, and occupy a narrower habitat breadth, were most vulnerable to extinction on Pulau Bintan. This was the only island where sufficient data existed to analyze the correlates of extinction. Forest preservation and restoration are needed on these islands to conserve the remaining forest avifauna. Our study of landbridge islands indicates that deforestation may increasingly threaten Southeast Asian biodiversity.     

Importance of Reserve Size and Landscape Context to Urban Bird Conservation

Abstract:  We tested whether reserve size, landscape surrounding the reserve, and their interaction affect forest songbirds in the metropolitan area of Seattle, Washington (U.S.A.), by studying 29 reserves of varying size (small, medium, large) and surrounding urbanization intensity (urban, suburban, exurban). Larger reserves contained richer and less even bird communities than smaller reserves. These size effects disappeared when we removed the positive correlation of shrub diversity with reserve size, suggesting that greater habitat diversity in large reserves supported additional species, some of which were rare. Standardizing the number of individuals detected among all reserve size classes reversed the effect of size on richness in exurban landscapes and reduced the magnitude of the effect in suburban or urban landscapes. The latter change suggested that richness increased with reserve size in most landscapes because larger areas also supported larger samples from the regional bird species pool. Most bird species associated with native forest habitat (native forest species) and with human activity (synanthropic species) were present in reserves larger than 42 ha and surrounded by >40% urban land cover, respectively. Thus, we recommend these thresholds as means for conserving the composition of native bird communities in this mostly forested region. Native forest species were least abundant and synanthropic species most abundant in urban landscapes, where exotic ground and shrub vegetation was most common. Therefore, control of exotic vegetation may benefit native songbird populations. Bird nests in shrubs were most dense in medium (suburban) and large reserves (urban) and tended to be most successful in medium (suburban) and large reserves (exurban), potentially supplying another mechanism by which reserve size increased retention of native forest species.     

Effects of Human Subsistence Activities on Forest Birds in Northern Kenya

Abstract: Indigenous tribes and conservation biologists may have common goals and may be able to collaborate on the maintenance of biodiversity, but few researchers have evaluated the impacts and potential benefits of human subsistence activities. I studied the effects of subsistence activities (primarily wood collection) of nomadic pastoralists in 3 Afromontane forests of northern Kenya. In surveys of 404, 25‐m‐radius plots, I recorded vegetation structure and composition of the forest bird community. Plots with higher levels of human activity had significantly different vegetation structure, with more‐open canopies, more grass, and fewer tree stems. Nectarivores (abundance +231%) and aerial insectivores (+66%) were more abundant in plots with more‐intense wood collecting than in plots with less human activity, whereas abundance of forest specialists (?28%) decreased in plots with more‐intense human activity. Abundance of 58% of the bird species either increased or decreased significantly in plots with more‐intense human activity. Generally, the number of individuals of forest specialists decreased (6 of 7 species showed significant responses) and the number of individuals of edge and nonforest species increased with increasing human activity. Canonical correspondence analysis showed that an intensification of human activities would favor nectarivores, aerial insectivores, granivores, and omnivores and would negatively affect large‐sized, ground‐foraging species and arboreal frugivores. Subsistence human activities favored the invasion of forest by edge species at the expense of forest specialists; thus, further intensification of forest exploitation by local peoples is not recommended. At the same time, however, subsistence activities in northern Kenya forests appeared to increase the structural diversity of the vegetation and provided suitable habitat for part (but not all) of the forest avifauna, which suggests that subsistence human activities may have a role in the maintenance of bird diversity.     

Effects of Habitat Fragmentation on Bird Species in a Relict Temperate Forest in Semiarid Chile

Abstract: We analyzed the structure and composition of a bird assemblage in a fragmented relict temperate forest located in northcentral Chile ( Fray Jorge National Park). In terms of species composition, the bird assemblage we found in Fray Jorge was more similar to southern temperate forest sites, located more than 1200 km south of Fray Jorge, than to localities found in nearby scrub habitats. The relict character and long-term isolation of the Fray Jorge forest provides a natural experiment with which to establish the potential long-term effects of fragmentation and isolation on southern Chilean temperate forests. Between May 1996 and March 1997, we conducted seasonal surveys of birds in six forest fragments, ranging in size from 0.5 to 22.5 ha, at Fray Jorge. The number of bird species at each forest fragment was positively correlated with fragment area during all seasons. The relict forest system had a steeper species-area slope than that reported for similar temperate-forest bird assemblages in forest fragments within Chiloé Island and for islands across the Chiloé Archipelago in southern Chile. In this regard, this bird fauna resembled a depauperate oceanic archipelago. This difference in area effects is likely a consequence of the minimization of rescue effects because of the absence of large source forest areas nearby and the long-term isolation of the system. In addition, the distribution of species among forest fragments in Fray Jorge was not random, showing a nested subset pattern. Thus, some species occur across all fragments, regardless of their area, and therefore are less affected by habitat fragmentation and less prone to local extinction. These results suggest that, for south-temperate forest birds, large fragments (or reserves) should afford better protection against extinction than small forest patches.