Nested Subsets and the Distribution of Birds on Isolated Woodlots

Abstract. Distribution of bird species among isolated habitat patches (e.g., woodlots in an agricultural landscape) often appears to be nonrandom; species present in small, species-poor patches also are found in larger patches that support more species. Bird communities form 'nested subsets' (after Patterson & Atmar 1986) if all species found in small faunas also are found in more species-rich assemblages. Occurrence of a nested subset pattern implies an underlying, nonrandom pattern of species distributions. I used computer simulations to analyze the degree of nestedness exhibited by bird communities in east-central Illinois. Results demonstrated that the distribution of bird species breeding in isolated woodlots (1.8 to 600 ha) differed significantly from that predicted by a random distribution model; species assemblages were more "nested" than expected by chance. Most species present in small, species-poor woodlots also were found in larger, species-rich woodlots. As groups, species requiring forest interior habitat for breeding and species wintering in the tropics showed highly nested distributions. In contrast, short-distance migrants and species breeding in forest edge habitat showed more variable distribution patterns, species recorded on smaller woodlots were not always recorded on larger, more species-rich wood-lots. Apparent absences from larger woodlots may have reflected real distribution patterns or insufficient sampling of edge habitats. These results support previous conclusions that small habitat patches are insufficient for preservation of many species.     

Using Montane Mammals to Model Extinctions Due to Global Change

We use data on the species-area relationship and the nested subset structure of the boreal mammal faunas inhabiting isolated mountaintops in the Great Basin to develop a simple quantitative model that predicts the number and identity of species that would go extinct under an assumed scenario of changing climate and vegetation. Global warming of 3°C is predicted to cause the loss of 9–62% of the species inhabiting each mountain range and the extinction of three of fourteen species throughout the region. These results suggest (1) that it is possible to make highly plausible predictions about the susceptibility of species to extinction without detailed information about their population biology, and (2) that global and regional environmental changes seriously threaten the survival of species that are restricted in distribution to both natural "habitat islands" and biological reserves.     

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.     

The Potential for Conservation of Polynesian Birds through Habitat Mapping and Species Translocation

Abstract: The dependency of highly endemic island floras on few potential pollinators in depauperate island faunas suggests that pollinators and seed dispersers may be crucial in the preservation of biodiversity in isolated oceanic islands. We discuss the hypothesis that flying foxes are "strong interactors" in South Pacific islands where they setwe as the principal pollinators and seed dispersers, This suggests that the ongoing decline and ultimate extinction of flying fox species on Pacific islands may lead to a cascade of linked plant extinctions. We propose an empirical test of this hypothesis: comparisons of plant reproductive success in Guam, which has virtually lost its flying fox populations, and Samoa, where signifcant populations remain.     

Flying Foxes as Strong Interactors in South Pacific Island Ecosystems: A Conservation Hypothesis

Abstract: The dependency of highly endemic island floras on few potential pollinators in depauperate island faunas suggests that pollinators and seed dispersers may be crucial in the preservation of biodiversity in isolated oceanic islands. We discuss the hypothesis that flying foxes are "strong interactors" in South Pacific islands where they serve as the principal pollinators and seed dispersers, This suggests that the ongoing decline and ultimate extinction of flying fox species on Pacific islands may lead to a cascade of linked plant extinctions. We propose an empirical test of this hypothesis: comparisons of plant reproductive success in Guam, which has virtually lost its flying fox populations, and Samoa, where significant populations remain.     

Nestedness of remnant sonoran desert plant communities in metropolitan Phoenix, Arizona

Urbanization can have profound effects on the plant communities persisting in remnant habitats. That process can be explored by examining patterns of nestedness. Species composition for a set of communities exhibits a nested pattern if species present in progressively richer assemblages form a series of subsets. Nestedness can form as a result of the dynamic processes of extinction or colonization. It can also reflect a nested distribution of habitats among the sites or the differential abundance properties of species through passive sampling. This study investigated whether Sonoran Desert woody vegetation in remnant islands within metropolitan Phoenix is nested and explored which mechanisms are responsible for the pattern. It also examined whether vegetation is nested in similar habitat types across islands, and how species abundance relates to the nested pattern and hypothesized mechanisms. All data sets were significantly nested, indicating a nested pattern at the island and habitat levels. Community-level analyses did not indicate a primary mechanism leading to the nested pattern. Among species with abundances correlated with the nested rank-order of sites, abundance properties were significantly related to different variables. This suggests that individual taxa respond to divergent ecological mechanisms, leading to nestedness. Thus, nestedness in plant communities can result from a complex set of contributors and may not be attributable to a single factor.     

Identifying Anomalous Reports of Putatively Extinct Species and Why It Matters

Abstract:  As species become very rare and approach extinction, purported sightings can stir controversy, especially when scarce management resources are at stake. We used quantitative methods to identify reports that do not fit prior sighting patterns. We also examined the effects of including records that meet different evidentiary standards on quantitative extinction assessments for four charismatic bird species that might be extinct: Eskimo Curlew ( Numenius borealis ), Ivory-billed Woodpecker ( Campephilus principalis ), Nukupu`u ( Hemignathus lucidus ), and O`ahu `Alauahio ( Paroreomyza maculata ). For all four species the probability of there being a valid sighting today, given the past pattern of verified sightings, was estimated to be very low. The estimates of extinction dates and the chance of new sightings, however, differed considerably depending on the criteria used for data inclusion. When a historical sighting record lacked long periods without sightings, the likelihood of new sightings declined quickly with time since the last confirmed sighting. For species with this type of historical record, therefore, new reports should meet an especially high burden of proof to be acceptable. Such quantitative models could be incorporated into the International Union for Conservation of Nature's Red List criteria to set evidentiary standards required for unconfirmed sightings of "possibly extinct" species and to standardize extinction assessments across species.     

Interspecific Patterns of Genetic Diversity in Birds: Correlations with Extinction Risk

Abstract:  Birds are frequently used as indicators of ecosystem health and are the most comprehensively studied class in the animal kingdom. Nevertheless, a comprehensive, interspecific assessment of the correlates of avian genetic diversity is lacking, even though indices of genetic diversity are of considerable interest in the conservation of threatened species. We used published data on variation at microsatellite loci from 194 bird species to examine correlates of diversity, particularly with respect to conservation status and population size. We found a significant decline in mean heterozygosity with increasing extinction risk, and showed, by excluding species whose heterozygosity values were calculated with heterospecific primers, that this relationship was not dependent on ascertainment bias. Results of subsequent regression analyses suggested that smaller population sizes of threatened species were largely responsible for this relationship. Thus, bird species at risk of extinction are relatively depauperate in terms of neutral genetic diversity, which is expected to make population recovery more difficult if it reflects adaptive genetic variation. Conservation policy will need to minimize further loss of diversity if the chances of saving threatened species are to be maximized.     

Climate Change, Elevational Range Shifts, and Bird Extinctions

Abstract:  Limitations imposed on species ranges by the climatic, ecological, and physiological effects of elevation are important determinants of extinction risk. We modeled the effects of elevational limits on the extinction risk of landbirds, 87% of all bird species. Elevational limitation of range size explained 97% of the variation in the probability of being in a World Conservation Union category of extinction risk. Our model that combined elevational ranges, four Millennium Assessment habitat-loss scenarios, and an intermediate estimate of surface warming of 2.8° C, projected a best guess of 400–550 landbird extinctions, and that approximately 2150 additional species would be at risk of extinction by 2100. For Western Hemisphere landbirds, intermediate extinction estimates based on climate-induced changes in actual distributions ranged from 1.3% (1.1° C warming) to 30.0% (6.4° C warming) of these species. Worldwide, every degree of warming projected a nonlinear increase in bird extinctions of about 100–500 species. Only 21% of the species predicted to become extinct in our scenarios are currently considered threatened with extinction. Different habitat-loss and surface-warming scenarios predicted substantially different futures for landbird species. To improve the precision of climate-induced extinction estimates, there is an urgent need for high-resolution measurements of shifts in the elevational ranges of species. Given the accelerating influence of climate change on species distributions and conservation, using elevational limits in a tested, standardized, and robust manner can improve conservation assessments of terrestrial species and will help identify species that are most vulnerable to global climate change. Our climate-induced extinction estimates are broadly similar to those of bird species at risk from other factors, but these estimates largely involve different sets of species.     

Missing Mammals on Great Basin Mountains: Holocene Extinctions and Inadequate Knowledge

The discontinuous distribution of boreal mammals across Great Basin mountains has provided a significant body of data for the analysis of biogeographic patterns in isolated populations of small mammals. Such use assumes that species unreported from particular ranges do not, in fact, occur there. Data from the Roberts and Diamond mountains (central Nevada) demonstrate that this assumption is incorrect. The new distributional records from these ranges further decrease the slope of the log species—log area relationship for Great Basin boreal mammals, but increase the nestedness of the montane faunas involved. These data underscore the need for more-thorough assessments of the mammalian faunas of Great Basin mountains, but they also suggest that extinction of boreal mammals on these mountains is even more predictable than has appeared in the past.     

Response of complex food webs to realistic extinction sequences

Although an ecosystem's response to biodiversity loss depends on the order in which species are lost, the extinction sequences generally used to explore such responses in food webs have been ecologically unrealistic. We investigate how several extinction orders affect the minimum number of secondary extinctions expected within pelagic food webs from 34 temperate freshwater lakes. An ecologically plausible extinction order is derived from the geographically nested pattern of species composition among the lakes and is corroborated by species' pH tolerances. Simulations suggest that lake communities are remarkably robust to this realistic extinction order and highly sensitive to the reverse sequence of species loss. This sensitivity is not well explained by the known sensitivity of networks to the loss of highly connected species but appears to be better explained by our observation that trophic specialists preferentially consume widely distributed species at low risk of extinction. Our results highlight an important aspect of community organization that may help to maintain biodiversity amidst changing environments.     

Black Rats, Island Characteristics, and Colonial Nesting Birds in the Mediterranean: Consequences of an Ancient Introduction

Abstract: The devastation of island faunas by alien species has been instrumental in raising concerns about the global threat to biological diversity. Colonial nesting species, often restricted to islands, have been affected severely. Eradication of introduced species as a means to alleviate the problem is usually done with little or no understanding of the mechanisms governing interactions between introduced and native species. Such an understanding could help target management action. We analyzed how island area, rock substrate, bird species biology, and presence of an introduced species, the black rat (   Rattus rattus ), interact to explain the distribution and abundance of colonial nesting birds on a set of 72 islands from six archipelagos in the western Mediterranean. Rats were introduced to this region over 2000 years ago, and these communities have had time to reach an equilibrium. Using general linear models, we show that rats have affected species distributions more on the smaller islands and on islands with an acidic or neutral rock substrate; larger bird species are more resistant. On the smaller islands, where rat densities are highest, larger colonial birds are negatively affected. On larger islands, where rat densities are lower and fluctuate form year to year, larger colonial birds can maintain healthy populations despite the presence of rats. Although rats caused local extinction or reduction in bird abundance, the presence of islands varying in size and/or substrate allowed most archipelagos to retain their suite of colonial nesting bird species, despite a presumably reduced abundance for most species.     

Effects of Human Activity on Global Extinction Risk

Both natural and anthropogenic factors are important in determining a species' risk of extinction. Little work has been done, however, to quantify the magnitude of current anthropogenic influences on the extinction process. The purpose of this study is to determine the extent to which measures of the intensity of human activity are related to the global variability of two measures of species' susceptibility to extinction. We observed six indices of human activities in 90 countries, and we tested their relationships to the proportion of threatened bird and mammal species in each country, as well as to mammalian population density. After correcting for area effects, latitudinal diversity gradients, and body size (for population density), 28 to 50% of the remaining variation was statistically attributable to anthropogenic variables. Different measures of anthropogenic influence were most closely related to extinction risk in birds and mammals. Human population density was the variable most closely related to the proportion of threatened bird species per country, whereas per capita GNP was more important for mammal species. Mammalian population density strongly correlates with the extent of protected area per country. Contrary to suggestions in earlier literature, our work does not support the hypothesis that habitat loss is a prime contributor to species loss because frequencies of threatened birds and mammals are not closely related to patterns of land use.     

Testing Hypotheses of Bird Extinctions at Rio Palenque,Ecuador, with Informal Species Lists

Abstract: Informally gathered species lists are a potential source of data for conservation biology, but most remain unused because of questions of reliability and statistical issues. We applied two alternative analytical methods (contingency tests and occupancy modeling) to a 35‐year data set (1973–2007) to test hypotheses about local bird extinction. We compiled data from bird lists collected by expert amateurs and professional scientists in a 2‐km² fragment of lowland tropical forest in coastal Ecuador. We tested the effects of the following on local extinction: trophic level, sociality, foraging specialization, light tolerance, geographical range area, and biogeographic source. First we assessed extinction on the basis of the number of years in which a species was not detected on the site and used contingency tests with each factor to compare the frequency of expected and observed extinction events among different species categories. Then we defined four multiyear periods that reflected different stages of deforestation and isolation of the study site and used occupancy modeling to test extinction hypotheses singly and in combination. Both types of analyses supported the biogeographic source hypothesis and the species‐range hypothesis as causes of extinction; however, occupancy modeling indicated the model incorporating all factors except foraging specialization best fit the data.     

Threats to Avifauna on Oceanic Islands

Abstract:  Results of the study by Blackburn et al. (2004 a ) of avifauna on oceanic islands suggest that distance from the mainland and time since European colonization have major influences on species extinctions and that island area is a significant but secondary contributing factor. After augmenting the data of the study on geographical properties for some of the islands they examined, we used a causal analysis approach with structural equation modeling to reexamine their conclusions. In our model geographical properties of islands, such as island area and isolation, were considered constraints on biological factors, such as the number of introduced mammalian predators and existing number of avifauna, that can directly or indirectly influence extinction. Of the variables we tested, island area had the greatest total influence on the threat of extinction due to its direct and indirect effects on the size of island avifauna. Larger islands had both a greater number of threatened bird species and more avifauna, increasing the number of species that could become threatened with extinction. Island isolation also had a significant, positive, and direct effect on threats to island avifauna because islands farther from the mainland had fewer current extant avifauna. Time since European colonization had a significant negative, but relatively weaker, influence on threats compared with the traditional biogeographic factors of island area and distance to the mainland. We also tested the hypothesis that the amount of threat is proportionally lower on islands that have had more extinctions (i.e., there is a "filter effect"). Because the proportion of bird extinctions potentially explained only 2.3% of the variation in the proportion of threatened species on islands, our results did not support this hypothesis. Causal modeling provided a powerful tool for examining threat of extinction patterns of known and hypothesized pathways of influence.     

Extinction debt of forest plants persists for more than a century following habitat fragmentation

Following habitat fragmentation individual habitat patches may lose species over time as they pay off their "extinction debt." Species with relatively low rates of population extinction and colonization ("slow" species) may maintain extinction debts for particularly prolonged periods, but few data are available to test this prediction. We analyzed two unusually detailed data sets on forest plant distributions and land-use history from Lincolnshire, United Kingdom, and Vlaams-Brabant, Belgium, to test for an extinction debt in relation to species-specific extinction and colonization rates. Logistic regression models predicting the presence-absence of 36 plant species were first parameterized using data from Lincolnshire, where forest cover has been relatively low (approximately 5-8%) for the past 1000 years. Consistent with extinction debt theory, for relatively slow species (but not fast species) these models systematically underpredicted levels of patch occupancy in Vlaams-Brabant, where forest cover was reduced from approximately 25% to <10% between 1775 and 1900 (it is presently 6.5%). As a consequence, the ability of the Lincolnshire models to predict patch occupancy in Vlaams-Brabant was worse for slow than for fast species. Thus, more than a century after forest fragmentation reached its current level an extinction debt persists for species with low rates of population turnover.     

Food webs are built up with nested subwebs

Nested structure, in which specialists interact with subsets of species with which generalists interact, has been repeatedly found in networks of mutualistic interactions and thus is considered a general feature of mutualistic communities. However, it is uncertain how exclusive nested structure is for mutualistic communities since few studies have evaluated nestedness in other types of networks. Here, we show that 31 published food webs consist of bipartite subwebs that are as highly nested as mutualistic networks, contradicting the hypothesis that antagonistic interactions disfavor nested structure. Our findings suggest that nested networks may be a common pattern of communities that include resource-consumer interactions. In contrast to the hypothesis that nested structure enhances biodiversity in mutualistic communities, we also suggest that nested food webs increase niche overlap among consumers and thus prevent their coexistence. We discuss potential mechanisms for the emergence of nested structure in food webs and other types of ecological networks.     

Rates of Movement of Threatened Bird Species between IUCN Red List Categories and toward Extinction

Abstract:  In recent centuries bird species have been deteriorating in status and becoming extinct at a rate that may be 2–3 orders of magnitude higher than in prehuman times. We examined extinction rates of bird species designated critically endangered in 1994 and the rate at which species have moved through the IUCN (World Conservation Union) Red List categories of extinction risk globally for the period 1988–2004 and regionally in Australia from 1750 to 2000. For Australia we drew on historical accounts of the extent and condition of species habitats, spread of invasive species, and changes in sighting frequencies. These data sets permitted comparison of observed rates of movement through the IUCN Red List categories with novel predictions based on the IUCN Red List criterion E, which relates to explicit extinction probabilities determined, for example, by population viability analysis. The comparison also tested whether species listed on the basis of other criteria face a similar probability of moving to a higher threat category as those listed under criterion E. For the rate at which species moved from vulnerable to endangered, there was a good match between observations and predictions, both worldwide and in Australia. Nevertheless, species have become extinct at a rate that, although historically high, is 2 (Australia) to 10 (globally) times lower than predicted. Although the extinction probability associated with the critically endangered category may be too high, the shortfall in realized extinctions can also be attributed to the beneficial impact of conservation intervention. These efforts may have reduced the number of global extinctions from 19 to 3 and substantially slowed the extinction trajectory of 33 additional critically endangered species. Our results suggest that current conservation action benefits species on the brink of extinction, but is less targeted at or has less effect on moderately threatened species.     

Temporal patterns of foraging activity in some wood warblers in relation to the availability of insect prey

Summary I monitored the temporal pattern of diurnal feeding activity in several wood warbler (Parulidae) species and concomitantly recorded the numbers of active (flying) insects in 2 willow habitats in the western United States. At one site the temporal relationship between the density of active and inactive (nonflying) insects was investigated. The diurnal patterns of insect and bird activity were inversely related and each pattern was significantly nonuniform throughout the day; the wood warblers were largely inactive during the middle of the day when insects were most active.As foliage-gleaning birds, wood warblers depend primarily on the availability of inactive (nonflying) insects that they pick from the foliage, and they appear to be limited in their foraging activity by the unavailability of such insects during midday. Interestingly, the duration of midday inactivity for a given bird species varied inversely with the proportion of time that species spent flycatching. Thus, food availability may play an important role in determining the temporal patterns of feeding activity in these insectivorous bird species.     

Use of historical and contemporary distribution of mammals in China to inform conservation

A systematic understanding of dynamic animal extinction trajectories for different regions in a nation like China is critically important to developing practical conservation strategies. We explored historical and contemporary changes in terrestrial mammalian diversity to determine how diversity in each of the 5 regions in China has changed over time and to examine the conservation potential of these regions. We used records from databases on Pleistocene mammalian fossils and historical distribution records (1175–2020) for Primates (as a case study) to reconstruct evolutionary and historical distribution trajectories of the 11 orders of terrestrial mammals and to predict their prospective survival based on the national conservation strategy applied. The results indicated that since the Pleistocene, 4–5 mammalian orders have been lost in the northeast, 3 in central China, 2 along the coast, and 1 in the northwest. In the southwest, all 11 orders were maintained. Contemporarily, the coast and southwest had the highest and second-highest species densities. The southwest region and southeastern sections of the northwest region were the most historically and contemporarily diverse areas, which suggests that they should be the first priority for protected area (PA) designation. The central and coastal areas should be secondarily prioritized. In these 2 regions, conservation should focus on human coexistence with nature. Less attention should be paid to the PA in the northeast and western northwest because in these areas ecosystems are depauperate and the climate is harsh. Conservation in these areas should focus principally on avoiding further human encroachment on natural areas. Article impact statement: Historical and contemporary patterns of extinction can be a basis for mammalian conservation strategies.