Variability in Population Abundance and the Classification of Extinction Risk

Abstract: Classifying species according to their risk of extinction is a common practice and underpins much conservation activity. The reliability of such classifications rests on the accuracy of threat categorizations, but very little is known about the magnitude and types of errors that might be expected. The process of risk classification involves combining information from many sources, and understanding the quality of each source is critical to evaluating the overall status of the species. One common criterion used to classify extinction risk is a decline in abundance. Because abundance is a direct measure of conservation status, counts of individuals are generally the preferred method of evaluating whether populations are declining. Using the thresholds from criterion A of the International Union for Conservation of Nature (IUCN) Red List (critically endangered, decline in abundance of >80% over 10 years or 3 generations; endangered, decline in abundance of 50–80%; vulnerable, decline in abundance of 30–50%; least concern or near threatened, decline in abundance of 0–30%), we assessed 3 methods used to detect declines solely from estimates of abundance: use of just 2 estimates of abundance; use of linear regression on a time series of abundance; and use of state‐space models on a time series of abundance. We generated simulation data from empirical estimates of the typical variability in abundance and assessed the 3 methods for classification errors. The estimates of the proportion of falsely detected declines for linear regression and the state‐space models were low (maximum 3–14%), but 33–75% of small declines (30–50% over 15 years) were not detected. Ignoring uncertainty in estimates of abundance (with just 2 estimates of abundance) allowed more power to detect small declines (95%), but there was a high percentage (50%) of false detections. For all 3 methods, the proportion of declines estimated to be >80% was higher than the true proportion. Use of abundance data to detect species at risk of extinction may either fail to detect initial declines in abundance or have a high error rate.     

Conserving the abundance of nonthreatened species

Human modification of the environment is driving declines in population size and distributional extent of much of the world's biota. These declines extend to many of the most abundant and widespread species, for which proportionally small declines can result in the loss of vast numbers of individuals, biomass, and interactions. These losses could have major localized effects on ecological and cultural processes and services without elevating a species’ global extinction risk. Although most conservation effort is directed at species threatened with extinction in the very near term, the value of retaining abundance regardless of global extinction risk is justifiable based on many biodiversity or ecosystem service metrics, including cultural services, at scales from local to global. The challenges of identifying conservation priorities for widespread and abundant species include quantifying the effects of species’ abundance on services and understanding how these effects are realized as populations decline. Negative effects of population declines may be disconnected from the threat processes driving declines because of species movements and environment flows (e.g., hydrology). Conservation prioritization for these species shares greater similarity with invasive species risk assessments than extinction risk assessments because of the importance of local context and per capita effects of abundance on other species. Because conservation priorities usually focus on preventing the extinction of threatened species, the rationale and objectives for incorporating declines of nonthreatened species must be clearly articulated, going beyond extinction risk to encompass the range of likely harmful effects (e.g., secondary extinctions, loss of ecosystem services) if declines persist or are not reversed. Research should focus on characterizing the effects of local declines in species that are not threatened globally across a range of ecosystem services and quantifying the spatial distribution of these effects through the distribution of abundance. The case for conserving abundance in nonthreatened species can be made most powerfully when the costs of losing this abundance are better understood.     

Biotic Impoverishment and Homogenization in Unfragmented Forest Understory Communities

Abstract:  Ecological change is often hard to document because of a lack of reliable baseline data. Several recent then-versus-now surveys of temperate forest and grassland communities demonstrate losses of local plant species, but most are based on data from a single site. We resurveyed understory communities in 62 upland forest stands in northern Wisconsin (U.S.A.) for which quantitative baseline data exist from 50 years ago. These stands are within a largely unfragmented region but vary in species composition and successional stage. We collected data on changes in (1) total and native species richness, (2) the ratio of exotic to native species, (3) the relative abundance of habitat generalists, and (4) community similarity among sites. We also compared how these rates of change varied over time. Over the past 50 years, native species density declined an average of 18.5% at the 20-m² scale, whereas the ratio of exotic species to native species increased at 80% of all sites. Habitat generalists increased, and habitat specialists declined, accounting in part for an 8.7% rise in average similarity in species composition among sites. Most of these changes cannot be related to succession, habitat loss, or invasion by exotic species. Areas without deer hunting showed the greatest declines in native species density, with parks and research natural areas faring no better than unprotected stands. Animal-pollinated and animal-dispersed species also declined, particularly at unhunted sites. These results demonstrate the power of quantitative multistand data for assessing ecological change and identify overabundant deer as a key driver of community change. Because maintaining forest habitats alone fails to preserve plant diversity at local scales, local biotic simplification seems likely to continue in the region unless active efforts are taken to protect diversity.     

Taxonomic and Geographic Patterns of Decline for Threatened and Endangered Species in the United States

Abstract: Species listed under the U.S. Endangered Species Act (i.e., listed species) have declined to the point that the probability of their extinction is high. The decline of these species, however, may manifest itself in different ways, including reductions in geographic range, number of populations, or overall abundance. Understanding the pattern of decline can help managers assess extinction probability and define recovery objectives. Although quantitative data on changes in geographic range, number of populations, and abundance usually do not exist for listed species, more often qualitative data can be obtained. We used qualitative data in recovery plans for federally listed species to determine whether each listed species declined in range size, number of populations, or abundance relative to historical levels. We calculated the proportion of listed species in each state (or equivalent) that declined in each of those ways. Nearly all listed species declined in abundance, and range size or number of populations declined in approximately 80% of species for which those data were available. Patterns of decline, however, differed taxonomically and geographically. Declines in range were more common among vertebrates than plants, whereas population extirpations were more common among plants. Invertebrates had high incidence of range and population declines. Narrowly distributed plants and invertebrates may be subject to acute threats that may result in population extirpations, whereas vertebrates may be affected by chronic threats that reduce the extent and size of populations. Additionally, in the eastern United States and U.S. coastal areas, where the level of land conversion is high, a greater percentage of species’ ranges declined and more populations were extirpated than in other areas. Species in the Southwest, especially plants, had fewer range and population declines than other areas. Such relations may help in the selection of species’ recovery criteria.     

Analyzing Variability and the Rate of Decline of Migratory Shorebirds in Moreton Bay,Australia

Abstract: Estimating the abundance of migratory species is difficult because sources of variability differ substantially among species and populations. Recently developed state‐space models address this variability issue by directly modeling both environmental and measurement error, although their efficacy in detecting declines is relatively untested for empirical data. We applied state‐space modeling, generalized least squares (with autoregression error structure), and standard linear regression to data on abundance of wetland birds (shorebirds and terns) at Moreton Bay in southeast Queensland, Australia. There are internationally significant numbers of 8 species of waterbirds in the bay, and it is a major terminus of the large East Asian‐Australasian Flyway. In our analyses, we considered 22 migrant and 8 resident species. State‐space models identified abundances of 7 species of migrants as significantly declining and abundance of one species as significantly increasing. Declines in migrant abundance over 15 years were 43–79%. Generalized least squares with an autoregressive error structure showed abundance changes in 11 species, and standard linear regression showed abundance changes in 15 species. The higher power of the regression models meant they detected more declines, but they also were associated with a higher rate of false detections. If the declines in Moreton Bay are consistent with trends from other sites across the flyway as a whole, then a large number of species are in significant decline.     

Batrachochytrium dendrobatidis and the Collapse of Anuran Species Richness and Abundance in the Upper Manu National Park,Southeastern Peru

Abstract: Amphibians are declining worldwide, but these declines have been particularly dramatic in tropical mountains, where high endemism and vulnerability to an introduced fungal pathogen, Batrachochytrium dendrobatidis (Bd), is associated with amphibian extinctions. We surveyed frogs in the Peruvian Andes in montane forests along a steep elevational gradient (1200–3700 m). We used visual encounter surveys to sample stream‐dwelling and arboreal species and leaf‐litter plots to sample terrestrial‐breeding species. We compared species richness and abundance among the wet seasons of 1999, 2008, and 2009. Despite similar sampling effort among years, the number of species (46 in 1999) declined by 47% between 1999 and 2008 and by 38% between 1999 and 2009. When we combined the number of species we found in 2008 and 2009, the decline from 1999 was 36%. Declines of stream‐dwelling and arboreal species (a reduction in species richness of 55%) were much greater than declines of terrestrial‐breeding species (reduction of 20% in 2008 and 24% in 2009). Similarly, abundances of stream‐dwelling and arboreal frogs were lower in the combined 2008–2009 period than in 1999, whereas densities of frogs in leaf‐litter plots did not differ among survey years. These declines may be associated with the infection of frogs with Bd. B. dendrobatidis prevalence correlated significantly with the proportion of species that were absent from the 2008 and 2009 surveys along the elevational gradient. Our results suggest Bd may have arrived at the site between 1999 and 2007, which is consistent with the hypothesis that this pathogen is spreading in epidemic waves along the Andean cordilleras. Our results also indicate a rapid decline of frog species richness and abundance in our study area, a national park that contains many endemic amphibian species and is high in amphibian species richness.     

Effects of an Invasive Plant on Population Dynamics in Toads

When populations decline in response to unfavorable environmental change, the dynamics of their population growth shift. In populations that normally exhibit high levels of variation in recruitment and abundance, as do many amphibians, declines may be difficult to identify from natural fluctuations in abundance. However, the onset of declines may be evident from changes in population growth rate in sufficiently long time series of population data. With data from 23 years of study of a population of Fowler's toad (Anaxyrus [ = Bufo] fowleri) at Long Point, Ontario (1989–2011), we sought to identify such a shift in dynamics. We tested for trends in abundance to detect a change point in population dynamics and then tested among competing population models to identify associated intrinsic and extrinsic factors. The most informative models of population growth included terms for toad abundance and the extent of an invasive marsh plant, the common reed (Phragmites australis), throughout the toads’ marshland breeding areas. Our results showed density‐dependent growth in the toad population from 1989 through 2002. After 2002, however, we found progressive population decline in the toads associated with the spread of common reeds and consequent loss of toad breeding habitat. This resulted in reduced recruitment and population growth despite the lack of significant loss of adult habitat. Our results underscore the value of using long‐term time series to identify shifts in population dynamics coincident with the advent of population decline. Efectos de una Planta Invasora sobre las Dinámica Poblacional de Sapos     

Contribution of Urban Expansion and a Changing Climate to Decline of a Butterfly Fauna

Butterfly populations are naturally patchy and undergo extinctions and recolonizations. Analyses based on more than 2 decades of data on California's Central Valley butterfly fauna show a net loss in species richness through time. We analyzed 22 years of phenological and faunistic data for butterflies to investigate patterns of species richness over time. We then used 18–22 years of data on changes in regional land use and 37 years of seasonal climate data to develop an explanatory model. The model related the effects of changes in land‐use patterns, from working landscapes (farm and ranchland) to urban and suburban landscapes, and of a changing climate on butterfly species richness. Additionally, we investigated local trends in land use and climate. A decline in the area of farmland and ranchland, an increase in minimum temperatures during the summer and maximum temperatures in the fall negatively affected net species richness, whereas increased minimum temperatures in the spring and greater precipitation in the previous summer positively affected species richness. According to the model, there was a threshold between 30% and 40% working‐landscape area below which further loss of working‐landscape area had a proportionally greater effect on butterfly richness. Some of the isolated effects of a warming climate acted in opposition to affect butterfly richness. Three of the 4 climate variables that most affected richness showed systematic trends (spring and summer mean minimum and fall mean maximum temperatures). Higher spring minimum temperatures were associated with greater species richness, whereas higher summer temperatures in the previous year and lower rainfall were linked to lower richness. Patterns of land use contributed to declines in species richness (although the pattern was not linear), but the net effect of a changing climate on butterfly richness was more difficult to discern. Contribución de la Expansión Urbana y un Clima Cambiante a la Declinación de la Fauna de Mariposas     

Long-Term Population Changes of Native and Introduced Birds in the Alaka'i Swamp, Kaua'i

Abstract:  Within the last 30 years, five endemic bird species of the Alaka'i Swamp, Kaua'i, Hawai'i, have likely gone extinct. We documented population trends of the remaining avifauna in this time period to identify a common pattern in the Hawaiian Islands: decline of native species and expansion of introduced species. We conducted bird surveys over 100 km² of the Alaka'i and Kōke'e regions of Kaua'i in March–April 2000 to estimate population size, distribution, and range limits of seven native and six introduced forest birds. We compared the results with four previous surveys conducted over the last 30 years. Five of the seven native species we studied have fared well, maintaining sizeable populations (>20,000 individuals) and unchanged or increasing numbers. The endemic 'Akikiki ( Oreomystis bairdi ), however, declined from 6296 (SE ± 1374) to 1472 (SE ± 680) individuals and exhibited range contraction from 88 to 36 km². The 'I'iwi ( Vestiaria coccinea ) also experienced a decline and contraction, though not as severe. Populations of several introduced forest birds are increasing, but all species, excluding the Japanese White-eye ( Zosterops japonicus ), were at low numbers (<5,500 individuals in survey area). One introduced species, the Japanese Bush-Warbler ( Cettia diphone ) recently invaded, whereas another, the Red-billed Leiothrix ( Leiothrix lutea ), has been extirpated. Two hurricanes in the past 20 years appear to have most strongly affected nectarivores and may have contributed to the decline or extinction of several other species. Overall, native bird populations on Kaua'i have exhibited species-specific responses to limiting factors. Although most native populations appear stable, the extant native avifauna is vulnerable as a result of limited distributions and the potential for widespread habitat degradation.     

Catastrophic Decline in Mollusc Diversity in Eastern Tasmania and Its Concurrence with Shellfish Fisheries

Abstract:  We used historical patterns of deposition of mollusc shells to infer changes to inshore benthic assemblages in the southeastern Tasmanian region over the past 120 years. We identified and counted shells in slices embedded within 1m long ²¹⁰Pb-dated sediment cores were collected at 13 sites in water depths of 8–16 m. Declines in mollusc species richness and shell production occurred during the past century at all sites studied, with a mean decline per 5-cm sediment slice from 21 species in 1890 to 7 species in 1990 and in shell abundance from 150 to 30 individuals over the same period. The time course of decline notably corresponded with the history of the scallop dredge fishery, presumably either because scallop dredging caused general declines in populations of mollusc species or because other factors caused a catastrophic regional decline in molluscs that included scallops. As a consequence, the fishery was forced to close. Of major concern is that losses had not previously been recognized but extended throughout the 100-km coastal span of the study. Given that fishing and other anthropogenic impacts, as well as a lack of observational data, are virtually ubiquitous for the coastal zone, major recent losses in mollusc biodiversity may be globally widespread but have gone unnoticed.     

Using citizen science butterfly counts to predict species population trends

Citizen scientists are increasingly engaged in gathering biodiversity information, but trade‐offs are often required between public engagement goals and reliable data collection. We compared population estimates for 18 widespread butterfly species derived from the first 4 years (2011–2014) of a short‐duration citizen science project (Big Butterfly Count [BBC]) with those from long‐running, standardized monitoring data collected by experienced observers (U.K. Butterfly Monitoring Scheme [UKBMS]). BBC data are gathered during an annual 3‐week period, whereas UKBMS sampling takes place over 6 months each year. An initial comparison with UKBMS data restricted to the 3‐week BBC period revealed that species population changes were significantly correlated between the 2 sources. The short‐duration sampling season rendered BBC counts susceptible to bias caused by interannual phenological variation in the timing of species’ flight periods. The BBC counts were positively related to butterfly phenology and sampling effort. Annual estimates of species abundance and population trends predicted from models including BBC data and weather covariates as a proxy for phenology correlated significantly with those derived from UKBMS data. Overall, citizen science data obtained using a simple sampling protocol produced comparable estimates of butterfly species abundance to data collected through standardized monitoring methods. Although caution is urged in extrapolating from this U.K. study of a small number of common, conspicuous insects, we found that mass‐participation citizen science can simultaneously contribute to public engagement and biodiversity monitoring. Mass‐participation citizen science is not an adequate replacement for standardized biodiversity monitoring but may extend and complement it (e.g., through sampling different land‐use types), as well as serving to reconnect an increasingly urban human population with nature.     

Risk-Based Viable Population Monitoring

Abstract:  We describe risk-based viable population monitoring, in which the monitoring indicator is a yearly prediction of the probability that, within a given timeframe, the population abundance will decline below a prespecified level. Common abundance-based monitoring strategies usually have low power to detect declines in threatened and endangered species and are largely reactive to declines. Comparisons of the population's estimated risk of decline over time will help determine status in a more defensible manner than current monitoring methods. Monitoring risk is a more proactive approach; critical changes in the population's status are more likely to be demonstrated before a devastating decline than with abundance-based monitoring methods. In this framework, recovery is defined not as a single evaluation of long-term viability but as maintaining low risk of decline for the next several generations. Effects of errors in risk prediction techniques are mitigated through shorter prediction intervals, setting threshold abundances near current abundance, and explicitly incorporating uncertainty in risk estimates. Viable population monitoring also intrinsically adjusts monitoring effort relative to the population's true status and exhibits considerable robustness to model misspecification. We present simulations showing that risk predictions made with a simple exponential growth model can be effective monitoring indicators for population dynamics ranging from random walk to density dependence with stable, decreasing, or increasing equilibrium. In analyses of time-series data for five species, risk-based monitoring warned of future declines and demonstrated secure status more effectively than statistical tests for trend.     

Climate correlates of 20 years of trophic changes in a high-elevation riparian system

The consequences of climate change for ecosystem structure and function remain largely unknown. Here, I examine the ability of climate variation to explain long-term changes in bird and plant populations, as well as trophic interactions in a high-elevation riparian system in central Arizona, USA, based on 20 years of study. Abundances of dominant deciduous trees have declined dramatically over the 20 years, correlated with a decline in overwinter snowfall. Snowfall can affect overwinter presence of elk, whose browsing can significantly impact deciduous tree abundance. Thus, climate may affect the plant community indirectly through effects on herbivores, but may also act directly by influencing water availability for plants. Seven species of birds were found to initiate earlier breeding associated with an increase in spring temperature across years. The advance in breeding time did not affect starvation of young or clutch size. Earlier breeding also did not increase the length of the breeding season for single-brooded species, but did for multi-brooded species. Yet, none of these phenology-related changes was associated with bird population trends. Climate had much larger consequences for these seven bird species by affecting trophic levels below (plants) and above (predators) the birds. In particular, the climate-related declines in deciduous vegetation led to decreased abundance of preferred bird habitat and increased nest predation rates. In addition, summer precipitation declined over time, and drier summers also were further associated with greater nest predation in all species. The net result was local extinction and severe population declines in some previously common bird species, whereas one species increased strongly in abundance, and two species did not show clear population changes. These data indicate that climate can alter ecosystem structure and function through complex pathways that include direct and indirect effects on abundances and interactions of multiple trophic components.     

Effects of Natura 2000 on nontarget bird and butterfly species based on citizen science data

The European Union's Natura 2000 (N2000) is among the largest international networks of protected areas. One of its aims is to secure the status of a predetermined set of (targeted) bird and butterfly species. However, nontarget species may also benefit from N2000. We evaluated how the terrestrial component of this network affects the abundance of nontargeted, more common bird and butterfly species based on data from long-term volunteer-based monitoring programs in 9602 sites for birds and 2001 sites for butterflies. In almost half of the 155 bird species assessed, and particularly among woodland specialists, abundance increased (slope estimates ranged from 0.101 [SD 0.042] to 3.51 [SD 1.30]) as the proportion of landscape covered by N2000 sites increased. This positive relationship existed for 27 of the 104 butterfly species (estimates ranged from 0.382 [SD 0.163] to 4.28 [SD 0.768]), although most butterflies were generalists. For most species, when land-cover covariates were accounted for these positive relationships were not evident, meaning land cover may be a determinant of positive effects of the N2000 network. The increase in abundance as N2000 coverage increased correlated with the specialization index for birds, but not for butterflies. Although the N2000 network supports high abundance of a large spectrum of species, the low number of specialist butterflies with a positive association with the N2000 network shows the need to improve the habitat quality of N2000 sites that could harbor open-land butterfly specialists. For a better understanding of the processes involved, we advocate for standardized collection of data at N2000 sites.     

Building phenological models from presence/absence data for a butterfly fauna.

Species phenology is increasingly being used to explore the effects of climate change and other environmental stressors. Long-term monitoring data sets are essential for understanding both patterns manifest by individual species and more complex patterns evident at the community level. This study used records of 78 butterfly species observed on 626 days across 27 years at a site in northern California, USA, to build quadratic logistic regression models of the observation probability of each species for each day of the year. Daily species probabilities were summed to develop a potential aggregate species richness (PASR) model, indicating expected daily species richness. Daily positive and negative contributions to PASR were calculated, which can be used to target optimum sampling time frames. Residuals to PASR indicate a rate of decline of 0.12 species per year over the course of the study. When PASR was calculated for wet and dry years, wet years were found to delay group phenology by up to 17 days and reduce the maximum annual expected species from 32.36 to 30. Three tests to determine how well the PASR model reflected the butterfly fauna dynamics were all positive: We correlated probabilities developed with species presence/absence data to observed abundance by species, tested species' predicted phenological patterns against known biological characteristics, and compared the PASR curve to a spline-fitted curve calculated from the original species richness observations. Modeling individual species' flight windows was possible from presence/absence data, an approach that could be used on other similar records for butterfly communities with seasonal phenologies, and for common species with far fewer dates than used here. It also provided a method to assess sample frequency guidelines for other butterfly monitoring programs.     

Estimation of spatiotemporal trends in bat abundance from mortality data collected at wind turbines

Renewable energy sources, such as wind energy, are essential tools for reducing the causes of climate change, but wind turbines can pose a collision risk for bats. To date, the population-level effects of wind-related mortality have been estimated for only 1 bat species. To estimate temporal trends in bat abundance, we considered wind turbines as opportunistic sampling tools for flying bats (analogous to fishing nets), where catch per unit effort (carcass abundance per monitored turbine) is a proxy for aerial abundance of bats, after accounting for seasonal variation in activity. We used a large, standardized data set of records of bat carcasses from 594 turbines in southern Ontario, Canada, and corrected these data to account for surveyor efficiency and scavenger removal. We used Bayesian hierarchical models to estimate temporal trends in aerial abundance of bats and to explore the effect of spatial factors, including landscape features associated with bat habitat (e.g., wetlands, croplands, and forested lands), on the number of mortalities for each species. The models showed a rapid decline in the abundance of 4 species in our study area; declines in capture of carcasses over 7 years ranged from 65% (big brown bat [Eptesicus fuscus]) to 91% (silver-haired bat [Lasionycteris noctivagans]). Estimated declines were independent of the effects of mitigation (increasing wind speed at which turbines begin to generate electricity from 3.5 to 5.5 m/s), which significantly reduced but did not eliminate bat mortality. Late-summer mortality of hoary (Lasiurus cinereus), eastern red (Lasiurus borealis), and silver-haired bats was predicted by woodlot cover, and mortality of big brown bats decreased with increasing elevation. These landscape predictors of bat mortality can inform the siting of future wind energy operations. Our most important result is the apparent decline in abundance of four common species of bat in the airspace, which requires further investigation.     

Assessing conservation status with extensive but low-resolution data: Application of frequentist and Bayesian models to endangered Athabasca River rainbow trout

Use of extensive but low-resolution abundance data is common in the assessment of species at-risk status based on quantitative decline criteria under International Union for Conservation of Nature (IUCN) and national endangered species legislation. Such data can be problematic for 3 reasons. First, statistical power to reject the null hypothesis of no change is often low because of small sample size and high sampling uncertainty leading to a high frequency of type II errors. Second, range-wide assessments composed of multiple site-specific observations do not effectively weight site-specific trends into global trends. Third, uncertainty in site-specific temporal trends and relative abundance are not propagated at the appropriate spatial scale. A common result is the propensity to underestimate the magnitude of declines and therefore fail to identify the appropriate at-risk status for a species. We used 3 statistical approaches, from simple to more complex, to estimate temporal decline rates for a designatable unit (DU) of rainbow trout in the Athabasca River watershed in western Canada. This DU is considered a native species for purposes of listing because of its genetic composition characterized as >0.95 indigenous origin in the face of continuing introgressive hybridization with introduced populations in the watershed. Analysis of abundance trends from 57 time series with a fixed-effects model identified 33 sites with negative trends, but only 2 were statistically significant. By contrast, a hierarchical linear mixed model weighted by site-specific abundance provided a DU-wide decline estimate of 16.4% per year and a 3-generation decline of 93.2%. A hierarchical Bayesian mixed model yielded a similar 3-generation decline trend of 91.3% and the posterior distribution showed that the estimate had a >99% probability of exceeding thresholds for an endangered listing. We conclude that the Bayesian approach was the most useful because it provided a probabilistic statement of threshold exceedance in support of an at-risk status recommendation.     

Effects of Selective Logging on the Butterflies of a Bornean Rainforest

Abstract: Selective logging has been the main cause of disturbance to tropical forests in Southeast Asia, so the extent to which biodiversity is maintained in selectively logged forest is of prime conservation importance. We compared the butterfly assemblages of Bornean primary rainforest to those of rainforest selectively logged 6 years previously. We sampled by means of replicated transects stratified into riverine and ridge forests and we included roads in the logged forest. There was a three-fold variation in species richness and abundance over the 8-month sampling period. More species and individuals were observed in the logged forest, although between-replicate variability was high. Rarefied species richness was positively correlated with canopy openness within the range of disturbance levels encountered at our forest sites. Within families, there was no significant difference in the number of species between primary and logged forest. There was a significant difference in the relative abundance of species, but this was due largely to the abundance of one or two species. Community ordination separated the sites along a gradient of disturbance and revealed strong differences between riverine and ridge-forest butterfly assemblages in primary forest that were obscured in logged forest. There was no evidence that logging has resulted in a change in the composition of the butterfly assemblages from species with a local distribution to more widespread species. We conclude that at a logged forest site in close proximity to primary forest, low intensities of logging do not necessarily reduce the species richness or abundance of butterflies, although assemblage composition is changed.     

Negative effects of nitrogen deposition on Swiss butterflies

Nitrogen (N) deposition from agriculture and combustion of fossil fuels is a major threat to plant diversity, but its effects on organisms at higher trophic levels are unclear. We investigated how N deposition may affect species richness and abundance (number of individuals per species) in butterflies. We reviewed the peer-reviewed literature on variables used to explain spatial variation in butterfly species richness and found that vegetation variables appeared to be as important as climate and habitat variables in explaining butterfly species richness. It thus seemed likely that increased N deposition could indirectly affect butterfly communities via its influence on plant communities. To test this prediction, we analyzed data from the Swiss biodiversity monitoring program for vascular plants and butterflies in 383 study sites of 1 km² that are evenly distributed throughout Switzerland. The area has a modeled N deposition gradient of 2–44 kg N ha⁻¹ year⁻¹. We used traditional linear models and structural equation models to infer the drivers of the spatial variation in butterfly species richness across Switzerland. High N deposition was consistently linked to low butterfly diversity, suggesting a net loss of butterfly diversity through increased N deposition. We hypothesize that at low elevations, N deposition may contribute to a reduction in butterfly species richness via microclimatic cooling due to increased plant biomass. At higher elevations, negative effects of N deposition on butterfly species richness may also be mediated by reduced plant species richness. In most butterfly species, abundance was negatively related to N deposition, but the strongest negative effects were found for species of conservation concern. We conclude that in addition to factors such as intensified agriculture, habitat fragmentation, and climate change, N deposition is likely to play a key role in negatively affecting butterfly diversity and abundance.     

Minimum viable metapopulation size, extinction debt, and the conservation of a declining species.

A key question facing conservation biologists is whether declines in species' distributions are keeping pace with landscape change, or whether current distributions overestimate probabilities of future persistence. We use metapopulations of the marsh fritillary butterfly Euphydryas aurinia in the United Kingdom as a model system to test for extinction debt in a declining species. We derive parameters for a metapopulation model (incidence function model, IFM) using information from a 625-km2 landscape where habitat patch occupancy, colonization, and extinction rates for E. aurinia depend on patch connectivity, area, and quality. We then show that habitat networks in six extant metapopulations in 16-km2 squares were larger, had longer modeled persistence times (using IFM), and higher metapopulation capacity (lambdaM) than six extinct metapopulations. However, there was a > 99% chance that one or more of the six extant metapopulations would go extinct in 100 years in the absence of further habitat loss. For 11 out of 12 networks, minimum areas of habitat needed for 95% persistence of metapopulation simulations after 100 years ranged from 80 to 142 ha (approximately 5-9% of land area), depending on the spatial location of habitat. The area of habitat exceeded the estimated minimum viable metapopulation size (MVM) in only two of the six extant metapopulations, and even then by only 20%. The remaining four extant networks were expected to suffer extinction in 15-126 years. MVM was consistently estimated as approximately 5% of land area based on a sensitivity analysis of IFM parameters and was reduced only marginally (to approximately 4%) by modeling the potential impact of long-distance colonization over wider landscapes. The results suggest a widespread extinction debt among extant metapopulations of a declining species, necessitating conservation management or reserve designation even in apparent strongholds. For threatened species, metapopulation modeling is a potential means to identify landscapes near to extinction thresholds, to which conservation measures can be targeted for the best chance of success.