Using a common commensal bacterium in endangered Takahe as a model to explore pathogen dynamics in isolated wildlife populations

Predicting and preventing outbreaks of infectious disease in endangered wildlife is problematic without an understanding of the biotic and abiotic factors that influence pathogen transmission and the genetic variation of microorganisms within and between these highly modified host communities. We used a common commensal bacterium, Campylobacter spp., in endangered Takahe (Porphyrio hochstetteri) populations to develop a model with which to study pathogen dynamics in isolated wildlife populations connected through ongoing translocations. Takahe are endemic to New Zealand, where their total population is approximately 230 individuals. Takahe were translocated from a single remnant wild population to multiple offshore and mainland reserves. Several fragmented subpopulations are maintained and connected through regular translocations. We tested 118 Takahe from 8 locations for fecal Campylobacter spp. via culture and DNA extraction and used PCR for species assignment. Factors relating to population connectivity and host life history were explored using multivariate analytical methods to determine associations between host variables and bacterial prevalence. The apparent prevalence of Campylobacter spp. in Takahe was 99%, one of the highest reported in avian populations. Variation in prevalence was evident among Campylobacter species identified. C. sp. nova 1 (90%) colonized the majority of Takahe tested. Prevalence of C. jejuni (38%) and C. coli (24%) was different between Takahe subpopulations, and this difference was associated with factors related to population management, captivity, rearing environment, and the presence of agricultural practices in the location in which birds were sampled. Modeling results of Campylobacter spp. in Takahe metapopulations suggest that anthropogenic management of endangered species within altered environments may have unforeseen effects on microbial exposure, carriage, and disease risk. Translocation of wildlife between locations could have unpredictable consequences including the spread of novel microbes between isolated populations.     

Spectral tuning and perceptual differences do not explain the rejection of brood parasitic eggs by American robins (Turdus migratorius)

By laying their eggs in the nests of other birds, avian brood parasites impose the cost of rearing young upon their hosts. The recognition and rejection of foreign eggs are primary host defenses against costly brood parasitism. Hosts of parasitic brown-headed cowbirds (Molothrus ater) challenge coevolutionary theory because most cowbird hosts accept parasitic eggs despite their drastically different appearance from the hosts’ own eggs. American robins (Turdus migratorius) are one of only 10 % of the over 200 potential cowbird host species to robustly reject parasitic eggs, but the mechanisms driving the sensory bases of foreign egg rejection remain elusive. Our research combined avian visual perceptual modeling and behavioral experimentation to investigate chromatic cues eliciting parasitic egg rejection in American robins. We assessed the effects of perceivable background color differences between real host and model parasite eggs, across all four avian photoreceptors, on rates of rejection of model eggs spanning in color across the entire avian spectral sensitivity range, and including immaculate model eggs matching the natural colors of robin and cowbird eggs. The results suggest that egg rejection in robins is driven by the overall perceivable difference in color between own and artificial eggs, and input from all four single-cone avian photoreceptors affects the rejection decision. The results, however, also reveal that when viewed by the avian eye, natural cowbird eggs appear more similar in background color to robin eggs than predicted by the high rejection rate of these parasitic eggs. This suggests that robins respond specifically to parasitism by cowbirds, despite an apparent lack of sensory tuning toward the detection of the background color of cowbird eggs.     

The Crumbling Infrastructure of Biodiversity: The Avian Example

The successful conservation of biodiversity depends in part upon an accurate assessment of the diversity to be preserved. This assessment is in the domain of systematics, taxonomy, and general comparative biology. Specimens play a central role in this science, and research collections thus represent the touchstone of biodiversity. The massive job of describing and understanding avian diversity is far from complete, yet the specimen basis for much-needed work is not being added to our collections; current holdings are inadequate. The dwindling influx of specimens is due primarily to opposition to collecting, which is fueled by (1) focusing conservation at the level of the individual; (2) unfamiliarity with population biology; (3) misunderstanding of scientific research; (4) typological thinking; and (5) misplaced morality. Specimen-based avian research has a long and scientifically strong history, and the benefits of this research have been extensive. Our research collections must serve as functional biological libraries. The majority of avian populations can easily withstand the relatively tiny levels of collecting required to keep this science vigorous. Insofar as avian conservation necessarily includes the preservation of a myriad of species comprising the ecosystems upon which birds rely, this problem has broad implications for the conservation of biodiversity.     

Interannual dynamics of antibody levels in naturally infected long-lived colonial birds

Little is known about the long-term persistence of specific antibodies (Ab) in natural host-parasite systems despite its potential epidemiological and ecological importance. In long-lived species, knowledge of the dynamics of individual immunological profiles can be important not only for interpreting serology results, but also for assessing transmission dynamics and the potential selective pressures acting on parasites. The aim of this paper was to investigate temporal variation in levels of specific Ab against the bacterium Borrelia burgdorferi sensu lato in adults of a long-lived colonial seabird, the Black-legged Kittiwake Rissa tridactyla. In wild populations, adults are naturally exposed each breeding season to a Borrelia vector, the tick Ixodes uriae. Breeding birds were captured during four consecutive breeding seasons, and parasite infestation quantified. Using enzyme-linked immunosorbent assay (ELISA) and immunoblots, we found that the immunological profiles of anti-Borrelia Ab were highly repeatable among years, reflecting the interannual persistence of Ab levels. We nevertheless also observed that year-to-year changes of Ab levels were related to exposure to ticks in the previous year. The long-term persistence of Ab levels may be an important mechanism of individual protection against future exposure to the microparasite. It will also affect the availability of susceptible hosts, and thus the transmission dynamics of the bacterium. These results illustrate the need to consider the dynamics of the immune response in order to better understand the evolutionary ecology of host-parasite interactions in natural populations.     

Parasite transmission in a migratory multiple host system

A transmission model was devised for trichostrongyloid nematodes of saiga antelopes and domestic sheep in Kazakhstan. The framework extends previous models by including seasonal migration of saigas, contact with separate populations of sheep, and climate-driven stochasticity in herbage biomass and in the development, survival and migration onto herbage of free-living larvae. The model was parameterised for the contrasting life histories of Marshallagia, Haemonchus and Nematodirus, three important parasites of saigas and sheep in the region, and was successful at predicting broad qualitative patterns of infection dynamics in sheep and saigas. Parasite transmission between saigas and sheep was predicted to be most important for Marshallagia (from sheep to saigas in the south in winter, and onward transmission to sheep in the north in summer) and Haemonchus (from sheep in the north in summer via saigas to sheep further south in autumn). Model predictions for winter transmission of Marshallagia infection in saigas were consistent with field data, which showed that saigas culled before they have grazed the winter range carry lower burdens of this parasite than older animals. The model provides a mechanistic explanation for its predictions, which will assist hypothesis formation, and further the epidemiological basis of efforts to control parasite transmission between wildlife and livestock in both directions. A similar modelling approach could prove useful in other situations where detailed mechanistic models of parasite transmission are inappropriate in the face of parameter uncertainty and spatio-temporal variation in climate and host density. This is likely to include the majority of wildlife-parasite systems.     

Impact of Invasive Cane Toads on Australian Birds

Abstract: The cane toad (Bufo marinus), a large, toxic, American anuran, was introduced to Australia in 1935. Populations of many of Australia's reptiles (snakes, varanid lizards, crocodiles) and carnivorous mammals (dasyurid marsupials) have declined because these predators are killed by the toad's powerful toxins. In contrast to these well‐studied species, little is known about the cane toads impacts on Australian birds. We reviewed published and unpublished data on behavioral interactions between Australian avian predators and cane toads and collated distributional and dietary information to identify avian taxa potentially at risk from cane toad invasion. Cane toads are sympatric with 172 frog‐eating bird species in Australia, and an additional 8 bird species overlap with the predicted future range of the toad. Although many bird species thus are potentially at risk, behavioral observations suggest the risk level is generally low. Despite occasional reports of Australian birds being killed when they ingest cane toads, most birds either ignore toads or survive the predation event. The apparently higher tolerance of Australian birds to toad toxins, compared with Australian reptiles and marsupials, may reflect genetic exchange between Australian birds and Asian populations that encounter other bufonid species regularly and hence have evolved the capacity to recognize or tolerate this toxic prey.     

Predation affects the susceptibility of hard clam Meretrix lusoria to Hg-stressed birnavirus

Predator–prey interaction in aquatic ecosystem is one of the simplest drivers affecting the species population dynamics. Predation controls are recognized as important aspects of ecosystem husbandry and management. In this paper we investigated how predation control cause an increase in host growth in the abundance of hard clam (Meretrix lusoria) populations subject to mercury (Hg)-stressed birnavirus. Here we linked predator–prey relationships with a bioenergetic matrix population model (MPM) associated with a susceptible–infectious–mortality (SIM) model based on a host–pathogen–predator framework to quantify the predator effects on population dynamics of disease in hard clam populations. Our results indicated that relative high predation rates could promote the hard clam abundances in relation to predators that selectively captured the infected hard clam, by which the disease transmission was suppressed. The results also demonstrated that predator-induced modifications in host behavior could have potential negative or positive effects on host growth depending on relative species density and resource dynamics. The most immediate implication of this study for the management of aquatic ecosystem is that, beyond the potential for causing a growth in abundance, predation might provoke greater predictability in aquatic ecosystem species populations and thereby increase the safety of ecosystem production from stochastic environmental events.     

Absence of Blood Parasites in Indigenous and Introduced Birds from the Cook Islands, South Pacific

Abstract: We found no protozoan parasites in 79 blood smears of birds from the Cook Islands, South Pacific. Our sample consisted of 55 smears from nine indigenous species of land and aquatic birds, and 24 smears from one introduced land bird The absence or scarcity of avian hematozoa in the Cook Islands is probably due to a very low prevalence of infection among introduced and naturally colonizing species of birds, rather than a scarcity of insect vectors. On one hand, the apparent absence or extreme scarcity of avian hematozoa, particularly the malaria-causing Plasmodium spp., is fortunate from a conservation standpoint, considering the devastating impact that Plasmodium from human-introduced mosquitoes and birds has had on the indigenous avifauna of Hawaii during the past century. On the other hand, our results suggest that the indigenous birds of the Cook Islands, much like those of Hawaii, would have little natural resistance to avian malaria should it be introduced. Thus precautions should be taken to prohibit the introduction of potentially infected nonnative birds or mosquitoes in the Cook Islands and elsewhere in Polynesia     

Joint modelling of breeding and survival in the kittiwake using frailty models

Assessment of population dynamics is central to population dynamics and conservation. In structured populations, matrix population models based on demographic data have been widely used to assess such dynamics. Although highlighted in several studies, the influence of heterogeneity among individuals in demographic parameters and of the possible correlation among these parameters has usually been ignored, mostly because of difficulties in estimating such individual-specific parameters. In the kittiwake (Rissa tridactyla), a long-lived seabird species, differences in survival and breeding probabilities among individual birds are well documented. Several approaches have been used in the animal ecology literature to establish the association between survival and breeding rates. However, most are based on observed heterogeneity between groups of individuals, an approach that seldom accounts for individual heterogeneity. Few attempts have been made to build models permitting estimation of the correlation between vital rates. For example, survival and breeding probability of individual birds were jointly modelled using logistic random effects models by [Cam, E., Link, W.A., Cooch, E.G., Monnat, J., Danchin, E., 2002. Individual covariation in life-history traits: seeing the trees despite the forest. Am. Naturalist, 159, in press]. This is the only example in wildlife animal populations we are aware of. Here we adopt the survival analysis approaches from epidemiology. We model the survival and the breeding probability jointly using a normally distributed random effect (frailty). Conditionally on this random effect, the survival time is modelled assuming a lognormal distribution, and breeding is modelled with a logistic model. Since the deaths are observed in year-intervals, we also take into account that the data are interval censored. The joint model is estimated using classic frequentist methods and also MCMC techniques in Winbugs. The association between survival and breeding attempt is quantified using the standard deviation of the random frailty parameters. We apply our joint model on a large data set of 862 birds, that was followed from 1984 to 1995 in Brittany (France). Survival is positively correlated with breeding indicating that birds with greater inclination to breed also had higher survival.     

Songbird abundance and parasitism differ between urban and rural shrublands.

Many studies have examined differences in avian community composition between urban and rural habitats, but few, if any, have looked at nesting success of urban shrubland birds in a replicated fashion while controlling for habitat. We tested factors affecting nest survival, parasitism by the Brown-headed Cowbird (Molothrus ater), and species abundance in shrubland habitat in rural and urban landscapes. We found no support for our hypothesis that nest survival was lower in urban landscapes, but strong support for the hypothesis that survival increased with nest height. We found strong support for our hypothesis that cowbird parasitism was greater in urban than rural landscapes; parasitism in urban sites was at least twice that of rural sites. We found strong support for an urban landscape effect on abundance for several species; Northern Cardinal (Cardinalis cardinalis) and Brown-headed Cowbirds were more abundant in urban landscapes, whereas Field Sparrow (Spizella pusilla) and Blue-winged Warbler (Vermivora pinus) were more abundant in rural sites. There was support for lower abundances of Blue-gray Gnatcatcher (Polioptila caerulea) and Indigo Bunting (Passerina cyanea) with increased housing density. For six other species, edge and trail density or vegetation parameters best explained abundance. Lower abundances and greater parasitism in habitat patches in urban landscapes are evidence that, for some species, these urban landscapes do not fulfill the same role as comparable habitats in rural landscapes. Regional bird conservation planning and local habitat management in urban landscapes may need to consider these effects in efforts to sustain bird populations at regional and local scales.     

Influence of host migration between woodland and pasture on the population dynamics of the tick Ixodes ricinus: A modelling approach

Ticks act as vectors of pathogens that can be harmful to animals and/or humans. Epidemiological models can be useful tools to investigate the potential effects of control strategies on diseases such as tick-borne diseases. The modelling of tick population dynamics is a prerequisite to simulating tick-borne diseases and the corresponding spread of the pathogen. We have developed a dynamic model to simulate changes in tick density at different stages (egg, larva, nymph and adult) under the influence of temperature. We have focused on the tick Ixodes ricinus, which is widespread in Europe. The main processes governing the biological cycles of ticks were taken into account: egg laying, hatching, development, host (small, mainly rodents, or large, like deer and cattle, mammals) questing, feeding and mortality. This model was first applied to a homogeneous habitat, where simulations showed the ability of the model to reproduce the general patterns of tick population dynamics. We considered thereafter a multi-habitat model, where three different habitats (woodland, ecotone and meadow) were connected through host migration. Based on this second application, it appears that migration from woodland, via the ecotone, is necessary to sustain the presence of ticks in the meadow. Woodland can therefore be considered as a source of ticks for the meadow, which in turn can be regarded as a sink. The influence of woodland on surrounding tick densities increases in line with the area of this habitat before reaching a plateau. A sensitivity analysis to parameter values was carried out and demonstrated that demographic parameters (sex ratio, development, mortality during feeding and questing, host finding) played a crucial role in the determination of questing nymph densities. This type of modelling approach provides insight into the influence of spatial heterogeneity on tick population dynamics.     

The Value of Rehabilitating Logged Rainforest for Birds

Abstract: The recent advent of carbon crediting has led to a rapid rise in biosequestration projects that seek to remove carbon from the atmosphere through afforestation and forest rehabilitation. Such projects also present an important potential opportunity to reverse biodiversity losses resulting from deforestation and forest degradation, but the biodiversity benefits of different forms of biosequestration have not been considered adequately. We captured birds in mist nets to examine the effects of rehabilitation of logged forest on birds in Sabah, Borneo, and to test the hypothesis that rehabilitation restores avian assemblages within regenerating forest to a condition closer to that seen in unlogged forest. Species richness and diversity were similar in unlogged and rehabilitated forest, but significantly lower in naturally regenerating forest. Rehabilitation resulted in a relatively rapid recovery of populations of insectivores within logged forest, especially those species that forage by sallying, but had a marked adverse effect on frugivores and possibly reduced the overall abundance of birds within regenerating forest. In view of these results, we advocate increased management for heterogeneity within rehabilitated forests, but we strongly urge an increased role for forest rehabilitation in the design and implementation of a biodiversity‐friendly carbon‐offsetting market.     

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.     

Managing the Abundance and Diversity of Breeding Bird Populations through Manipulation of Deer Populations

Abstract: Deer densities in forests of eastern North America are thought to have significant effects on the abundance and diversity of forest birds through the role deer play in structuring forest understories. We tested the ability of deer to affect forest bird populations by monitoring the density and diversity of vegetation and birds for 9 years at eight 4-ha sites in northern Virginia, four of which were fenced to exclude deer. Both the density and diversity of understory woody plants increased following deer exclosure. The numerical response of the shrubs to deer exclosure was significantly predicted by the soil quality (ratio of organic carbon to nitrogen) at the sites. Bird populations as a whole increased following exclosure of deer, particularly for ground and intermediate canopy species. The diversity of birds did not increase significantly following exclosure of deer, however, primarily because of replacement of species as understory vegetation proceeded through successional processes. Changes in understory vegetation accounted for most of the variability seen in the abundance and diversity of bird populations. Populations of deer in protected areas are capable of causing significant shifts in the composition and abundance of bird communities. These shifts can be reversed by increasing the density and diversity of understory vegetation, which can be brought about by reducing deer density.     

Potential Misuse of Avian Density as a Conservation Metric

Abstract: Effective conservation metrics are needed to evaluate the success of management in a rapidly changing world. Reproductive rates and densities of breeding birds (as a surrogate for reproductive rate) have been used to indicate the quality of avian breeding habitat, but the underlying assumptions of these metrics rarely have been examined. When birds are attracted to breeding areas in part by the presence of conspecifics and when breeding in groups influences predation rates, the effectiveness of density and reproductive rate as indicators of habitat quality is reduced. It is beneficial to clearly distinguish between individual‐ and population‐level processes when evaluating habitat quality. We use the term reproductive rate to refer to both levels and further distinguish among levels by using the terms per capita fecundity (number of female offspring per female per year, individual level) and population growth rate (the product of density and per capita fecundity, population level). We predicted how density and reproductive rate interact over time under density‐independent and density‐dependent scenarios, assuming the ideal free distribution model of how birds settle in breeding habitats. We predicted population density of small populations would be correlated positively with both per capita fecundity and population growth rate due to the Allee effect. For populations in the density‐dependent growth phase, we predicted no relation between density and per capita fecundity (because individuals in all patches will equilibrate to the same success rate) and a positive relation between density and population growth rate. Several ecological theories collectively suggest that positive correlations between density and per capita fecundity would be difficult to detect. We constructed a decision tree to guide interpretation of positive, neutral, nonlinear, and negative relations between density and reproductive rates at individual and population levels.     

Demographic Responses by Birds to Forest Fragmentation

Abstract:  Despite intensive recent research on the effects of habitat loss and fragmentation on bird populations, our understanding of underlying demographic causes of population declines is limited. We reviewed avian demography in relation to habitat fragmentation. Then, through a meta-analysis, we compared specific demographic responses by forest birds to habitat fragmentation, providing a general perspective of factors that make some species and populations more vulnerable to fragmentation than others. We obtained data from the scientific literature on dispersal, survival, fecundity, and nesting success of birds. Birds were divided into subgroups on the basis of region, nest site, biogeographical history, and migration strategy. Species most sensitive to fragmentation were ground- or open-nesters nesting in shrubs or trees. Residents were equally sensitive to fragmentation in the Nearctic and Palearctic regions, but Nearctic migrants were more sensitive than Palearctic migrants. Old World species were less sensitive than New World species, which was predicted based on the history of forest fragmentation on these two continents. Pairing success was the variable most associated with fragmentation, suggesting an important role of dispersal. Fledgling number or condition, timing of nesting, and clutch size were not associated with sensitivity to fragmentation, suggesting that negative fragmentation effects on birds do not generally result from diminished food resources with increasing level of fragmentation. Future studies on demographic responses of birds to habitat fragmentation would be more effective if based on a combination of measures that can distinguish among the demographic mechanisms underlying population changes related to habitat fragmentation.     

Making the Leap from Researcher to Planner: Lessons from Avian Conservation Planning in the Dominican Republic

Abstract: As research biologists move into conservation biology, especially in foreign countries, providing support for conservation planning efforts presents unique challenges. Published accounts of national, multidisciplinary planning efforts and priority setting for avian conservation are not common. I describe the process and results of a broad-based, grassroots-oriented avian conservation planning workshop held in the Dominican Republic in which we designed a coordinated strategy for avian conservation in the country. The planning process sought to (1) increase communication and cooperation among conservationists; (2) familiarize participants with resources pertinent to avian conservation; (3) encourage the transfer of information between researchers and managers; (4) promote the concepts of long-term avian monitoring, avian conservation plans, and species management plans; and (5) develop a common, multidisciplinary strategy to promote the conservation of birds in the Dominican Republic. The workshop highlighted group discussions among research biologists, managers, educators, and public policy specialists to assess avian conservation needs and priorities with respect to each discipline and has since galvanized a significant portion of the conservation community around several cooperative projects involving diverse segments of the community. Avian biologists can play a significant role in conservation efforts through a willingness to work with key players in diverse fields and to envision holistic, multidisciplinary approaches to conservation issues.     

Emerging prion disease drives host selection in a wildlife population

Infectious diseases are increasingly recognized as an important force driving population dynamics, conservation biology, and natural selection in wildlife populations. Infectious agents have been implicated in the decline of small or endangered populations and may act to constrain population size, distribution, growth rates, or migration patterns. Further, diseases may provide selective pressures that shape the genetic diversity of populations or species. Thus, understanding disease dynamics and selective pressures from pathogens is crucial to understanding population processes, managing wildlife diseases, and conserving biological diversity. There is ample evidence that variation in the prion protein gene (PRNP) impacts host susceptibility to prion diseases. Still, little is known about how genetic differences might influence natural selection within wildlife populations. Here we link genetic variation with differential susceptibility of white-tailed deer to chronic wasting disease (CWD), with implications for fitness and disease-driven genetic selection. We developed a single nucleotide polymorphism (SNP) assay to efficiently genotype deer at the locus of interest (in the 96th codon of the PRNP gene). Then, using a Bayesian modeling approach, we found that the more susceptible genotype had over four times greater risk of CWD infection; and, once infected, deer with the resistant genotype survived 49% longer (8.25 more months). We used these epidemiological parameters in a multi-stage population matrix model to evaluate relative fitness based on genotype-specific population growth rates. The differences in disease infection and mortality rates allowed genetically resistant deer to achieve higher population growth and obtain a long-term fitness advantage, which translated into a selection coefficient of over 1% favoring the CWD-resistant genotype. This selective pressure suggests that the resistant allele could become dominant in the population within an evolutionarily short time frame. Our work provides a rare example of a quantifiable disease-driven selection process in a wildlife population, demonstrating the potential for infectious diseases to alter host populations. This will have direct bearing on the epidemiology, dynamics, and future trends in CWD transmission and spread. Understanding genotype-specific epidemiology will improve predictive models and inform management strategies for CWD-affected cervid populations.     

Area-based assessment of extinction risk

Underpinning the International Union for Conservation of Nature (IUCN) Red List is the assessment of extinction risk as determined by the size and degree of loss of populations. The IUCN system lists a species as Critically Endangered, Endangered, or Vulnerable if its population size declines 80%, 50%, or 30% within a given time frame. However, effective implementation of the system faces substantial challenges and uncertainty because geographic scale data on population size and long-term dynamics are scarce. I develop a model to quantify extinction risk using a measure based on a species' distribution, a much more readily obtained quantity. The model calculates the loss of the area of occupancy that is equivalent to the loss of a given proportion of a population. It is a very simple yet general model that has no free parameters and is independent of scale. The model predicted well the distributions of 302 tree species at a local scale and the distributions of 348 species of North American land birds. This area-based model provides a solution to the long-standing problem for IUCN assessments of lack of data on population sizes, and thus it will contribute to facilitating the quantification of extinction risk worldwide.     

Dissecting the drivers of population cycles: Interactions between parasites and mountain hare demography

There is a growing awareness that cyclic population dynamics in vertebrate species are driven by a complex set of interactions rather than a single causal factor. While theory suggests that direct host-parasite interactions may destabilise population dynamics, the interaction between host and parasite may also influence population dynamics through indirect effects that result in delayed responses to either density or to life-history traits. Using empirical data on mountain hares (Lepus timidus) infected with a nematode parasite (Trichostrongylus retortaeformis), we developed an individual-based model (IBM) that incorporated direct effects and delayed life-history effects (DLHEs) of a macroparasite, alternative transmission mechanisms and seasonality in host population dynamics. The full model describes mean characteristics of observed mountain hare time series and parasite abundance, but by systematically removing model structure we dissect out dynamic influences of DLHEs. The DLHEs were weakly destabilising, increasing the propensity for cyclic dynamics and suggesting DLHEs could be important processes in host-parasite systems. Further, by modifying model structure we identify a strong influence of parasite transmission mechanism on host population stability, and discuss the implications for parasite aggregation mechanisms, host movement and natural geographical variation in host population dynamics. The effect of T. retortaeformis on mountain hares likely forms part of a complex set of interactions that lead to population cycles.