Extension of landscape-based population viability models to ecoregional scales for conservation planning

Landscape-based population models are potentially valuable tools in facilitating conservation planning and actions at large scales. However, such models have rarely been applied at ecoregional scales. We extended landscape-based population models to ecoregional scales for three species of concern in the Central Hardwoods Bird Conservation Region and compared model projections against long-term trend data from the North American Breeding Bird Survey. We used a spatially-explicit demographic model and structured the regional population into ecological subsections on the basis of habitat, landscape patterns, and demographic rates to assess species viability. Our model projections were within 2% of the Breeding Bird Survey trends over the last 40 years for each species. Wood thrush (Hylocichla mustelina) populations remained relatively stable over the simulation and worm-eating warbler (Helmitheros vermivorus) abundance increased throughout most of the time period until reaching carrying capacity. In contrast, the prairie warbler (Dendroica discolor) population steadily declined by 0.59% annually. The combination of habitat and demographic modeling allowed us to create models that address processes driving these populations at all scales, which is critical to understanding how regional populations respond to landscape processes such as habitat loss and fragmentation. Therefore, because it is spatially explicit and directly addresses population growth and viability, this approach provides a valuable foundation to planning conservation strategies, offering the ability to identify the most salient risks to viability and explore ways to address them.     

Demography of Asian elephants (Elephas maximus) at Uda Walawe National Park, Sri Lanka based on identified individuals

We provide estimates of population size and other demographic variables for individually-identified Asian elephants (Elephas maximus) in Uda Walawe National Park (UWNP), Sri Lanka based on systematic year-round observations. Two hundred and eighty-six adult females and 241 adult males were identified, of which four adults (2% of males) had tusks. Sightings-based demographic models showed seasonal immigration and emigration from the study area. The total population, including non-adults, was between 804 and 1160 individuals. Density ranged from 102 to 116 adult females per 100 km² and remains at this level throughout the year. This large, un-fragmented population of Asian elephants should be of high conservation priority. We find that estimates of survivorship and migration rates should be based on long sampling intervals when possible, but estimates of density and population size can still be made when observations are constrained to shorter intervals, if spatial data are available. We offer suggestions to guide census design for other elephant populations or cryptic species. We urge that other locations be systematically surveyed as well using photographic identification.     

Predicting Life-History Traits for Female New Zealand Sea Lions, <Emphasis Type="Italic">Phocarctos hookeri</Emphasis>: Integrating Short-Term Mark-Recapture Data and Population Modeling

The trade-off between survival and reproduction by individuals is central to understanding life-history parameters of a species. Few mammal species have life-history information from long-term research. Instead, demographic models are commonly utilized to investigate an individual’s life-history strategy, species dynamics, and population trends. This research investigates age-related survival and reproductive performance of adult female New Zealand (NZ) sea lions (Phocarctos hookeri), using multi-state mark-recapture data from known-age branded individuals over five years. The mark-recapture analysis was integrated with a population model to predict the lifetime reproductive output of female NZ sea lions. The integration of an analysis of short-term datasets with population modeling allows for the prediction of life-history parameters of long lived animals when long-term information is not available. While such approaches involve some caveats, it provides a framework for investigating population dynamics and is preferential to unsubstantiated assumptions. This technique can lead to better design and implementation of conservation management for long lived species.     

Population trends in a community of large Procellariiforms of Indian Ocean: Potential effects of environment and fisheries interactions

Despite the worrying conservation status of several albatross and petrel population, the long-term trends of many populations remain largely unknown and the causes of decline in many cases are known or very strongly suspected to be incidental mortality in fisheries. Here we combine long-term monitoring of population trends, breeding success and band recoveries to examine the past and current status of five species of albatrosses and giant petrels breeding at the same site: sooty albatross (Phoebetria fusca), light-mantled albatross (Phoebetria palpebrata), wandering albatross (Diomedea exulans), northern giant (Macronectes halli) and southern giant petrels (Macronectes giganteus) on Possession Island, Crozet archipelago. We identified three groups of trends over a 25-years period (1980-2005) suggesting common underlying causes for these species in relation to their bioclimatic foraging ranges. The Antarctic species - light-mantled albatross and southern giant petrel - appeared stable and increased recently, the Sub-Antarctic species - wandering albatross and northern giant petrel - declined with intermediate periods of increase, and finally the subtropical species - sooty albatross - declined all over the period. Breeding success, indicative of environmental conditions, showed two kinds of pattern (low and fluctuating versus high and/or increasing) which were consistent with oceanographic variations as found in a previous study. We present the analysis of fisheries-related recoveries, indicative of fisheries bycatch risks showing specific catch rates. No direct relationship between population trends and longline fishing effort was detected, probably because census data alone are not sufficient to capture the potentially complex response of demographic parameters of different life stages to environmental variation. This study highlights the contrasted changes of procellariiform species and the particularly worrying status of the subtropical sooty albatrosses, and in a lesser extent of Sub-Antarctic species.     

Full Open Population Capture–Recapture Models With Individual Covariates

Traditional analyses of capture–recapture data are based on likelihood functions that explicitly integrate out all missing data. We use a complete data likelihood (CDL) to show how a wide range of capture–recapture models can be easily fitted using readily available software JAGS/BUGS even when there are individual-specific time-varying covariates. The models we describe extend those that condition on first capture to include abundance parameters, or parameters related to abundance, such as population size, birth rates or lifetime. The use of a CDL means that any missing data, including uncertain individual covariates, can be included in models without the need for customized likelihood functions. This approach also facilitates modeling processes of demographic interest rather than the complexities caused by non-ignorable missing data. We illustrate using two examples, (i) open population modeling in the presence of a censored time-varying individual covariate in a full robust design, and (ii) full open population multi-state modeling in the presence of a partially observed categorical variable. Supplemental materials for this article are available online.     

Conservation biology for suites of species: Demographic modeling for Pacific island kingfishers

Conservation practitioners frequently extrapolate data from single-species investigations when managing critically endangered populations. However, few researchers initiate work with the intent of making findings useful to conservation efforts for other species. We presented and explored the concept of conducting conservation-oriented research for suites of geographically separated populations with similar natural histories, resource needs, and extinction threats. An example was provided in the form of an investigation into the population demography of endangered Micronesian kingfishers (Todiramphus cinnamominus). We provided the first demographic parameter estimates for any of the 12 endangered Pacific Todiramphus species, and used results to develop a population projection matrix model for management throughout the insular Pacific. Further, we used the model for elasticity and simulation analyses with demographic values that randomly varied across ranges that might characterize congener populations. Results from elasticity and simulation analyses indicated that changes in breeding adult survival exerted the greatest magnitude of influence on population dynamics. However, changes in nestling survival were more consistently correlated with population dynamics as demographic rates were randomly altered. We concluded that conservation practitioners working with endangered Pacific kingfishers should primarily focus efforts on factors affecting nestling and breeder survival, and secondarily address fledgling juveniles and helpers. Further, we described how the generalized base model might be changed to focus on individual populations and discussed the potential application of multi-species models to other conservation situations.     

Applying IUCN criteria to invertebrates: How red is the Red List of European butterflies?

The IUCN is the leading authority on assessing species’ extinction risks worldwide and introduced the use of quantitative criteria for the compilation of Red Lists of threatened species. Recently, we assessed the threat status of the 483 European butterfly species, using semi-quantitative data on changes in distribution and in population sizes provided by national butterfly experts. We corrected distribution trends for the observation that coarse-scale grid cells underestimate actual population trends by 35%. To account for uncertainty, we included a 5% error margin on the distribution and population trends provided. The new Red List of European butterflies determined one species as Regionally Extinct, 37 species as threatened (Critically Endangered, Endangered or Vulnerable) and a further 44 as Near Threatened. The use of semi-quantitative data on distribution and population trends permitted us to use IUCN criteria to compile a scientifically underpinned Red List of butterflies in Europe. However, a comparison of detailed monitoring data for some grassland species showed that coarse-scale grid cell data and population trends strongly underestimate extinction risks, and the list should be taken as a conservative estimate of threat. Finally, combining the new Red List status with the data provided by the national butterfly experts, allowed us to determine simple criteria to delineate conservation priorities for butterflies in Europe, so called SPecies of European conservation Concern (SPEC’s). Using European butterflies, our approach illustrated how Red Listing can be performed when data are incomplete for some IUCN criteria or vary strongly among countries.     

Potential biological removal of albatrosses and petrels with minimal demographic information

Seabirds such as albatrosses and petrels are frequently caught in longline and trawl fisheries, but limited demographic data for many species creates management challenges. A method for estimating the potential biological removal (the PBR method) for birds requires knowledge of adult survival, age at first breeding, a conservation goal, and the lower limit of a 60% confidence interval for the population size. For seabirds, usually only the number of breeding pairs is known, rather than the actual population size. This requires estimating the population size from the number of breeding pairs when important demographic variables, such as breeding success, juvenile survival, and the proportion of the adult population that engages in breeding, are unknown. In order to do this, a simple population model was built where some demographic parameters were known while others were constrained by considering plausible asymptotic estimates of the growth rate. While the median posterior population estimates are sensitive to the assumed population growth rate, the 20th percentile estimates are not. This allows the calculation of a modified PBR value that is based on the number of breeding pairs instead of the population size. For threatened albatross species, this suggests that human-caused mortalities should not exceed 1.5% of the number of breeding pairs, while for threatened petrel species, mortalities should be kept below 1.2% of the number of breeding pairs. The method is applied to 22 species and sub-species of albatrosses and petrels in New Zealand that are of management concern, of which at least 10 have suffered mortalities near or above these levels.     

Integrating demographic and genetic approaches in plant conservation

We summarize the problems that populations of formerly common plants may encounter when habitat fragmentation isolates them and reduces population size. Genetic erosion, inbreeding depression, Allee-effects on reproductive success, catastrophes and environmental stochasticity are illustrated with studies on species that have recently become rare in The Netherlands due to habitat fragmentation. These clearly indicate that population viability is negatively affected. We also show that in the recent literature (since 1980), most studies on the conservation of rare plants have addressed population genetic structure and relationships between genetic variation and population size. Though important, these studies are not suitable for assessing the importance of genetics for population viability. In turn, demographic studies can detect changes in vital rates in small populations, but cannot reveal underlying genetic causes. Fitness and demographic studies are also well-represented in the literature, but remarkably few studies have attempted to integrate empirical demographic and genetic studies. We discuss two approaches to fill this very important lacuna in our knowledge. One of these constructs matrix-projection models on the basis of demographic censuses of—if possible—large and viable populations, and combines these with the results of experiments to determine inbreeding effects on demographic transitions and, subsequently, population growth and extinction. The other approach is to demographically monitor experimentally created small and large populations with low and high genetic variation and measure their actual growth rates and probabilities of extinction. We conclude that demography and demographic-genetic experiments should play a central role in plant conservation genetics.     

Massive immigration balances high anthropogenic mortality in a stable eagle owl population: Lessons for conservation

The modern anthropized landscape is a major source of hazards for large animals such as raptors. Collisions with cables, vehicles and trains, as well as electrocution cause casualties, which may negatively impact populations. Yet, demographic studies of that impact remain scarce, which is an impediment to evidence-based conservation action. We studied the dynamics of an eagle owl (Bubo bubo) population in the northwestern Alps (Switzerland). We estimated, firstly, its demographic parameters using a Bayesian integrated population model; secondly, the frequency of different types of casualty through radio-tracking. Thirdly, we investigated the effects of reductions of human-related mortality on population trends. The breeding population was small but remained fairly stable during 20 years, suggesting that it was apparently in a good shape. However, survival probabilities of all age classes were very low (?0.61), productivity fairly good (0.93), and immigration very high (1.6 females per pair and year), indicating that the population operated as a sink. Half of the mortality was caused by infrastructure, with electrocution accounting for 24% of all fatalities. The elimination of electrocution would result in a strong population increase (17% annually). Under that scenario, immigration rate could decline by 60% and the population would still remain stable. Given that the supply of recruits from elsewhere is likely to continue, we can expect a rapid local population recovery if dangerous electric pylons are mitigated systematically. Our study demonstrates that detailed demographic analyses are necessary to diagnose problems occurring in populations and to identify efficient conservation actions.     

Variations in the age of mates as an early warning signal of changes in population trends? The case of Bonelli's eagle in Andalusia

Methods to evaluate population trends have recently received particular attention because of the decreasing patterns shown by several species since the nineteenth century. We have studied demographic traits of the Bonelli's eagle (Hieraaetus fasciatus) population of Andalusia (southern Spain) for 20 years (1980-2000). This species is endangered and is suffering a rapid decline in most of its distribution range. Although our study population has remained stable for the last 10 years, both age at first breeding and productivity have decreased during the last 5 years. We propose that age at first breeding could be used as a potential early warning signal to detect possible future changes in population trends of long-lived species with deferred maturation age. In particular, in this species spatial variation in demographic traits could help to detect whether unnatural events such as shooting are decreasing age at first breeding and productivity. If occupation of territories by non-adults is caused by human persecution, conservation efforts in the particular case of Bonelli's eagle have to be directed to protect those breeding territories where an increase in the percentage of pairs containing at least one non-adult individual would be detected.     

Usefulness of volunteer data to measure the large scale decline of “common” toad populations

Measuring a species decline is pivotal to evaluate their conservation status, but an accurate assessment of demographic trends requires observations collected across broad spatial and temporal scales. Volunteers can help to collect information over large scales, but their data may be affected by heterogeneity for sampling efforts and protocols, which may influence detection probability. Ignoring this issue may conduct to misleading conclusions. Here we show that data collected by different volunteer groups can be integrated with measures of sampling efforts, to obtain information on large scale demographic trends. We collected data on 33 common toad (Bufo bufo) populations across Italy for the period 1993–2010. We used two approaches (meta-analysis; analysis of average change in population size) to evaluate the overall demographic trend. We incorporated measures of volunteer sampling efforts into analyses, to take into account changes in detection probability. Toad abundance significantly declined in the last decade. From 2000 to 2010, 70% of populations showed a strong decline, and only 10% increased. Trends were heterogeneous among populations, but taking into account sampling effort reduced heterogeneity by 40%. We detected a 76% cumulative average decline of toad populations, despite an increasing mean sampling effort. The widespread toad decline rises concern for its future, also because the causes remain unclear. Volunteer data can be extremely useful to identify large scale population trends, if information on sampling effort are recorded and used to adjust counts.     

Conservation priorities differ at opposing species borders of a European orchid

How populations from different regions within the distribution of a species contribute to the adaptive potential and survival of that species has important implications for formulating conservation actions. We test assumptions of concepts on geographic population structure (e.g. central-marginal concept and ‘rear edge versus leading edge’ model) that could be used to inform conservation of plant species under climatic changes. We analyze a comprehensive dataset of demographic traits (e.g. population size, flowering, δ¹³C of plant leaves) of up to 32 sites of Himantoglossum hircinum (L.) Spreng. (Orchidaceae) located within six sub-regions of its European distribution range. Soil and climate parameters are employed as environmental predictors of variation in measured population traits. Climate is the main driver of demographic variability overriding central-marginal gradients that might be present. Warming of the climate at high latitudes paves the way for northward range expansion of species. Populations at the north and north-eastern range peripheries partly show exponential population growth and high genetic diversity and are likely to be the source of immigrants for colonization of newly suitable habitats as the climate continues to change. In recent times, populations at the southern range periphery have suffered from intensification of land use and decreasing rainfall, but in the case of Southern Italy are important because they contain genetically unique traits. Populations at both, ‘leading’ and ‘rear’, edges ought to be at the focus of conservation planning. Different conservation strategies are proposed at opposing species borders taking into account spatial variation in population needs on a geographic scale, projected population response to expected environmental changes and genetic characteristics.     

An Integrated Population Model From Constant Effort Bird-Ringing Data

Data from annual bird-ringing programs, in which catch effort is standardized, are routinely used to index abundance, productivity, and adult survival. Efficient models have been developed for each. Such monitoring schemes, based on ringing across a number of sites, are perhaps unique in providing this combination of demographic information and make the data particularly amenable to an integrated approach to population modeling. We develop a Bayesian approach and a deterministic population model uniting abundance, productivity, and survival. The method is applied to sedge warbler Acrocephalus schoenobaenus data from the British Trust for Ornithology’s Constant Effort Sites scheme. The possibility of “transient” birds needs to be incorporated within this analysis. We demonstrate how current methodology can efficiently be extended to use additional data from multiple within year recaptures when controlling for transience. Supplemental materials for this article are available online.     

Facilitating the evolution of resistance to avian malaria in Hawaiian birds

Research has shown that avian malaria plays an important role in limiting the distribution and population sizes of many Hawaiian birds, and that projected climate change is likely to eliminate most disease-free habitat in Hawai’i in the next century. I used a modeling approach, parameterized with demographic data from the literature and the field, to examine alternate management scenarios for the conservation of native Hawaiian birds. I examined the feasibility of using management in the form of rodent control to facilitate the evolution of resistance to malaria by increasing the survival and reproduction of native birds. Analysis of demographic data from seven native species, Akepa (Loxops coccineus), ‘Akohekohe (Palmeria dolei), Elepaio (Chasiempis sandwichensis), Hawai’i’amakihi (Hemignathus virens), Hawai’i creeper (Oreomystis mana), Omao (Myadestes obscurus), and Palila (Loxioides bailleui), suggest that differences in life history cause some species to be more susceptible to local extinctions from the transmission of malaria. Modeling results demonstrated that rodent control at middle, but not high, elevations can facilitate the evolution of resistance to malaria in several species of Hawaiian birds. Advocating a management approach that encourages evolutionary change in endangered species contrasts with the traditional conservation paradigm but it may be the best strategy to reduce the impacts of one of the multiple stressors that have devastated the native bird community of Hawai’i.     

Bayesian methods in plant conservation biology

We provide an introduction to Bayesian methods in conservation biology, illustrating inferences, prediction and decision making issues. After presenting the basic framework with a recovery plan evaluation problem, we illustrate more complex issues related to forecasting trends of structured populations using matrix population models and, finally, describe relevant topics in spatial and logistic regression problems. Computational and other implementation difficulties are also discussed.     

New Anatidae population estimates for eastern China: Implications for current flyway estimates

Soundly-based conservation plans for Anatidae require abundance and distributional data to provide accurate estimation of population sizes and trends and to identify key sites for protection. Here, we report the first ever extensive surveys throughout eastern China’s wetlands for Anatidae, 80% of which occurred in the Yangtze floodplain. Population estimates for 24 species with sufficient data are provided, allowing assessment of the accuracy of current flyway population estimates; fourteen species are far less numerous than previously believed and three far more numerous. Further improvement of the population estimates for eastern China will need coordinated counts across the region requiring a large, skilled counter network.     

The ecology of extinction: population fluctuation and decline in amphibians

Even among widespread species with high reproductive potentials and significant dispersal abilities, the probability of extinctions should be correlated both with population size variance and with the extent of population isolation. To address how variation in demographic characteristics and habitat requirements may reflect on the comparative risk of species decline, I examined 617 time series of population census data derived from 89 amphibian species using the normalized estimate of the realized rate of increase, ΔN, and its variance. Amphibians are demonstrably in general decline and exhibit a great range of dispersal abilities, demographic characteristics, and population sizes. I compared species according to life-history characteristics and habitat use. Among the populations examined, census declines outnumbered increases yet the average magnitudes for both declines and increases were not demonstrably different, substantiating findings of amphibian decline. This gives no support for the idea that amphibian population sizes are dictated by regimes featuring relatively rare years of high recruitment offset by intervening years of gradual decline such that declines may outnumber increases without negative effect. For any given population size, those populations living in large streams or in ponds had significantly higher variance than did populations of completely terrestrial or other stream-dwelling amphibians. This could not be related to life-history complexity as all the stream-breeding species examined have larvae and all of the wholly terrestrial species have direct development without a larval stage. Variance in ΔN was highest amongst the smallest populations in each comparison group. Estimated local extinction rates averaged 3.1% among pond-breeding frogs, 2.2% for pond-breeding salamanders, and negligible for both stream-breeding and terrestrial direct-developing species. Recoveries slightly exceeded extinctions among European pond-breeding frogs but not among North American pond-breeding frogs. Less common species had greater negative disparities between extinctions and recoveries. Species with highly fluctuating populations and high frequencies of local extinctions living in changeable environments, such pond- and torrent-breeding amphibians, may be especially susceptible to curtailment of dispersal and restriction of habitat.     

Effects of community-level grassland management on the non-target rare annual Agalinis auriculata

The objective of grassland management is to maintain keystone species, as well as species diversity, to promote a particular community structure, or to preserve specific ecosystem processes. Studies of grassland management rarely ascertain the effects on rare plant species, although conservation plans for rare herbaceous plants often recommend habitat management as well as restoration activities. Indeed, conservation biology has evolved from a relatively species-specific discipline into one more focused on larger scale issues such as ecosystem function, community composition and habitat restoration. Few studies have tested whether species of concern are adequately managed with a community or ecosystem-level approach. In this study, we evaluate whether community-level management of white-tailed deer and woody brush improves the viability of the rare annual plant species, Agalinis auriculata (Earleaf false foxglove; Orobanchaceae). Reducing deer browse and removing brush each positively affect plant vital rates. Brush removal increases the proportion of plants that reach the largest size class. Reducing deer browse increases the fertility of plants, particularly those in the largest size class. We report on demographic matrix models created with data from five populations of A. auriculata studied across 4 years. We find that both management activities positively affect the non-target plant species and are, in fact, necessary for A. auriculata to persist. Conservation of A. auriculata at our study sites requires both reducing the density of white-tailed deer and brush removal. Our study demonstrates that management at the community and ecosystem levels can be compatible with conservation goals at the species and population levels.     

Population viability and species interactions: Life outside the single-species vacuum

Population viability analysis (PVA) has become a widely used set of tools for evaluating relative extinction risk and prioritizing management options among imperiled populations. While PVA is a widely sanctioned tool in conservation biology, the field of population viability is in its infancy with respect to species interactions. In this paper, I review available methods for evaluating extinction risk when species interactions contribute significantly to population viability. This review includes an evaluation of six broad categories of species interactions (predation, disease, competition, mutualism, parasitism and host-parasitoid interactions) in population viability analysis, with a particular focus on predation as a case study. I first evaluated how often species interactions are considered when PVA is applied to population data from imperiled species. I identified 378 articles in commonly cited conservation journals, of which 24 attempted a viability analysis for populations threatened by interactions with other species. Most of these PVA’s treat a putative species interaction as a constant source of mortality rather than a coupled, dynamic population process. Second, I reviewed the literature to identify the availability of time-series of abundance data for two interacting species in which at least one species was threatened or endangered. Adequate time-series data were available for both species comprising an interacting pair in only 9 out of 407 papers reviewed. Third, I used a stochastic, fully stage-structured predator prey model to create time-series data (vital rates and projection matrices) in order to quantify the efficacy of two matrix-based, single-species PVA approaches. Simple single-species PVAs confound stochastic variation with population cycles induced by species interactions (in this case predation). As a result these models provide conservatively biased forecasts of viability. Unfortunately, the data needed to construct more complex PVA’s with feedback and multi-species stochasticity are rarely collected. I close with a discussion of key advances needed to “escape the population vacuum” in a move toward more realistic estimates of extinction risk.