Diagnosing Mechanisms of Decline and Planning for Recovery of an Endangered Brown Bear (Ursus arctos) Population

Background

The usual paradigm for translocations is that they should not take place in declining populations until the causes(s) of the decline has been reversed. This approach sounds intuitive, but may not apply in cases where population decline is caused by behavioral or demographic mechanisms that could only be reversed by translocation itself.

Methodology/Principal Findings

We analyzed a decade of field data for Pyrenean brown bears (Ursus arctos) from two small populations: the growing Central population - created from a previous translocation and the endemic Western population - believed to be declining because of excessive human-caused mortality. We found that adult survival rates for both populations were as high as those observed for most other protected brown bear populations. However, the Western population had much lower reproductive success than the Central population. Adult breeding sex ratio was male-biased in the Western population and female-biased in the Central population. Our results exclude high anthropogenic mortality as a cause for population decline in the West but support low reproductive success, which could result from sexually selected infanticide induced by a male-biased adult sex ratio or inbreeding depression. Using a stochastic demographic model to compute how many bears should be released to ensure viability, we show that the Western population could recover provided adequate numbers of new females are translocated.

Conclusions/Significance

We suggest that a translocation could take place, even if the decline has not yet been reversed, if the translocation itself removes the biological mechanisms behind the decline. In our case, the ultimate cause of low reproductive success remained unknown (infanticide or inbreeding), but our proposed translocation strategies should eliminate the proximate cause (low reproductive success) of the decline and ensure population recovery and viability.     

A data-driven demographic model to explore the decline of the Bathurst caribou herd

The Bathurst herd of barren-ground caribou (Rangifer tarandus groenlandicus) in the Canadian central arctic declined from an estimated 203,800 to 16,400 breeding females from 1986 to 2009, with the most rapid decline from 2006 to 2009. A key research and management question was whether the decline was mainly due to decreases in productivity alone or also due to reduced adult female survival. Investigating causes of the decline was hampered by a lack of direct estimates of caribou demographic parameters. We developed a demographic model that could be objectively fitted to field data to explore the mechanisms for the Bathurst decline, with a focus on the recent accelerated decline from 2006 to 2009. Our modeling indicated that the decline was driven by increasing negative trends in adult female and calf survival rates and possibly reduced fecundity The effect of a constant hunter harvest on the declining herd was one potential cause for the recent accelerated decline in adult survival. The demographic model detected negative trends in adult female survival that were not detected using standalone analyses of collar-based survival data. The model allowed rigorous interpretation of trends in productivity by controlling for the simultaneous influence of trends in adult, calf, and yearling survival and adult fecundity on field-based calf–cow ratios. Stochastic simulations suggested that large increases in adult survival and productivity would be needed for the herd to recover. Our methods enable objective modeling of caribou demography that can assist in caribou management based upon all sources of available data. © 2011 The Wildlife Society.     

Population decline in tree swallows ( Tachycineta bicolor ) linked to climate change and inclement weather on the breeding ground

Population decline and the threat of extinction are realities currently facing many species. Yet, in most cases, the detailed demographic data necessary to identify causes of population decline are unavailable. Using 43 years (1975−2017) of data from a box-nesting population of tree swallows (Tachycineta bicolor), we identified reduced survival of offspring as a probable demographic cause of population decline. Poor fledging success was associated with increased predation and poor weather conditions during early nestling development. Low juvenile survival and subsequent recruitment was linked to poor weather conditions during the post-fledging period and may also be linked to conditions on the wintering grounds. Regional weather conditions during critical stages of breeding (early nestling and post-fledging) have become progressively worse over the 43-year study period. None of the other factors linked to offspring survival have similarly deteriorated. Overall, our results suggest tree swallows should be added to the growing list of species challenged by climate change, and that other species of aerial insect specialists may face similar impacts of climate change.     

Characterising demographic contributions to observed population change in a declining migrant bird

Populations of Afro‐Palearctic migrant birds have shown severe declines in recent decades. To identify the causes of these declines, accurate measures of both demographic rates (seasonal productivity, apparent survival, immigration) and environmental parameters will allow conservation and research actions to be targeted effectively. We used detailed observations of marked breeding birds from a ‘stronghold’ population of whinchats Saxicola rubetra in England (stable against the declining European trend) to reveal both on‐site and external mechanisms that contribute to population change. From field data, a population model was developed based on demographic rates from 2011 to 2014. Observed population trends were compared to the predicted population trends to assess model‐accuracy and the influence of outside factors, such as immigration. The sensitivity of the projected population growth rate to relative change in each demographic rate was also explored. Against expectations of high productivity, we identified low seasonal breeding success due to nocturnal predation and low apparent first‐year survival, which led to a projected population growth rate (λ) of 0.818, indicating a declining trend. However, this trend was not reflected in the census counts, suggesting that high immigration was probably responsible for buffering against this decline. Elasticity analysis indicated λ was most sensitive to changes in adult survival but with covariance between demographic rates accounted for, most temporal variation in λ was due to variation in productivity. Our study demonstrates that high quality breeding habitat can buffer against population decline but high immigration and low productivity will expose even such stronghold populations to potential decline or abandonment if either factor is unsustainable. First‐year survival also appeared low, however this result is potentially confounded by high natal dispersal. First‐year survival and/or dispersal remains a significant knowledge gap that potentially undermines local solutions aimed at counteracting low productivity.     

The demography of an increasing insular Eurasian Scops Owl (Otus scops) population in western France

Rapid population declines of many long-distance Afro-Palaearctic migratory bird species are ongoing across Europe but the demographic drivers are often poorly understood, thereby limiting the development of appropriate conservation actions. Using long-term population monitoring (39 years), capture–mark–recapture data and a matrix model, we estimated demographic parameters and the effect of climate variables on adult survival, and modelled the dynamics of an increasing population of Eurasian Scops Owls Otus scops in a landscape with agricultural abandonment in western France. The observed mean annual population growth rate was 1.055 (from 68 to 523 territorial males between 1981 and 2019). Over the study period, clutch size and hatching success were stable, but fledging success and breeding success showed slight negative trends, probably due to density-dependence. Survival varied with age, with an increase during early life and evidence for rapid senescence from 4 years old. Adult survival remained stable and was positively linked to the amount of autumn rainfall in the Sahel and to the winter North Atlantic Oscillation. Survival of younger age-classes made the largest contribution to the variance of the population growth rate, followed by clutch size, fledging success and survival of older birds. Such a long-term population increase in a landscape where intensive agriculture has decreased by 64.6% sheds some new light on the causes of the decline of European Scops Owl and other Afro-Palaearctic bird populations. We infer some of the possible causes of this large-scale decline, in particular food shortage, and discuss conservation measures that could be applicable to reverse this trend.     

Pup Mortality in a Rapidly Declining Harbour Seal (Phoca vitulina) Population

The harbour seal population in Orkney, off the north coast of Scotland, has reduced by 65% between 2001 and 2010. The cause(s) of this decline are unknown but must affect the demographic parameters of the population. Here, satellite telemetry data were used to test the hypothesis that increased pup mortality could be a primary driver of the decline in Orkney. Pup mortality and tag failure parameters were estimated from the duration of operation of satellite tags deployed on harbour seal pups from the Orkney population (n = 24) and from another population on the west coast of Scotland (n = 24) where abundance was stable. Survival probabilities from both populations were best represented by a common gamma distribution and were not different from one another, suggesting that increased pup mortality is unlikely to be the primary agent in the Orkney population decline. The estimated probability of surviving to 6 months was 0.390 (95% CI 0.297 – 0.648) and tag failure was represented by a Gaussian distribution, with estimated mean 270 (95% CI = 198 – 288) and s.d. 21 (95% CI = 1 – 66) days. These results suggest that adult survival is the most likely proximate cause of the decline. They also demonstrate a novel technique for attaining age-specific mortality rates from telemetry data.     

Small-scale spatial and temporal variation in the demographic processes underlying the large-scale decline of eiders in the Baltic Sea

The application of uniform conservation schemes often fails to account for small-scale spatial variation in the drivers of population decline. Demographic comparisons of imperilled populations across locations are therefore crucial for successful conservation, but progress is hampered by lack of long-term data from more than a single population. The recent large-scale decline of eider ducks (Somateria mollissima) in the Baltic Sea is ideal for determining to what extent mechanisms underlying population decline can be extrapolated over larger areas. We utilized stochastic demographic methods incorporating both environmental and sampling variation to assess small-scale spatial and temporal variation in the population dynamics of eiders at Söderskär (eastern range-margin) and Tvärminne (core breeding area), situated 130 km apart. The stochastic growth rate models accurately predicted the observed differences in the rate of decline between sites and time periods. At Söderskär, established breeder survival had by far the greatest elasticity, whereas elasticity was more evenly distributed among vital rates at Tvärminne. Although the study sites showed the single largest difference in fecundity, stochastic life table response experiment analyses revealed that reduced adult female survival at Tvärminne mainly determined the observed difference in growth rates between sites. In contrast, reduced fecundity primarily differentiated the past population increase from the present population decline at Söderskär. Our results demonstrate that different mechanisms may be associated with population decline across adjacent geographic locations, and indicate that dispersal of first-time breeders may be important for population dynamics. Safeguarding adult female survival and/or fecundity should be prioritized in management efforts.     

Early reproductive investment,senescence and lifetime reproductive success in female Asian elephants

The evolutionary theory of senescence posits that as the probability of extrinsic mortality increases with age, selection should favour early‐life over late‐life reproduction. Studies on natural vertebrate populations show early reproduction may impair later‐life performance, but the consequences for lifetime fitness have rarely been determined, and little is known of whether similar patterns apply to mammals which typically live for several decades. We used a longitudinal dataset on Asian elephants (Elephas maximus) to investigate associations between early‐life reproduction and female age‐specific survival, fecundity and offspring survival to independence, as well as lifetime breeding success (lifetime number of calves produced). Females showed low fecundity following sexual maturity, followed by a rapid increase to a peak at age 19 and a subsequent decline. High early life reproductive output (before the peak of performance) was positively associated with subsequent age‐specific fecundity and offspring survival, but significantly impaired a female's own later‐life survival. Despite the negative effects of early reproduction on late‐life survival, early reproduction is under positive selection through a positive association with lifetime breeding success. Our results suggest a trade‐off between early reproduction and later survival which is maintained by strong selection for high early fecundity, and thus support the prediction from life history theory that high investment in reproductive success in early life is favoured by selection through lifetime fitness despite costs to later‐life survival. That maternal survival in elephants depends on previous reproductive investment also has implications for the success of (semi‐)captive breeding programmes of this endangered species.     

Effects of Climate Change and Fisheries Bycatch on Shy Albatross (Thalassarche cauta) in Southern Australia

The impacts of climate change on marine species are often compounded by other stressors that make direct attribution and prediction difficult. Shy albatrosses (Thalassarche cauta) breeding on Albatross Island, Tasmania, show an unusually restricted foraging range, allowing easier discrimination between the influence of non-climate stressors (fisheries bycatch) and environmental variation. Local environmental conditions (rainfall, air temperature, and sea-surface height, an indicator of upwelling) during the vulnerable chick-rearing stage, have been correlated with breeding success of shy albatrosses. We use an age-, stage- and sex-structured population model to explore potential relationships between local environmental factors and albatross breeding success while accounting for fisheries bycatch by trawl and longline fisheries. The model uses time-series of observed breeding population counts, breeding success, adult and juvenile survival rates and a bycatch mortality observation for trawl fishing to estimate fisheries catchability, environmental influence, natural mortality rate, density dependence, and productivity. Observed at-sea distributions for adult and juvenile birds were coupled with reported fishing effort to estimate vulnerability to incidental bycatch. The inclusion of rainfall, temperature and sea-surface height as explanatory variables for annual chick mortality rate was statistically significant. Global climate models predict little change in future local average rainfall, however, increases are forecast in both temperatures and upwelling, which are predicted to have detrimental and beneficial effects, respectively, on breeding success. The model shows that mitigation of at least 50% of present bycatch is required to offset losses due to future temperature changes, even if upwelling increases substantially. Our results highlight the benefits of using an integrated modeling approach, which uses available demographic as well as environmental data within a single estimation framework, to provide future predictions. Such predictions inform the development of management options in the face of climate change.     

Negative population trend for Chapada Flycatchers (Suiriri islerorum) despite high apparent annual survival

ABSTRACT Population size and trends are demographic parameters that can be used to help determine the threat of extirpation or extinction of bird populations and in determining management strategies. Chapada Flycatchers (Suiriri islerorum) are endemic to the Cerrado region of Brazil, dependent on open cerrado habitat, defend large territories, and their populations seem to be declining. From 2003 to 2007, we analyzed demographic parameters of a declining population of Chapada Flycatchers, using the rate at which territories became vacant and the size of the breeding population to ascertain the status of the population, and evaluated the relative contributions of apparent annual survival and recruitment rates to these trends. Territories became vacant at a mean rate of 13% per year, and 32% of all territories were vacated during our study. The current breeding population (20 pairs) was at least seven times smaller than the estimated carrying capacity of the reserve (141 territorial pairs). Estimated apparent annual survival probabilities (0.77 for breeders; 0.67 for nonbreeders) based on model averaging were comparable to those reported for other Neotropical passerines. Survival rates did not differ between the sexes and the estimated recruitment rate was 0.21. In many species, adult survival is the factor that most strongly influences population growth rates. However, the population of Chapada Flycatchers we studied is declining despite high annual survival rates, with low and variable breeding success probably causing the decline. Our results improve our understanding of the possible role of adult survival and breeding success in the decline of populations of small passerines in isolated reserves in the tropics.     

The decline of the Spotted Flycatcher Muscicapa striata in the UK: an integrated population model

The Spotted Flycatcher has become a species of great conservation concern in Europe and the UK following a period of prolonged and accelerating decline. We consider a range of population models for UK census data between 1965 and 1996, along with independent survey data designed to estimate demographic parameters. Population declines are similar in the major habitats and regions of the UK, indicating that a broad-scale factor has influenced the population. Nest record data show that changes in individual nest success are not the cause of the increasing decline. This is more likely the result of a fall in the survival rates of fledged birds prior to the next breeding season; we show by comparison with the recoveries of ringed birds that the most likely cause is an increased mortality rate of birds either shortly after fledging or within the subsequent first year of life.     

Changing demography and population decline in the Common Starling Sturnus vulgaris: a multisite approach to Integrated Population Monitoring

Understanding demography is critical for understanding the causes underlying population declines, and for initiating and monitoring policies to reverse them. A method of fitting demographic models directly to avian count data recorded at a sample of census sites is described. The model is applied to national and regional counts of Common Starling Sturnus vulgaris in Britain for the period 1965–2000. Starlings have declined markedly during this time and are now on the list of birds of highest conservation concern. Although there have been small changes in various aspects of breeding success over this time, the analysis shows that these are unlikely to have made much impact on the long-term status of the species. By incorporating estimates of survival, based upon recoveries of ringed birds, the model suggests that the pronounced national decline in Starling numbers since a period of stability in the 1970s is most likely to be due to changes in the survival of first-year birds. This narrows the candidate range of environmental factors which might influence the decline and implicates changes in resource availability.     

An integrated population model sheds light on the complex population dynamics of a unique colonial breeder

Climate models forecast increasing climatic variation and more extreme events, which could increase the variability in animal demographic rates. More variable demographic rates generally lead to lower population growth and can be detrimental to wild populations, especially if the particular demographic rates affected are those to which population growth is most sensitive. We investigated the population dynamics of a metapopulation of 25 colonies of a semi-arid bird species, the sociable weaver Philetairus socius, and how it was influenced by seasonal weather during 1993–2014. We constructed an integrated population model which estimated population sizes similar to observed population counts, and allowed us to estimate annual fecundity and recruitment. Variance in fecundity contributed most to variance in population growth, which showed no trend over time. No weather variables explained overall demographic variation at the population level. However, a separate analysis of the largest colony showed a clear decline with a high extinction probability (0.05 to 0.33) within 5 years after the study period. In this colony, juvenile survival was lower when summers were hot, and adult survival was lower when winters were cold. Rainfall was also negatively correlated with adult survival. These weather effects could be due to increased physiological demands of thermoregulation and rainfall-induced breeding activity. Our results suggest that the dynamics of the population on the whole are buffered against current weather variation, as individual colonies apparently react in different ways. However, if more and increasingly extreme weather events synchronize colony dynamics, they are likely to have negative effects.     

Short- and long-term population dynamical consequences of asymmetric climate change in black grouse

Temporal asymmetry in patterns of regional climate change may jeopardize the match between the proximate and ultimate cues of the timing of breeding. The consequences on short- and long-term population dynamics and trends as well as the underlying mechanisms are, however, often unknown. Using long-term data from Finland, we demonstrate that black grouse (Tetrao tetrix) have responded to spring warming by advancing both egg-laying and hatching. However, early summer (the time of hatching) has not advanced, and chicks have to face colder post-hatching conditions. Demonstrating that these conditions are critical to post-hatching survival, we show that chicks are increasingly suffering higher mortality because they hatch too early. Consequently, breeding success and population size has severely declined over the past four decades. Finally, we modelled the impact of this particular climate change scenario on population dynamics and show that the mismatch can further explain the observed collapse of cyclic fluctuations. Because the evolutionary response of grouse is lagging behind the novel selective pressures, seasonally asymmetric climate change is likely to constitute an important determinant of future short- and long-term changes in the dynamics of black grouse populations.     

Modelling urban populations of the Australian White Ibis (Threskiornis molucca) to inform management

Since the 1970s, populations of the Australian White Ibis (Threskiornis molucca) have dramatically increased in many Australian urban centres. Managers of ibis are currently focusing on limiting this bird's reproductive success in order to reduce population sizes or at least halt further increases in urban areas. Here we use data on nesting success and survival for three populations of ibis around greater Sydney to develop an age-structured population model. The estimated growth rate for all populations combined was about 1.5 % per year and for individual sites were more variable at −1, −7, and 9 %. For all populations, growth rates were most sensitive (based on elasticity analyses) to the survival of adults and least sensitive to fecundity, especially of 3 year olds. Further exploration of the importance of fecundity rates, which are relatively poorly known for these populations, suggests that rates of <0.4 fledglings per nest per year is very likely to lead to a population decline (λ less than lower bound of 95 % CI). Conversely, positive population growth is nearly assured (λ greater than upper bound of 95 % CI) for fecundities of >0.7 fledgling per nest per year. The results suggest that ibis from other locations (probably their traditional breeding areas in inland Australia) have immigrated into urban environments as estimated growth rates cannot account for current population sizes. Management strategies must take these findings into account and also consider that ibis are declining in their traditional habitats to avoid exacerbating their decline at a regional scale.     

Senescence and carryover effects of reproductive performance influence migration,condition, and breeding propensity in a small shorebird

Breeding propensity, the probability that an animal will attempt to breed each year, is perhaps the least understood demographic process influencing annual fecundity. Breeding propensity is ecologically complex, as associations among a variety of intrinsic and extrinsic factors may interact to affect an animal's breeding decisions. Individuals that opt not to breed can be more difficult to detect than breeders, which can (1) lead to difficulty in estimation of breeding propensity, and (2) bias other demographic parameters. We studied the effects of sex, age, and population reproductive success on the survival and breeding propensity of a migratory shorebird, the piping plover (Charadrius melodus ), nesting on the Missouri River. We used a robust design Barker model to estimate true survival and breeding propensity and found survival decreased as birds aged and did so more quickly for males than females. Monthly survival during the breeding season was lower than during migration or the nonbreeding season. Males were less likely to skip breeding (range: 1–17%) than females (range: 3–26%; β_sex = ?0.21, 95% CI: ?0.38 to ?0.21), and both sexes were less likely to return to the breeding grounds following a year of high reproductive success. Birds that returned in a year following relatively high population‐wide reproductive output were in poorer condition than following a year with lower reproductive output. Younger adult birds and females were more likely to migrate from the breeding area earlier than older birds and males; however, all birds stayed on the breeding grounds longer when nest survival was low, presumably because of renesting attempts. Piping plovers used a variety of environmental and demographic cues to inform their reproduction, employing strategies that could maximize fitness on average. Our results support the “disposable soma” theory of aging and follow with predictions from life history theory, exhibiting the intimate connections among the core ecological concepts of senescence, carryover effects, and life history.     

Demographic consequences of heavy metals and persistent organic pollutants in a vulnerable long-lived bird,the wandering albatross

Seabirds are top predators of the marine environment that accumulate contaminants over a long life-span. Chronic exposure to pollutants is thought to compromise survival rate and long-term reproductive outputs in these long-lived organisms, thus inducing population decline. However, the demographic consequences of contaminant exposure are largely theoretical because of the dearth of long-term datasets. This study aims to test whether adult survival rate, return to the colony and long-term breeding performance were related to blood mercury (Hg), cadmium (Cd) and persistent organic pollutants (POPs), by using a capture–mark–recapture dataset on the vulnerable wandering albatross Diomedea exulans. We did not find evidence for any effect of contaminants on adult survival probability. However, blood Hg and POPs negatively impacted long-term breeding probability, hatching and fledging probabilities. The proximate mechanisms underlying these deleterious effects are likely multifaceted, through physiological perturbations and interactions with reproductive costs. Using matrix population models, we projected a demographic decline in response to an increase in Hg or POPs concentrations. This decline in population growth rate could be exacerbated by other anthropogenic perturbations, such as climate change, disease and fishery bycatch. This study gives a new dimension to the overall picture of environmental threats to wildlife populations.     

Modelling the effect of fishing on southern Buller's albatross using a 60-year dataset

Abstract

Many albatross populations are declining and a major cause is believed to be incidental mortality from fishing. We investigated the effect of fishing on southern Buller's albatross Thalassarche bulleri bulleri, using a new approach to seabird population modelling that allows estimation of demographic parameters from multiple data types. Three types of data were used: a 60-year set of mark–recapture observations, four censuses of the breeding population, and estimates of fishing effort and bycatch. The fisheries risk to the viability of this population over the last 60 years appears to have been small, since the adult population is estimated to have increased about five-fold over that time. There is some cause for concern in recent changes (population growth has slowed, and perhaps reversed, and adult survival rates are falling). The most common age at first breeding was 12 years, and about 80% of adults breed each year. Annual survival was estimated to be 0.91 for juveniles, and varied between this value and 1 for adults. Though this population is not in immediate danger from fishing, there is a need for continued monitoring to see whether the recent fall in survival rates persists and causes a decline in abundance. Our analysis showed that when, as is common, mark–recapture data do not provide good estimates of all demographic rates, the assessment of seabird population trends can be improved by the use of other types of data, particularly abundance.