Biases in the estimation of the demographic parameters of a snowshoe hare population

1. Survival rates and natalities for a population of snowshoe hares in the Yukon were estimated independently of and simultaneously with estimates of population change during the increase phase of a hare cycle.
2. Simple demographic models are used to show that even though the estimated survival rates and natalities were high relative to previously published estimates, the observed demographic parameters are unable to explain the extent of population increase, and we conclude that some of these parameters must be underestimates.
3. A sensitivity analysis is used to examine the potential influence of changes in these demographic parameters on the population growth rate. During most years of the hare cycle the population growth rate is potentially most sensitive to changes in juvenile postweaning survival. Only during crash years is adult survivorship likely to be a more important determinant of the rate of population change.
4. Examination of previously published data sets on two full population cycles suggests that while survival rates are positively correlated with population growth rates, their incorporation into demographic models results in frequent underestimation of the rate of population increase.     

Sea buckthorn berries Hippophae rhamnoides L. predict size and composition of a great tit population Parus major L

In seasonal environments variation in food abundance in the non‐breeding season is thought to affect songbird population dynamics. In a unique tit‐sea buckthorn berry system we can estimate the berry abundance and both the tit consumption and population dynamics. Six hundred nest boxes were available to great and blue tits Cyanistes caeruleus for breeding in spring and roosting in winter. We followed the dynamics including the recapture histories of individually marked great tits from 2008 to 2014. In each year we estimated 1) the winter sea buckthorn berry availability, 2) an index of berry consumption in December based on the colour of the faeces of roosting birds, 3) the number of breeding great and blue tits, 4) both recapture probability and the return rate of the great tits and 5) immigration rates. December berry abundance positively predicted the number of breeding pairs of both species in the subsequent season and great tit return rates in the second half of the winter. There was support for a sex specific berry effect on the adult return rate in the great tit: female return rate was associated less strongly to berry abundance than male return rate. This skewed the sex ratio of the local breeders in the following breeding season. Intriguingly, annual berry consumption in December was not related to berry abundance, and individuals consuming more berries tended to have slightly lower return rates. Reproductive rate was not related to berry abundance. There was hardly support for a relation between immigration rates of first year breeders and berry abundance. Taken together these results imply that berry stock not only affected population size but also the population composition through sex specific exchange with the surroundings. Since population density covaried with berry abundance, density dependent effects provide an alternative explanation for the patterns observed.     

An assessment of overwinter food limitation in a snowshoe hare population at a cyclic low

Snowshoe hares (Lepus americanus) undergo 8- to 11-year population cycles caused by direct and/or interactive effects of overwinter food shortage and predation. However, the demographic significance of food shortage during cyclic population lows remains unclear. I evaluated the importance of overwinter food limitation to the demography (numbers, age and sex ratios) of low-density hare populations during two winters in Manitoba. Also, I examined whether the hypothesized differences in demography of fed and unfed hare populations could be explained by altered movement patterns or social dynamics. Bimonthly live-trapping revealed that food failed to have a direct long-term effect on the number, or change in number, of hares estimated to be on the three supplemented areas, relative to three control areas. Modest numerical responses to supplementation tended to be short-term (i.e., restricted to winter) and related to pre-supplementation densities, with the study area characterized by the highest hare density displaying the strongest and most consistent response to added food. During winter the percentage of females was remarkably variable among study areas and time periods, but added food may have augmented slightly the proportion of females captured in traps. There tended to be slightly more juveniles on supplemented areas during winter periods, and this effect was strongest during the first winter (1991–1992). I found that immigration rates and percentage of hares that were considered to be transient animals were similar on supplemented and control areas, and that spatial distribution of radio-collared animals on versus off of study areas also was similar. Because the overall effect of food on hare populations was small and short-lived, and could be explained largely by small increases in survival and reproduction, I conclude that the study population was not subject to overwinter food limitation. Received: 22 February 1998 / Accepted: 12 February 1999     

Density‐dependent demography and movements in a cyclic brown lemming population

Theoretical modeling predicts that both direct and delayed density‐dependence are key factors to generate population cycles. Deciphering density‐dependent processes that lead to variable population growth characterizing different phases of the cycles remains challenging. This is particularly the case for the period of prolonged low densities, which is inherently data deficient. However, demographic analyses based on long‐term capture–mark–recapture datasets can help resolve this question. We relied on a 16‐year (2004–2019) live‐trapping program to analyze the summer demography and movements of a cyclic brown lemming population in the Canadian Arctic. More specifically, we examined if inversely density‐dependent processes could explain why population growth can remain low during the prolonged low phase. We found that the proportion of females in the population was inversely density‐dependent with a strong male‐biased sex ratio at low densities but not at high densities. However, survival of adult females was higher than adult males, but both had lower survival at low densities than at high ones. Distances moved by both adult males and females were density‐dependent, and proportion of females in reproductive condition was weakly density‐dependent as it tended to increase at low density. Individual body condition, measured as monthly change in body mass, was not density‐dependent. Overall, the strong male‐biased sex ratio at very low densities suggests a loss of reproductive potential due to the rarity of females and appears to be the most susceptible demographic factor that could contribute to the prolonged low phase in cyclic brown lemmings. What leads to this sex‐bias in the first place is still unclear, potentially owing to our trapping period limited to the summer, but we suggest that it could be due to high predation rate on breeding females in winter.     

Source populations in carnivore management: cougar demography and emigration in a lightly hunted population

Wildlife agencies typically attempt to manage carnivore numbers in localized game management units through hunting, and do not always consider the potential influences of immigration and emigration on the outcome of those hunting practices. However, such a closed population structure may not be an appropriate model for management of carnivore populations where immigration and emigration are important population parameters. The closed population hypothesis predicts that high hunting mortality will reduce numbers and densities of carnivores and that low hunting mortality will increase numbers and densities. By contrast, the open population hypothesis predicts that high hunting mortality may not reduce carnivore densities because of compensatory immigration, and low hunting mortality may not result in more carnivores because of compensatory emigration. Previous research supported the open population hypothesis with high immigration rates in a heavily hunted (hunting mortality rate=0.24) cougar population in northern Washington. We test the open population hypothesis and high emigration rates in a lightly hunted (hunting mortality rate=0.11) cougar population in central Washington by monitoring demography from 2002 to 2007. We used a dual sex survival/fecundity Leslie matrix to estimate closed population growth and annual census counts to estimate open population growth. The observed open population growth rate of 0.98 was lower than the closed survival/fecundity growth rates of 1.13 (deterministic) and 1.10 (stochastic), and suggests a 12–15% annual emigration rate. Our data support the open population hypothesis for lightly hunted populations of carnivores. Low hunting mortality did not result in increased numbers and densities of cougars, as commonly believed because of compensatory emigration.     

The demography of a newly established Osprey Pandion haliaetus population in France

Using long‐term mark–resighting data acquired over 27 years in continental France, we estimated demographic parameters and modelled the dynamics of a newly established population of Ospreys Pandion haliaetus using a life‐history model. We then performed prospective and retrospective analyses to estimate the sensitivity of the population growth rate to demographic parameters, and to quantify their contribution to the observed variation in abundance. The observed population growth rate was estimated at 1.150 (from one to 38 pairs in the period 1985–2011), and the stochastic population growth rate was estimated at 1.156. The number of fledglings per nest made the largest contribution to the variance of the observed population growth rate. Breeding productivity was stable across years. In contrast, the prospective analysis indicated that the sensitivity of the population growth rate was greatest for immigration and adult survival. Our results suggest that the increase of a new and recently established breeding population of Ospreys was mainly driven by local dynamics (high productivity and high proportion of breeding individuals), with no sign of density‐dependence except for juvenile survival. This probably reflects highly favourable conditions for breeding. Our results show that productivity can be a major driver in recovering raptor populations, and conservation work should aim to protect occupied nest‐sites and their surrounding habitat and to maintain highly favourable foraging areas in the vicinity of breeding sites.     

Growth and demography of a re‐introduced population of White‐tailed Eagles Haliaeetus albicilla

White‐tailed Eagles Haliaeetus albicilla became extinct in Britain in 1918 following prolonged persecution. Intensive conservation efforts since the 1970s have included the re‐introduction of the species to Britain through two phases of release of Norwegian fledglings in western Scotland in 1975–85 and 1993–98. Population growth and breeding success have been monitored closely to the present day, aided by the use of patagial tags to individually mark most released birds as well as a high proportion of wild‐bred nestlings. This study reviews the growth and demography of this re‐introduced population, and makes comparisons with other European populations. For the first time, we compare the demographic rates of released and wild‐bred birds in the Scottish population. Breeding success in the Scottish population has increased over time as the average age and experience of individuals in the population have increased, and success tends to be higher where one or both adults are wild‐bred. Current levels of breeding success remain low compared with some other populations in Europe, but similar to those in Norway where weather conditions and food availability are likely to be most similar. Survival rates in Scotland are similar to those recorded elsewhere, but survival rates of released birds are lower than those of wild‐bred birds, especially during the first 3 years of life. Despite the effect of lower survival rates of released birds in limiting overall population growth rate, the recent rate of growth of the Scottish population remains high relative to other recovering populations across Europe. Differences in demographic rates of wild‐bred and released birds suggest that in future re‐introduction programmes, steps to maximize the success and output of the earliest breeding attempts would help ensure the most rapid shift to a population composed largely of wild‐bred birds, which should then have a higher rate of increase.     

不同藻密度下布洛芬浓度对萼花臂尾轮虫生命表统计学参数的影响

在不同斜生栅藻(Scenedesmus obliquus)密度(1.0×10~6、2.0×10~6个细胞/m L和4.0×10~6个细胞/m L)下,研究了不同浓度的(0、1、10、100、1000μg/L和5000μg/L)布洛芬对萼花臂尾轮虫(Brachionus calyciflorus)生命表统计学参数的影响。结果表明,与各藻密度下的对照组相比,当藻密度为1.0×106个细胞/m L时,100—5000μg/L布洛芬处理组中轮虫的生命期望和平均寿命显著缩短,100μg/L布洛芬处理组中轮虫的世代时间显著缩短,1.0μg/L布洛芬处理组中轮虫的净生殖率显著提高,10—5000μg/L布洛芬处理组中轮虫的后代混交率显著提高。当藻密度为2.0×10~6个细胞/mL时,10—5000μg/L布洛芬处理组中轮虫的生命期望和平均寿命显著缩短,1000μg/L和5000μg/L布洛芬处理组中轮虫的世代时间显著缩短,1、10、1000μg/L和5000μg/L布洛芬处理组中轮虫的种群内禀增长率显著提高,1μg/L和1000μg/L布洛芬处理组中轮虫的后代混交率显著提高。当藻密度为4.0×10~6个细胞/mL时,10—5000μg/L布洛芬处理组中轮虫的生命期望、平均寿命和世代时间显著缩短。藻密度对轮虫的世代时间、净生殖率和种群内禀增长率有显著性影响(P0.05),布洛芬浓度对轮虫的生命期望、平均寿命、世代时间、净生殖率和种群内禀增长率有显著性影响(P0.05),藻密度和布洛芬浓度的交互作用对轮虫的生命期望、平均寿命和后代混交率有显著性影响(P0.05)。在实验设置的布洛芬浓度范围内,2.0×10~6个细胞/m L藻密度下,轮虫的生命期望、平均寿命和世代时间与布洛芬浓度之间均具有显著的剂量—效应关系(P0.05); 4.0×10~6个细胞/m L藻密度下,轮虫的生命期望、平均寿命和净生殖率与布洛芬浓度之间均具有显著的剂量—效应关系(P0.05)。     

Camera-trapping density estimates suggest critically low population sizes for the Wet Tropics subspecies of the spotted-tailed quoll (Dasyurus maculatus gracilis)

Accurate estimates of distribution and population density are critical for the management of threatened species. This is particularly pertinent for mammalian predators, whose generally low population density, elusive nature, and large home range requirements make it difficult to detect declines. We aimed to refine population estimates of the northern spotted-tailed quoll (Dasyurus maculatus gracilis) in the Wet Tropics bioregion, to estimate the total number of adults, the likely size of subpopulations across the known distribution of the subspecies, and its associated conservation status. We performed targeted upland camera-trapping surveys from June 2017 to May 2019. To calculate population densities, we used a combination of the number of individuals identified from each survey and the mean maximum distance moved from three life history stages. We then extrapolated these estimates to modelled suitable habitat areas, refined by the camera-trapping surveys. Population sizes for the six defined subpopulations were estimated, and ranged from approximately 5 to 105 individuals. The total population was estimated to be 221 individuals. This total population estimate, and the estimates for each of the subpopulations, are lower than previous published estimates and are cause for concern. Given the low population estimates presented here and unresolved threats driving declines in some subpopulations, we suggest elevation of this subspecies to Critically Endangered under the EPBC Act.     

Long-term demography of a zebra mussel (Dreissena polymorpha) population

1. We used long‐term data and a simulation model to investigate temporal fluctuations in zebra mussel populations, which govern the ecological and economic impacts of this pest species. 2. The size of the zebra mussel (Dreissena polymorpha) population in the Hudson River estuary fluctuated approximately 11‐fold across a 13‐year period, following a cycle with a 2–4 year period. 3. This cycling was caused by low recruitment during years of high adult population size, rapid somatic growth of settled animals, and adult survivorship of 50% per year. 4. Adult growth and body condition were weakly correlated with phytoplankton biomass. 5. The habitat distribution of the Hudson's population changed over the 13‐year period, with an increasing proportion of the population spreading onto soft sediments over time. The character of soft‐sediment habitats in the Hudson changed because of large amounts (mean = 34 g DM m^?2) of empty zebra mussel shells now in the sediments. 6. Simulation models show that zebra mussel populations can show a range of long‐term trajectories, depending on the balance between adult space limitation, larval food limitation, and disturbance. 7. Effective understanding and management of the effects of zebra mussels and other alien species depend on understanding of their long‐term demography, which may vary across ecosystems.     

不同斜生栅藻密度下布洛芬浓度对多刺裸腹溞生命表统计学参数的影响

以多刺裸腹溞为受试生物,在较低(0.5×10⁶ cells·mL^-1)、中等(1.0×10⁶ cells·mL^-1)和较高(2.0×10⁶ cells·mL^-1)的斜生栅藻密度下研究了不同浓度(0、0.45、0.9、1.8、3.6、7.2 和14.4 mg·L^-1)的布洛芬对多刺裸腹溞生命表统计学参数的影响.结果表明: 在较低和较高的食物密度下,布洛芬浓度分别对多刺裸腹溞的种群内禀增长率和生命期望具有显著性影响;但在3个食物密度下,布洛芬浓度对多刺裸腹溞的世代时间、净生殖率和总生殖率均无显著性影响.与对照组相比,在较低的食物密度下,暴露于0.45、0.9、3.6和14.4 mg·L^-1布洛芬溶液中的多刺裸腹溞的种群内禀增长率显著降低;在较高的食物密度下,暴露于3.6 和14.4 mg·L^-1布洛芬溶液中的多刺裸腹溞的生命期望显著延长.表明在较低的食物密度下,布洛芬对多刺裸腹溞的种群增长具有不利的影响;但在中等和较高的食物密度下其不利影响消失.在较高的食物密度下,布洛芬对多刺裸腹溞的存活具有促进作用;但在较低和中等食物密度下其促进作用并未出现.布洛芬对多刺裸腹溞部分生命表统计学参数的影响因食物密度的不同而异.     

Sex-specific population dynamics and demography of capercaillie (Tetrao urogallus L.) in a patchy environment

Modeling the population dynamics of patchily distributed species is a challenge, particularly when inference must be based on incomplete and small data sets such as those from most species of conservation concern. Here, we develop an open population spatial capture–recapture (SCR) model with sex-specific detection and population dynamics parameters to investigate population trend and sex-specific population dynamics of a capercaillie (Tetrao urogallus) population in Switzerland living in eight distinct forest patches totaling 22 km² within a region of 908 km² and sampled via scat collection. Our model accounts for the patchy distribution of habitat and the uncertainty introduced by collecting data only every third year, while producing sex by patch population trajectories. The estimated population trajectory was a decline of 2% per year; however, the sex specificity of the model revealed a decline in the male population only, with no evidence of decline in the female population. The decline observed in males was explained by the demography of just two of the eight patches. Our study highlights the flexibility of open population SCR models for assessing population trajectories through time and across space and emphasizes the desirability of estimating sex-stratified population trends especially in species of conservation concern.     

Effects of food supplementation on home-range size, reproductive success, productivity and recruitment in a small population of Iberian lynx

In a conservation context, food supplementation is a management tool used to reverse the decline of food-limited populations by means of positive changes in behaviour and fitness that may be reflected in population parameters. The critically endangered Iberian lynx Lynx pardinus has suffered a dramatic decline primarily because of the severe drop of its main prey, the European wild rabbit Oryctolagus cuniculus . To reverse this situation, a food supplementation programme has been implemented in Doñana, south-west Spain, since 2002. In this study, we assess the utility of providing artificial food to reduce home-range (HR) size, and to increase productivity, survival and recruitment in a scenario of low lynx density, as compared with reference data from the same population in the absence of extra food. Food supplementation produced a significant contraction of core areas, but not of complete lynx HRs. We did not detect any significant change in productivity or dispersal rates, but supplementation could have helped transient adult lynx to settle down. The positive effects of food supplementation may have been partly countered by factors such as inbreeding, Allee effects and disease outbreaks, whose effects may have been exacerbated in this small lynx population. Food supplementation, however, proved useful to retain individuals, to keep range sizes within their normal range of values, thus maintaining spatial organization, and to allow lynx reproduction and kitten survival in areas with very low prey density. Therefore, we recommend keeping an extensive and intensive supplementary feeding programme until the density of wild rabbits will enable the viability of this endangered lynx population.