Demography and dynamics of three wild horse populations in the Australian Alps

Wild horses (Equus caballus) are a non‐native species occupying over 2800 km² of the nationally significant Australian Alps National Parks. We estimated key demographic parameters (fecundity, adult and juvenile survival and annual finite population growth rate) over 3 years and related these to horse body condition and available food for three populations under natural conditions, and found a trend consistent with food limitation. The populations were independent, with different site characteristics and occupied areas, identified by land managers, as areas of concern about possible conservation impacts. Annual fecundity and juvenile survival varied across sites averaging between 0.21 and 0.31 female young per adult female, and 0.83 and 0.90 per annum, respectively, and annual adult survival was consistent across sites averaging 0.91 per annum. One population was increasing (λ = 1.09 year^?1; 95% CI 1.04–1.14) and two populations were stable (λ ～ 1.0 year^?1). Mean body condition of horses was positively correlated with mean pasture biomass rank. Across the three populations, fecundity, recruitment, body condition and annual finite population growth rate were lowest when mean pasture biomass rank was lowest and conversely highest when pasture rank was highest. We conclude that food limitation appears to be operating across these three sites. We used our results to assess the sensitivity of annual finite rate of increase (λ) to changes in key demographic parameters and found that λ was most sensitive to a change in adult survival, with the second most sensitive parameter being fecundity. Thus, if the aim of management is to reduce the size of the wild horse population then targeting adult survival is most important, followed by fecundity. Finally, we estimated the linear, negative, numerical response for wild horses between annual λ and horses per unit pasture biomass.     

Forest Fragmentation Alters the Population Dynamics of a Late‐successional Tropical Tree

Tropical late‐successional tree species are at high risk of local extinction due to habitat loss and fragmentation. Population‐growth rates in fragmented populations are predicted to decline as a result of reduced fecundity, survival and growth. We examined the demographic effects of habitat fragmentation by comparing the population dynamics of the late‐successional tree Poulsenia armata (Moraceae) in southern Mexico between a continuous forest and several forest fragments using integral projection models (IPMs) during 2010–2012. Forest fragmentation did not lead to differences in population density and even resulted in a higher population‐growth rate (λ) in fragments compared to continuous forests. Habitat fragmentation had drastic effects on the dynamics of P. armata, causing the population structure to shift toward smaller sizes. Fragmented populations experienced a significant decrease in juvenile survival and growth compared to unaltered populations. Adult survival and growth made the greatest relative contributions to λ in both habitat types during 2011–2012. However, the relative importance of juvenile survival and growth to λ was highest in the fragmented forest in 2010–2011. Our Life Table Response Experiment analysis revealed that positive contributions of adult fecundity explained most of the variation of λ between both habitats and annual periods. Finally, P. armata has a relatively slow speed of recovery after disturbances, compromising persistence of fragmented populations. Developing a mechanistic understanding of how forest fragmentation affects plant population dynamics, as done here, will prove essential for the preservation of natural areas.     

The Australian White Ibis (Threskiornis molucca) as a Reservoir of Zoonotic and Livestock Pathogens

Over the last 20 years, Australian white ibis populations (Threskiornis molucca) have expanded into urban areas, leading to increased contact between ibis, domestic animals, and humans. This has led to concern that ibis may transmit pathogens that threaten public health or food production. Here we report results from a study of ibis viral serology and bacterial culture that indicate that ibis are hosts of zoonotic and livestock pathogens such as Salmonella spp., Newcastle disease virus, avian influenza virus, and flaviviruses in Australia. We also performed a behavioral study to measure contact rates among ibis, people, and livestock that determine the potential for disease transmission.     

Life history variation between high and low elevation subspecies of horned larks Eremophila spp.

Environmental variation along elevational gradients can strongly influence life history strategies in vertebrates. We investigated variation in life history patterns between a horned lark subspecies nesting in high elevation alpine habitat Eremophila alpestris articola and a second subspecies in lower elevation grassland and sandy shoreline habitats E. a. strigata. Given the shorter breeding season and colder climate at the northern alpine site we expected E. a. articola to be larger, have lower fecundity and higher apparent survival than E. a. strigata. As predicted, E. a. articola was larger and the trend was toward higher apparent adult survival for E. a. articola than E. a. strigata (0.69 vs 0.51). Contrary to our predictions, however, there was a trend toward higher fecundity for E. a. articola (1.75 female fledglings/female/year vs 0.91). The larger clutch sizes, higher nest survival, and shorter renesting intervals have apparently allowed E. a. articola to compensate for the short breeding season. Estimates of population growth rate (λ) predicted a stable population for E. a. articola (λ=1.00) and a rapidly declining population for E. a. strigata (λ=0.62) this may provide an explanation for the deviations from our expectation of higher reproduction in E. a. strigata. We suggest that anthropogenic influences (e.g. habitat loss and degradation and increased nest predator abundance) to E. a. strigata nesting sites may be responsible for reducing annual fecundity to the point where it is almost half that of E. a. articola. This result suggests that human influenced habitat changes that, in turn, change demographic rates may result in vital rates that do not accurately reflect historically divergent life histories between the two groups. Our results underscore the importance of including estimates of multiple traits in life history studies to provide insight into compensatory interactions among components of demographic rates and to identify recent changes to demographic rates that might result in a mismatch between observed and predicted life history strategies.     

Long-term population dynamics of a managed burrowing owl colony

We analyzed the population dynamics of a burrowing owl (Athene cunicularia) colony at Mineta San Jose International Airport in San Jose, California, USA from 1990–2007. This colony was managed by using artificial burrows to reduce the occurrence of nesting owls along runways and within major airport improvement projects during the study period. We estimated annual reproduction in natural and artificial burrows and age-specific survival rates with mark–recapture techniques, and we estimated the relative contribution of these vital rates to population dynamics using a life table response experiment. The breeding colony showed 2 distinct periods of change: high population growth from 7 nesting pairs in 1991 to 40 pairs in 2002 and population decline to 17 pairs in 2007. Reproduction was highly variable: annual nesting success (pairs that raised ≥1 young) averaged 79% and ranged from 36% to 100%, whereas fecundity averaged 3.36 juveniles/pair and ranged from 1.43 juveniles/pair to 4.54 juveniles/pair. We estimated annual adult survival at 0.710 during the period of colony increase from 1996 to 2001 and 0.465 during decline from 2002 to 2007, but there was no change in annual survival of juveniles between the 2 time periods. Long-term population growth rate (λ) estimated from average vital rates was λ_a = 1.072 with λ_i = 1.288 during colony increase and λ_d = 0.921 (Δλ = 0.368) during decline. A life table response experiment showed that change in adult survival rate during increasing and declining phases explained more than twice the variation in growth rate than other vital rates. Our findings suggest that management and conservation of declining burrowing owl populations should address factors that influence adult survival. © 2011 The Wildlife Society.     

Comparison of asymptotic population growth rates with heterogeneous variances

This report explores how the heterogeneity of variances affects randomization tests used to evaluate differences in the asymptotic population growth rate, λ. The probability of Type I error was calculated in four scenarios for populations with identical λ but different variance of λ: (1) Populations have different projection matrices: the same λ may be obtained from different sets of vital rates, which gives room for different variances of λ. (2) Populations have identical projection matrices but reproductive schemes differ and fecundity in one of the populations has a larger associated variance. The two other scenarios evaluate a sampling artifact as responsible for heterogeneity of variances. The same population is sampled twice, (3) with the same sampling design, or (4) with different sampling effort for different stages. Randomization tests were done with increasing differences in sample size between the two populations. This implies additional differences in the variance of λ. The probability of Type I error keeps at the nominal significance level (α = .05) in Scenario 3 and with identical sample sizes in the others. Tests were too liberal, or conservative, under a combination of variance heterogeneity and different sample sizes. Increased differences in sample size exacerbated the difference between observed Type I error and the nominal significance level. Type I error increases or decreases depending on which population has a larger sample size, the population with the smallest or the largest variance. However, by their own, sample size is not responsible for changes in Type I errors.     

A review of vital rates and cause‐specific mortality of elk Cervus elaphus populations in eastern North America

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Variation in vital-rate sensitivity between populations of Texas horned lizards

Demographic studies of imperiled populations can aid managers in planning conservation actions. However, applicability of findings for a single population across a species’ range is sometimes questionable. We conducted long-term studies (8 and 9 years, respectively) of 2 populations of the lizard Phrynosoma cornutum separated by 1000 km within the historical distribution of the species. The sites were a 15-ha urban wildlife reserve on Tinker Air Force Base (TAFB) in central Oklahoma and a 6000-ha wildland site in southern Texas, the Chaparral Wildlife Management Area (CWMA). We predicted a trade-off between the effect of adult survival and fecundity on population growth rate (λ), leading to population-specific contributions of individual vital rates to λ and individualized strategies for conservation and management of this taxon. The CWMA population had lower adult survival and higher fecundity than TAFB. As predicted, there was a trade-off in the effects of adult survival and fecundity on λ between the two sites; fecundity affected λ more at CWMA than at TAFB. However, adult survival had the smallest effect on λ in both populations. We found that recruitment in P. cornutum most affected λ at both sites, with hatchling survival having the strongest influence on λ. Management strategies focusing on hatchling survival would strongly benefit both populations. As a consequence, within the constraint of the need to more accurately estimate hatchling survival, managers across the range of species such as P. cornutum could adopt similar management priorities with respect to stage classes, despite intra-population differences in population vital rates.     

Linking vital rates to invasiveness of a perennial herb

Invaders generally show better individual performance than non-invaders and, therefore, vital rates (survival, growth, fecundity) could potentially be used to predict species invasiveness outside their native range. Comparative studies have usually correlated vital rates with the invasiveness status of species, while few studies have investigated them in relation to population growth rate. Here, I examined the influence of five vital rates (plant establishment, survival, growth, flowering probability, seed production) and their variability (across geographic regions, habitat types, population sizes and population densities) on population growth rate (λ) using data from 37 populations of an invasive, iteroparous herb (Lupinus polyphyllus) in a part of its invaded range in Finland. Variation in vital rates was often related to habitat type and population density. The performance of the populations varied from declining to rapidly increasing independently of habitat type, population size or population density, but differed between regions. The population growth rate increased linearly with plant establishment, and with the survival and growth of vegetative individuals, while the survival of flowering individuals and annual seed production were not related to λ. The vital rates responsible for rapid population growth varied among populations. These findings highlight the importance of both regional and local conditions to plant population dynamics, demonstrating that individual vital rates do not necessarily correlate with λ. Therefore, to understand the role of individual vital rates in a species ability to invade, it is necessary to quantify their effect on population growth rate.     

Direct and indirect effects of an insect outbreak increase the reproductive output for an avian insectivore and nest‐cavity excavator,the red‐breasted nuthatch Sitta canadensis

Community‐wide food pulses may ameliorate food constraints but may also result in increased competition for other resources and predation rates. In cavity‐nesting vertebrate communities, where the availability of tree cavities can limit reproduction and the reuse of cavities can increase nest predation by squirrels, excavators may maximize their fecundity by creating new cavities in competitor‐ and predator‐rich habitats that undergo food pulses. The reproductive cost associated with excavation (i.e. increased energy allocation early in the breeding season that often delays laying and thereby reduces clutch size), may be reduced if food pulses allow for a longer breeding season and larger clutches. A large‐scale mountain pine beetle Dendroctonus ponderosae outbreak that occurred during our long‐term study (1995–2009) provided a natural food supplementation experiment across 27 sites in British Columbia, Canada. We examined the effects of a reduction in food constraints accompanied with increases in excavation rates, conspecific density and nest predation risk on the fecundity of a facultative excavator, the red‐breasted nuthatch Sitta canadensis. We found a total of 420 nests in tree cavities. Nuthatch clutch sizes ranged from two to nine eggs, and broods from one to nine fledglings per nest. Later clutches were larger at sites and in years with high beetle abundance (mean clutch size of six eggs did not decline later in the season), second broods were produced in outbreak years (usually only one nesting attempt/normal year), and the number of fledglings per successful nest increased with increasing beetle abundance and nuthatch densities, but declined with increased squirrel densities. Since fecundity did not differ between new and reused cavities, the costs and benefits of excavation versus cavity reuse may be neutralized for nuthatches during strong resource pulses. Overall, the beetle outbreak reduced food constraints for nuthatches and provided alternate food for nest predators, allowing increased annual fecundity.     

Optimal predator management for mountain sheep conservation depends on the strength of mesopredator release

Large predators often suppress ungulate population growth, but they may also suppress the abundance of smaller predators that prey on neonatal ungulates. Antagonistic interactions among predators may therefore need to be integrated into predator–prey models to effectively manage ungulate–predator systems. We present a modeling framework that examines the net impact of interacting predators on the population growth rate of shared prey, using interactions among wolves Canis lupus, coyotes Canis latrans and Dall sheep Ovis dalli dalli as a case study. Wolf control is currently employed on approximately 16 million ha in Alaska to increase the abundance of ungulates for human harvest. We hypothesized that the positive effects of wolf control on Dall sheep population growth could be counteracted by increased levels of predation by coyotes. Coyotes and Dall sheep adult females (ewes) and lambs were radiocollared in the Alaska Range from 1999–2005 to estimate fecundity, age‐specific survival rates, and causes of mortality in an area without wolf control. We used stage‐structured population models to simulate the net effect of wolf control on Dall sheep population growth (λ). Our models accounted for stage‐specific predation rates by wolves and coyotes, compensatory mortality, and the potential release of coyote populations due to wolf control. Wolves were the main predators of ewes, coyotes were the main predators of lambs, and wolves were the main source of mortality for coyotes. Population models predicted that wolf control could increase sheep λ by 4% per year in the absence of mesopredator release. However, if wolf control released coyote populations, our models predicted that sheep λ could decrease by up to 3% per year. These results highlight the importance of integrating antagonistic interactions among predators into predator–prey models, because the net effect of predator management on shared prey can depend critically on the strength of mesopredator release.     

Demographic response of plant populations to habitat fragmentation and temporal environmental variability

Many plant species currently exist in fragmented populations of different sizes, while they also experience unpredictable climatic fluctuation over time. However, we still understand little about how plant demography responds to such spatial and temporal environmental variability. We studied population dynamics of an understory perennial herb Trillium camschatcense in the Tokachi plain of Hokkaido, Japan, where a significant effect of forest fragmentation on seedling recruitment was previously reported. Four populations across a range of fragment sizes were studied for 6 years, and the data were analyzed using matrix population models. Per capita fecundity (the number of recruits per plant) varied greatly among populations, but the variation in population growth rates (λ) was mainly driven by the variation in stasis and growth rates, suggesting that the general trend of reduced fecundity in fragmented populations may not be readily translated into subsequent dynamics. Temporal variation in λ among years was more than 2 times larger than spatial variation among populations, and this result was likely attributable to the contrasting response of correlation structures among demographic rates. The among-population variation in λ was dampened by negative covariation between matrix elements possibly due to density-dependent regulation as well as an inherent constraint that some elements are not independent, whereas positive covariation between matrix elements resulted in large temporal variation in λ. Our results show that population dynamics responded differently to habitat fragmentation and temporal variability of the environment, emphasizing the need to discriminate these spatial and temporal variations in demographic models. Although no populations were projected to be declining in stochastic simulations, correlation between current habitat size and plant density implies historical λ is positively related to habitat size.     

Inbreeding levels and prey abundance interact to determine fecundity in natural populations of two species of wolf spider

Long-term effective population size is expected, and has been shown, to correlate positively with various measures of population fitness. Here we examine the interacting effects of population size (as a surrogate for genetic factors) and prey consumption rates (as a surrogate for environmental quality) on fecundity in two sympatric species of wolf spider, Rabidosa punctulata and Rabidosa rabida. Population size was estimated in each of seven genetically isolated populations in each of 3 years using mark-recapture methods. Fecundity was estimated as the mean number of live offspring produced by ∼15 females sampled from each population of each species each year for 3 years. Prey consumption rates were estimated by sampling ∼300 spiders per population per year and assaying the proportion of spiders with prey. Larger populations have higher fecundity and more genetic diversity than smaller populations. Variation among populations in fecundity for a given year could be attributed most strongly to differences in population size, with variation in prey consumption rates and the interaction between population size and prey consumption playing smaller but still important roles. During the most stressful environmental conditions, the smallest populations of both species experienced disproportionately low-fecundity rates, more than doubling the estimated number of lethal equivalents during those years. The evidence presented in this paper for inbreeding-environment interactions at the population level and further evidence for a log-linear relationship between population size and fitness have important implications for conservation.     

Integrated population model reveals that kestrels breeding in nest boxes operate as a source population

The identification of the source–sink status of a population is critical for the establishment of conservation plans and enacting smart management decisions. We developed an integrated population model to formally assess the source status of a kestrel Falco tinnunculus population breeding in nest boxes in Switzerland. We estimated juvenile and adult survival, reproduction and net dispersal (emigration/immigration) by jointly analyzing capture–recapture, dead recovery, breeding monitoring and population survey data. We also investigated the role of nest boxes on kestrel demography and assessed the contributions of vital rates to realized population growth rates. The results indicate that the kestrel population breeding in nest boxes has acted as a source over the 15 years of the study duration. A quantitative approach suggests that a substantial number of individuals have emigrated annually from this population likely affecting the population dynamics outside the management area. Variation in fecundity explained 34% of the temporal variability of the population growth rate. Moreover, a literature review suggests that kestrel pairs produce on average 1.4 chicks more per breeding attempt in nest boxes compared to natural open nests. Together, these findings suggest that fecundity was an important driver for the dynamics of this population and that nest boxes have contributed to its raise. Nest boxes are regularly used as an efficient tool for conservation management. We suggest that such a conservation action can result in the establishment of a source population being beneficial for populations both inside and outside the managed area.     

Rain,prey and predators: climatically driven shifts in frog abundance modify reproductive allometry in a tropical snake

To predict the impacts of climate change on animal populations, we need long-term data sets on the effects of annual climatic variation on the demographic traits (growth, survival, reproductive output) that determine population viability. One frequent complication is that fecundity also depends upon maternal body size, a trait that often spans a wide range within a single population. During an eight-year field study, we measured annual variation in weather conditions, frog abundance and snake reproduction on a floodplain in the Australian wet-dry tropics. Frog numbers varied considerably from year to year, and were highest in years with hotter wetter conditions during the monsoonal season (“wet season”). Mean maternal body sizes, egg sizes and post-partum maternal body conditions of frog-eating snakes (keelback, Tropidonophis mairii, Colubridae) showed no significant annual variation over this period, but mean clutch sizes were higher in years with higher prey abundance. Larger females were more sensitive to frog abundance in this respect than were smaller conspecifics, so that the rate at which fecundity increased with body size varied among years, and was highest when prey availability was greatest. Thus, the link between female body size and reproductive output varied among years, with climatic factors modifying the relative reproductive rates of larger (older) versus smaller (younger) animals within the keelback population.     

Behavioural Adaptation of a Bird from Transient Wetland Specialist to an Urban Resident

Dramatic population increases of the native white ibis in urban areas have resulted in their classification as a nuisance species. In response to community and industry complaints, land managers have attempted to deter the growing population by destroying ibis nests and eggs over the last twenty years. However, our understanding of ibis ecology is poor and a question of particular importance for management is whether ibis show sufficient site fidelity to justify site-level management of nuisance populations. Ibis in non-urban areas have been observed to be highly transient and capable of moving hundreds of kilometres. In urban areas the population has been observed to vary seasonally, but at some sites ibis are always observed and are thought to be behaving as residents. To measure the level of site fidelity, we colour banded 93 adult ibis at an urban park and conducted 3-day surveys each fortnight over one year, then each quarter over four years. From the quarterly data, the first year resighting rate was 89% for females (n = 59) and 76% for males (n = 34); this decreased to 41% of females and 21% of males in the fourth year. Ibis are known to be highly mobile, and 70% of females and 77% of males were observed at additional sites within the surrounding region (up to 50 km distant). Our results indicate that a large proportion of ibis have chosen residency over transience both within the study site and across the broader urban region. Consequently the establishment of refuge breeding habitat should be a priority localised management may be effective at particular sites, but it is likely to have an impact across the broader population.     

Effects of black bear relocation on elk calf recruitment at Great Smoky Mountains National Park

Previous research from 2001 to 2006 on an experimentally released elk (Cervus elaphus) population at Great Smoky Mountains National Park (GSMNP or Park) indicated that calf recruitment (i.e., calves reaching 1 yr of age per adult female elk) was low (0.306, total SE = 0.090) resulting in low or negative population growth (λ = 0.996, 95% CI = 0.945–1.047). Black bear (Ursus americanus) predation was the primary calf mortality factor. From 2006 to 2008, we trapped and relocated 49 bears (30 of which were radiocollared) from the primary calving areas in the Park and radiomonitored 67 (28 M:39 F) adult elk and 42 calves to compare vital rates and population growth with the earlier study. A model with annual calf recruitment rate correlating with the number of bears relocated each year was supported (ΔAIC_c = 0.000; β = 0.070, 95% CI = 0.028–0.112) and a model with annual calf recruitment differing from before to during bear relocation revealed an increase to 0.544 (total SE = 0.098; β = −1.092, 95% CI = −1.180 to −0.375). Using vital rates and estimates of process standard errors observed during our study, 25-yr simulations maintained a mean positive growth rate in 100% of the stochastic trials with λ averaging 1.118 (95% CI = 1.096–1.140), an increase compared with rates before bear relocation. A life table response experiment revealed that increases in population growth were mostly (67.1%) due to changes in calf recruitment. We speculate that behavioral adaptation of the elk since release also contributed to the observed increases in recruitment and population growth. Our results suggest that managers interested in elk reintroduction within bear range should consider bear relocation as a temporary means of increasing calf recruitment. © 2011 The Wildlife Society.     

Population demography of frillneck lizards (Chlamydosaurus kingii,Gray 1825) in the wet‐dry tropics of Australia

We explore the effects of biotic and abiotic factors on the population demography of frillneck lizards (Chlamydosaurus kingii) in the Australian wet‐dry tropics. Annual growth rates of males were significantly higher across all body sizes compared to females, resulting in a significant larger maximum body size in males. Both male and female lizards were highly philopatric and 81% of the among‐year recapture distances were less than 200 m. Juvenile and adult frillnecks were subjected to low but highly variable annual survival rates. Both proportion of juveniles and relative proportion of reproductive females showed extensive among‐year variations. No relationship was, however, observed between proportion of gravid females and that of juveniles captured during the subsequent year. High rainfall in January was negatively correlated with recruitment most likely caused by increased egg/embryo mortality due to flooding of nest sites. We therefore suggest that the lack of association between female reproduction and juvenile recruitment was due to the effects of stochastic variation in January rainfall. Lizard numbers increased during the first five years of the study followed by a decline during the subsequent four years. Our analyses show that annual variation in survival constituted the main determinant in driving the annual change in frillneck numbers. Surprisingly, no relationship was observed between fillneck population dynamics and annual variation in juvenile recruitment. We suggest that the 7‐years over which these analyses were conducted were insufficient to detect any significant effects of recruitment on lizard numbers, demonstrating the need for long‐term studies to accurately document vertebrate population demographic processes in areas experiencing stochastic variations rainfall such as the Australian wet‐dry tropics.     

Population viability analysis of Lower Missouri River shovelnose sturgeon with initial application to the pallid sturgeon

Demographic models for the shovelnose (Scaphirhynchus platorynchus) and pallid (S. albus) sturgeons in the Lower Missouri River were developed to conduct sensitivity analyses for both populations. Potential effects of increased fishing mortality on the shovelnose sturgeon were also evaluated. Populations of shovelnose and pallid sturgeon were most sensitive to age‐0 mortality rates as well as mortality rates of juveniles and young adults. Overall, fecundity was a less sensitive parameter. However, increased fecundity effectively balanced higher mortality among sensitive age classes in both populations. Management that increases population‐level fecundity and improves survival of age‐0, juveniles, and young adults should most effectively benefit both populations. Evaluation of reproductive values indicated that populations of pallid sturgeon dominated by ages ≥35 could rapidly lose their potential for growth, particularly if recruitment remains low. Under the initial parameter values portraying current conditions the population of shovelnose sturgeon was predicted to decline by 1.65% annually, causing the commercial yield to also decline. Modeling indicated that the commercial yield could increase substantially if exploitation of females in ages ≤12 was highly restricted.