Population ecology of the velvet gecko,Oedura lesueurii in south eastern Australia: Implications for the persistence of an endangered snake

Abstract Ecological specialization, such as major dependence upon a single‐prey species, can render a predator taxon vulnerable to extinction. In such cases, understanding the population dynamics of that prey type is important for conserving the predator that relies upon it. In eastern Australia, the endangered broad‐headed snake Hoplocephalus bungaroides feeds largely on velvet geckos (Oedura lesueurii). We studied growth, longevity and reproduction in a population of velvet geckos in Morton National Park in south‐eastern Australia. We marked 458 individual geckos over a 3‐year period (1992–1995) and made yearly visits to field sites from 1995–2006 to recapture marked individuals. Female geckos grew larger than males, and produced their first clutch at age 4 years. Males can mature at 2 years, but male–male combat for females probably forces males to delay reproduction until age 3 years. Females lay a single clutch of two eggs in communal nests in November, and up to 22 females deposited eggs in a single nest. Egg hatching success was high (100%), and juveniles had high survival (76%) during their first 6 months of life. Velvet geckos are long‐lived, and the mean age of marked animals recaptured after 1995 was 6.1 years (males) and 8.4 years (females). Older females (7.5–9.5 years) were all gravid when last recaptured. Like other temperate‐climate gekkonids, O. lesueurii has a ‘slow’ life history, and population viability could be threatened by any factors that increase egg or adult mortality. Two such factors – the removal of ‘bush rocks’ for urban gardens, and the overgrowth of rock outcrops by vegetation – could render small gecko populations vulnerable to extinction. In turn, the reliance of predatory broad‐headed snakes on this slow‐growing lizard species may increase its vulnerability to extinction.     

Differential adult survival at close seabird colonies: The importance of spatial foraging segregation and bycatch risk during the breeding season

Marine megafauna, including seabirds, are critically affected by fisheries bycatch. However, bycatch risk may differ on temporal and spatial scales due to the uneven distribution and effort of fleets operating different fishing gear, and to focal species distribution and foraging behavior. Scopoli's shearwater Calonectris diomedea is a long‐lived seabird that experiences high bycatch rates in longline fisheries and strong population‐level impacts due to this type of anthropogenic mortality. Analyzing a long‐term dataset on individual monitoring, we compared adult survival (by means of multi‐event capture–recapture models) among three close predator‐free Mediterranean colonies of the species. Unexpectedly for a long‐lived organism, adult survival varied among colonies. We explored potential causes of this differential survival by (1) measuring egg volume as a proxy of food availability and parental condition; (2) building a specific longline bycatch risk map for the species; and (3) assessing the distribution patterns of breeding birds from the three study colonies via GPS tracking. Egg volume was very similar between colonies over time, suggesting that environmental variability related to habitat foraging suitability was not the main cause of differential survival. On the other hand, differences in foraging movements among individuals from the three colonies expose them to differential mortality risk, which likely influenced the observed differences in adult survival. The overlap of information obtained by the generation of specific bycatch risk maps, the quantification of population demographic parameters, and the foraging spatial analysis should inform managers about differential sensitivity to the anthropogenic impact at mesoscale level and guide decisions depending on the spatial configuration of local populations. The approach would apply and should be considered in any species where foraging distribution is colony‐specific and mortality risk varies spatially.     

Using complementary approaches to estimate survival of juvenile and adult Eastern Kingbirds

ABSTRACT Survival rates of young birds during the period between nest departure and their first breeding season is an important but difficult statistic to measure because of low natal site fidelity, especially for long‐distance migrants. From 2002 to 2008, we conducted a capture‐mark‐resight study of Eastern Kingbirds (Tyrannus tyrannus), a Nearctic‐Neotropical migrant, at Malheur National Wildlife Refuge, Oregon, to estimate juvenile (S_J) and adult (S_A) survival. The return rate of juvenile kingbirds was high (0.224) and not significantly different from program MARK's estimate of S_J (0.291). On average, and for both sexes, program MARK's estimate of S_A for birds banded as nestlings (0.64) was similar to that for birds first banded as adults (0.65). Enumeration methods and MARK yielded similar estimates of S_A, especially for males banded as adults. We attribute the similarity of resighting rate of birds banded as nestlings to S_J and the similarity of S_A estimated using program MARK and by enumeration methods to the high site fidelity of most juvenile and adult kingbirds at our ecologically isolated study site. An independent estimate of S_J calculated using local estimates of population growth and average annual production of young per year suggested that true S_J and S_A were probably slightly higher than program MARK's estimates because of undocumented permanent emigration, especially of birds first banded as adults. Demographic balancing suggests that true S_A and S_J were roughly 0.70 and 0.30, respectively. In general, our multiple estimates of S_J were similar. However, program MARK estimates of S_J tended to be lower than those produced by demographic balancing. Because of the difficulty in differentiating permanent emigration and mortality, and the affect it has on empirical estimates of survival, we urge researchers to use multiple methods of survival estimation, when possible, to validate the precision of their estimates.     

Age‐dependent breeding dispersal and adult survival within a metapopulation of Common Terns Sterna hirundo

Dispersal is increasingly recognized as a process of fundamental importance in population dynamics and other aspects of biology. Concurrently, interest in age‐dependent effects on survival, including actuarial senescence, has increased, especially in studies of long‐lived seabirds. Nevertheless, datasets necessary for studying dispersal and age‐dependent effects are few, as these require simultaneous data collection at two or more sites over many years. We conducted a 22‐year capture‐mark‐recapture study of Common Terns Sterna hirundo at three breeding colonies 10–26 km apart in Buzzards Bay, Massachusetts, USA. All birds in the study were of known age (range 2–28 years, median 7 years, n = 3290) and 77% were of known sex. Estimates of adult recapture, survival and breeding dispersal rates were obtained for all age‐classes from 2 to 20 years. The model that acquired 100% of the QAIC_c (Akaike's Information Criterion adjusted for small sample size and overdispersion) weight in our analysis included age‐specificity in all parameters but no relationship with sex. Our study may be the first to demonstrate age‐specificity in recapture, survival and breeding dispersal rates simultaneously, using a single model. Annual rates of breeding dispersal ranged from <0.01 to 0.27, with a population‐weighted mean of 0.065; they decreased with increasing distance between colony sites and, unexpectedly, increased with age. Breeding dispersal did not increase consistently after years with predation on adults or after an attempt to displace birds from an oiled site. Survival rates did not vary among sites or years. Annual adult survival increased from 0.80 in 2‐year‐old birds to a maximum of approximately 0.88 around age 8 years and then declined to 0.76 at age 20 years, yielding strong evidence for actuarial senescence. The peak annual survival rate of 0.88 is at the low end of other estimates for Common Tern and in the lower part of the range recorded for other terns, but total numbers in the three colonies increased seven‐fold during the study. This was part of a slower increase in the regional population, with net immigration into the study colonies. Our results demonstrate the biological significance of breeding dispersal in local population dynamics and age‐related effects on survival and dispersal from a metapopulation of a long‐lived seabird.     

Projecting demographic responses to climate change: adult and juvenile survival respond differently to direct and indirect effects of weather in a passerine population

Few studies have quantitatively projected changes in demography in response to climate change, yet doing so can provide important insights into the processes that may lead to population declines and changes in species distributions. Using a long‐term mark‐recapture data set, we examined the influence of multiple direct and indirect effects of weather on adult and juvenile survival for a population of Song Sparrows (Melospiza melodia) in California. We found evidence for a positive, direct effect of winter temperature on adult survival, and a positive, indirect effect of prior rainy season precipitation on juvenile survival, which was consistent with an effect of precipitation on food availability during the breeding season. We used these relationships, and climate projections of significantly warmer and slightly drier winter weather by the year 2100, to project a significant increase in mean adult survival (12–17%) and a slight decrease in mean juvenile survival (4–6%) under the B1 and A2 climate change scenarios. Together with results from previous studies on seasonal fecundity and postfledging survival in this population, we integrated these results in a population model and projected increases in the population growth rate under both climate change scenarios. Our results underscore the importance of considering multiple, direct, and indirect effects of weather throughout the annual cycle, as well as differences in the responses of each life stage to climate change. Projecting demographic responses to climate change can identify not only how populations will be affected by climate change but also indicate the demographic process(es) and specific mechanisms that may be responsible. This information can, in turn, inform climate change adaptation plans, help prioritize future research, and identify where limited conservation resources will be most effectively and efficiently spent.     

Survival of Afro‐Palaearctic passerine migrants in western Europe and the impacts of seasonal weather variables

Populations of migratory songbirds in western Europe show considerable variation in population trends between species and regions. The demographic and environmental causes of these large‐scale patterns are poorly understood. Using data from Constant Effort mist‐netting studies, we investigated relationships between changes in abundance, adult survival and seasonal weather conditions among 35 western European populations of eight species of migratory warblers (Sylviidae). We used cross‐species and within‐species comparisons to assess whether annual variation in survival was correlated with weather conditions during passage or winter. We estimated survival using CJS mark‐recapture models accounting for variation in the proportion of transient individuals and recapture rates. Species wintering in the humid bioclimatic zone of western Africa had significantly higher annual survival probabilities than species wintering in the arid bioclimatic zone of Africa (the Sahel). Rainfall in the Sahel was positively correlated with survival in at least some populations of five species. We found substantially fewer significant relationships with indices of weather during the autumn and spring passage periods, which may be due to the use of broad‐scale indices. Annual population changes were correlated with adult survival in all of our study species, although species undergoing widespread declines showed the weakest relationships.     

Apparent survival of tropical birds in a wet,premontane forest in Costa Rica

Despite the importance of tropical birds in the development of life history theory, we lack information about demographic rates and drivers of population dynamics for most species. We used a 7‐year (2007–2013) capture‐mark‐recapture dataset from an exceptionally wet premontane forest at mid‐elevation in Costa Rica to estimate apparent survival for seven species of tropical passerines. For four of these species, we provide the first published demographic parameters. Recapture probabilities ranged from 0.21 to 0.53, and annual estimates of apparent survival varied from 0.23 to 1.00. We also assessed the consequences of inter‐annual variation in rainfall on demographic rates. Our results are consistent with inter‐annual rainfall increasing estimates of apparent survival for two species and decreasing estimates for three species. For the three species where we could compare our estimates of apparent survival to estimates from drier regions, our estimates were not consistently higher or lower than those published previously. The temporal and spatial variability in demographic rates we document within and among species highlights the difficulties of generalizing life history characteristics across broad biogeographic gradients. Most importantly, this work emphasizes the context‐specific role of precipitation in shaping tropical avian demographic rates and underscores the need for mechanistic studies of environmental drivers of tropical life histories.     

High juvenile annual survival probabilities in Southern Rockhopper Penguins Eudyptes chrysocome are independent of individual fledging traits

Juvenile survival is an important demographic parameter. Southern Rockhopper Penguins Eudyptes chrysocome have undergone a dramatic population decline in the past century across their distribution, but the demographic processes are poorly understood. To estimate juvenile annual survival probabilities, Rockhopper Penguin chicks from two cohorts on New Island, Falkland Islands, were marked with transponders and recorded in subsequent years using an automated gateway. We first estimated annual survival and detection probabilities using a Cormack‐Jolly‐Seber (CJS) model, and found that both probabilities were extremely high (81% in the first and 98% in the second, third and fourth years of life), even in comparison with adult birds. Because detection probability after 3 years was effectively 1, and our sample size (n = 114) was too small to explore the effects of individual traits on survival in a CJS model, we assessed whether sex, cohort, body mass and laying sequence affected whether juveniles returned to the colony during their first 3 years of life using a simple generalized linear model that assumed perfect detection. Juveniles from the first cohort and males showed a higher return probability than juveniles from the second cohort and females. There was no clear effect of fledging body mass on return rate, probably related to the favourable environmental conditions during the study period. The laying sequence did not markedly affect the return probability of chicks, indicating that, once fledged, first‐laid A‐chicks have the same probability to return as second‐laid B‐chicks despite a much larger initial maternal investment in B‐eggs in this species. This study demonstrates extraordinarily high juvenile survival probabilities and will help to understand the recent changes in the population dynamics of the Falkland Islands Southern Rockhopper Penguins.     

Seasonal abundance and adult survival of bottlenose dolphins (Tursiops truncatus) in a community that cooperatively forages with fishermen in southern Brazil

A subgroup of a population of Tursiops truncatus in southern Brazil is known for a cooperative behavior with artisanal fishermen whereby the dolphins shoal fish towards net‐casting fishermen. Combining photo‐identification data collected between September 2007 and 2009 with mark‐recapture and Pollock's robust design models, we assessed abundance within seasons and survival and temporary emigration rates of dolphins between seasons. We also reanalyzed a previous data set collected during 1989–1991, and Cormack‐Jolly‐Seber models were applied to estimate survival rates for each of the study periods. The abundance of marked “cooperative” dolphins varied between seasons from 18 (CI: 17–24) to 21 (CI: 20–24). The total abundance varied from 59 in the winter of 2008 (CI: 49–72) to 50 in the autumn of 2009 (CI: 40–62). The annual adult survival was estimated to be 0.917 (CI: 0.876–0.961), close to that estimated from data collected in the 1990s (0.941; CI: 0.888–0.998). The emigration probability was low (0.031; CI: 0.011–0.084) and different capture probabilities between the “cooperative” and “noncooperative” dolphins indicated a degree of behavioral segregation. The precision of our estimates is likely to provide sufficient power to detect population change, but we recommend a precautionary management approach to protect this vulnerable dolphin community and its unique cooperative feeding tradition.     

No effect of blood sampling or phytohaemagglutinin injection on postfledging survival in a wild songbird

The injection of phytohaemagglutinin (PHA) and sampling of blood are widely used in studies of wild vertebrates to assess components of immune and endocrine function and health state and to obtain genetic material. Despite the pervasive use of these techniques in the life sciences, their potential effects on survival are rarely considered. For example, whether injection of the immunogen PHA into body parts critical for locomotion (e.g., the prepatagium, or wing web, in birds) affects survival has not been tested. Here, we test whether injection of PHA into the wing web and blood sampling from nestling house wrens affects their subsequent recruitment and survival as breeding adults. Capture‐mark‐recapture analysis on a large sample of young (N = 20,152 fledglings from 3959 broods) treated over 10 years revealed that neither PHA injection nor blood sampling affected individual survival and detection probability. Recruitment as a breeder varied among years, but this variation was not attributable to sampling effort, or the percent of all adults identified at the nest during a given year. Variation in the percent of adults identified was primarily attributable to the effect of nest depredation on our ability to capture nesting pairs. Our results indicating lack of an effect of blood sampling and immune stimulation on survival are encouraging, but we recommend further work to assess the potential negative effects of all commonly used techniques on the survival of study subjects in the wild, including the potential costs associated with mounting various immunological responses.     

Climate anomalies affect annual survival rates of swifts wintering in sub‐Saharan Africa

Several species of migratory swifts breed in the Western Palearctic, but they differ in reproductive traits and nonbreeding areas explored in Africa. We examined survival and recapture probabilities of two species of swifts by capture–mark–recapture data collected in northern Italy (Pallid Swift Apus pallidus in Carmagnola, Turin, and Common Swift Apus apus in Guiglia, Modena) in the breeding season (May–July). Apparent survival rates were relatively high (>71%), comparable to other studies of European swifts, but showed marked annual variations. We used geolocators to establish the exact wintering areas of birds breeding in our study colonies. Common Swifts explored the Sahel zone during migration and spent the winter in SE Africa, while the Pallid Swifts remained in the Sahel zone for a longer time, shifting locations southeast down to Cameroun and Nigeria later in winter. These movements followed the seasonal rains from north to south (October to December). In both species, we found large yearly differences in survival probabilities related to different climatic indices. In the Pallid Swift, wintering in Western Africa, the Sahel rainfall index best explained survival, with driest seasons associated with reduced survival. In the Common Swift, wintering in SE Africa, the El Niño–Southern Oscillation (ENSO) cycle performed significantly better than Sahel rainfall or North Atlantic Oscillation (NAO). Extreme events and precipitation anomalies in Eastern Africa during La Niña events resulted in reduced survival probabilities in Common Swifts. Our study shows that the two species of swifts have similar average annual survival, but their survival varies between years and is strongly affected by different climatic drivers associated with their respective wintering areas. This finding could suggest important ecological diversification that should be taken into account when comparing survival and area use of similar species that migrate between temperate breeding areas and tropical wintering areas.     

Effects of river temperature and climate warming on stock‐specific survival of adult migrating Fraser River sockeye salmon (Oncorhynchus nerka)

Mean summer water temperatures in the Fraser River (British Columbia, Canada) have increased by ～1.5 °C since the 1950s. In recent years, record high river temperatures during spawning migrations of Fraser River sockeye salmon (Oncorhynchus nerka) have been associated with high mortality events, raising concerns about long‐term viability of the numerous natal stocks faced with climate warming. In this study, the effect of freshwater thermal experience on spawning migration survival was estimated by fitting capture–recapture models to telemetry data collected for 1474 adults (captured in either the ocean or river between 2002 and 2007) from four Fraser River sockeye salmon stock‐aggregates (Chilko, Quesnel, Stellako‐Late Stuart and Adams). Survival of Adams sockeye salmon was the most impacted by warm temperatures encountered in the lower river, followed by that of Stellako‐Late Stuart and Quesnel. In contrast, survival of Chilko fish was insensitive to the encountered river temperature. In all stocks, in‐river survival of ocean‐captured sockeye salmon was higher than that of river‐captured fish and, generally, the difference was more pronounced under warm temperatures. The survival–temperature relationships for ocean‐captured fish were used to predict historic (1961–1990) and future (2010–2099) survival under simulated lower river thermal experiences for the Quesnel, Stellako‐Late Stuart and Adams stocks. A decrease of 9–16% in survival of all these stocks was predicted by the end of the century if the Fraser River continues to warm as expected. However, the decrease in future survival of Adams sockeye salmon would occur only if fish continue to enter the river abnormally early, towards warmer periods of the summer, as they have done since 1995. The survival estimates and predictions presented here are likely optimistic and emphasize the need to consider stock‐specific responses to temperature and climate warming into fisheries management and conservation strategies.     

Effect of diet on survival,in the laboratory and the field,of sterile male Mediterranean fruit flies

To improve the effectiveness of the sterile insect technique against the Mediterranean fruit fly, Ceratitis capitata (Wiedemann) (Diptera: Tephritidae), our objectives in this study were two‐fold. First, to evaluate the ability of sterile males of the Vienna‐8 strain to survive starvation, we compared them to wild males under laboratory conditions. The second objective was to determine the effect of protein‐rich nutrition on sterile male fly survival, under starvation conditions in the laboratory, under semi‐natural conditions in a field enclosure, and under natural conditions in the open field. Therefore, we released marked sterile flies of the two diet regimes, protein‐fed or protein‐deprived, and monitored their survival by recapturing them after 4, 6, and 7 days. In the laboratory, wild males endured starvation significantly better than sterile ones and protein addition to sterile fly diet resulted in even greater reduced capability to endure starvation. On the other hand, the addition of protein to sterile‐male diet did not affect their ability to survive in a field enclosure or in the open field. We conclude that under natural conditions, where food is available, sterile male fly survival is unaffected by protein‐rich pre‐release diet.     

Vertical and horizontal movements of Fuller's rose weevil (Coleoptera: Curculionidae) in Australian citrus groves

Fuller's rose weevil (FRW; Asynonychus cervinus (Boheman)) is a pest of quarantine concern in many Asian markets for Australian citrus. We investigated vertical (tree climbing) and horizontal movements (inter‐tree movements) of FRW adults in a citrus orchard during 2009–2011 with mark‐recapture experiments. Trunk traps were used to intercept tree‐climbing adults. The results showed that the majority of the adults released on the foliage stayed in the canopy, however considerable numbers did drop and some dropped multiple times. The rate of drops appeared to be fixed, independent of how many times the weevils had dropped before. Inter‐tree movements of adults in the canopy occurred mainly within rows, probably because of touching foliage between within‐row neighboring trees. Movement directions on the ground were more random, when reliance on touching foliage was not necessary. According to fitted dispersal equations, the average dispersal distance was 38 m for weevils released on foliage and 127 m for those released on the ground. Trunk banding with chemicals is an important control tactic for FRW, and based on our results, chemicals should be applied at a high enough rate (or in a wide enough band) to ensure the weevils are killed by a single crossing. For maximum reductions of FRW populations in citrus, control actions should be applied not only to infested blocks but also to neighboring blocks in the orchard.     

Evidence of reduced individual heterogeneity in adult survival of long‐lived species

The canalization hypothesis postulates that the rate at which trait variation generates variation in the average individual fitness in a population determines how buffered traits are against environmental and genetic factors. The ranking of a species on the slow‐fast continuum – the covariation among life‐history traits describing species‐specific life cycles along a gradient going from a long life, slow maturity, and low annual reproductive output, to a short life, fast maturity, and high annual reproductive output – strongly correlates with the relative fitness impact of a given amount of variation in adult survival. Under the canalization hypothesis, long‐lived species are thus expected to display less individual heterogeneity in survival at the onset of adulthood, when reproductive values peak, than short‐lived species. We tested this life‐history prediction by analysing long‐term time series of individual‐based data in nine species of birds and mammals using capture‐recapture models. We found that individual heterogeneity in survival was higher in species with short‐generation time (< 3 years) than in species with long generation time (> 4 years). Our findings provide the first piece of empirical evidence for the canalization hypothesis at the individual level from the wild.     

Effect of sex and bright coloration on survival and predator‐induced wing damage in an aposematic lantern fly with startle display

1. Aposematic coloration in prey promotes its survival by conspicuously advertising unpalatability to predators. Although classical examples of aposematic signals involve constant presentation of a signal at a distance, some animals suddenly display warning colours only when they are attacked. 2. Characteristics of body parts suddenly displayed, such as conspicuous coloration or eyespot pattern, may increase the survival of the prey by startling the predator, and/or by signalling unpalatability to the predators at the moment of attack. 3. The adaptive value of such colour patterns suddenly displayed by unpalatable prey has not been studied. We experimentally blackened the red patch in the conspicuous red–white–black hindwing pattern displayed by an unpalatable insect Lycorma delicatula White (Hemiptera: Fulgoridae) in response to predator's attack. 4. There was no evidence that the presence of the red patch increased prey survival over several weeks. We hypothesise that predators generalised from the red–white–black patches on the hindwings of unpalatable L. delicatula to any similar wing display as a signal of unpalatability. Because a higher proportion of males than females stay put at their resting sites, displaying their wings in response to repeated attacks by predators, wing damage was more frequent in males than in females. 5. To our knowledge, this is the first experimental test of an adaptive role of aposematic signals presented by unpalatable prey during sudden displays triggered by direct predatory attack.     

Variation in fine‐scale genetic structure and local dispersal patterns between peripheral populations of a South American passerine bird

The distribution of suitable habitat influences natal and breeding dispersal at small spatial scales, resulting in strong microgeographic genetic structure. Although environmental variation can promote interpopulation differences in dispersal behavior and local spatial patterns, the effects of distinct ecological conditions on within‐species variation in dispersal strategies and in fine‐scale genetic structure remain poorly understood. We studied local dispersal and fine‐scale genetic structure in the thorn‐tailed rayadito (Aphrastura spinicauda), a South American bird that breeds along a wide latitudinal gradient. We combine capture‐mark‐recapture data from eight breeding seasons and molecular genetics to compare two peripheral populations with contrasting environments in Chile: Navarino Island, a continuous and low density habitat, and Fray Jorge National Park, a fragmented, densely populated and more stressful environment. Natal dispersal showed no sex bias in Navarino but was female‐biased in the more dense population in Fray Jorge. In the latter, male movements were restricted, and some birds seemed to skip breeding in their first year, suggesting habitat saturation. Breeding dispersal was limited in both populations, with males being more philopatric than females. Spatial genetic autocorrelation analyzes using 13 polymorphic microsatellite loci confirmed the observed dispersal patterns: a fine‐scale genetic structure was only detectable for males in Fray Jorge for distances up to 450 m. Furthermore, two‐dimensional autocorrelation analyzes and estimates of genetic relatedness indicated that related males tended to be spatially clustered in this population. Our study shows evidence for context‐dependent variation in natal dispersal and corresponding local genetic structure in peripheral populations of this bird. It seems likely that the costs of dispersal are higher in the fragmented and higher density environment in Fray Jorge, particularly for males. The observed differences in microgeographic genetic structure for rayaditos might reflect the genetic consequences of population‐specific responses to contrasting environmental pressures near the range limits of its distribution.     

Factors Influencing Survival of Radiotagged and Banded Northern Bobwhites in Georgia

Abstract: Numerous studies of behavior and ecology of northern bobwhites (Colinus virginianus) have depended on radiotagging and telemetry for data collection. Excluding the presumably short-term effects of trapping, handling, and attaching radiotransmitters, researchers often assume that little bias is associated with estimating survival and behavioral parameters associated with this technique. However, researchers have not adequately examined these effects on organisms being investigated and have thus assumed demographic information obtained from such methods are valid. In light of this conjecture, it is imperative to evaluate methodological assumptions to ensure research is statistically valid and biologically meaningful. Therefore, we used Burnham's model and program MARK to analyze survival estimates of individually banded and radiotagged bobwhites during an 8-year period (1997–2004) consisting of 6,568 individuals (2,527 radiotagged) via combined analysis of mark—recapture, dead recovery (via harvest), and radiotelemetry data to test the effects of radiotransmitters on bobwhite survival. We also compared band—recapture survival estimates to Kaplan—Meier survival estimates, and we examined the effects of various other factors (e.g., temporal, spatial) on bobwhite survival. Based on Akaike's model selection criterion, the best model including the radiotransmitter covariate (Akaike's Information Criterion adjusted for small sample size bias and overdispersion relative value = 0.72) did not explain more of the variation in survival than models without this effect. Thus, we found the effect of radiotransmitters as negligible. Bobwhite survival varied relative to spatial (e.g., site), temporal (e.g., yr and season), and gender effects. Average annual survival for the 8-year period was 22.76% (1.50 SE) for banded-only and 21.72% (1.49 SE) for radiotagged birds. Survival rate varied annually, ranging from 12.42% (7.51 SE) to 37.16% (8.27 SE), and seasonally, ranging from 23.82% (2.71 SE) to 65.06% (3.23 SE); however, between group (banded-only, radiotagged) survival differences were still inconsequential. We conclude that for our study, radiotelemetry provided reliable survival estimates of an intensively managed bobwhite population, where supplemental food was provided, and this information provided useful data to make practical habitat management decisions. We believe that future radiotelemetry studies would benefit as a whole if researchers conducted similar analyses prior to presenting their results from radiotelemetry data, especially for populations that are more food limited.     

Estimability of migration survival rates from integrated breeding and winter capture–recapture data

Long‐distance migration is a common phenomenon across the animal kingdom but the scale of annual migratory movements has made it difficult for researchers to estimate survival rates during these periods of the annual cycle. Estimating migration survival is particularly challenging for small‐bodied species that cannot carry satellite tags, a group that includes the vast majority of migratory species. When capture–recapture data are available for linked breeding and non‐breeding populations, estimation of overall migration survival is possible but current methods do not allow separate estimation of spring and autumn survival rates. Recent development of a Bayesian integrated survival model has provided a method to separately estimate the latent spring and autumn survival rates using capture–recapture data, though the accuracy and precision of these estimates has not been formally tested. Here, I used simulated data to explore the estimability of migration survival rates using this model. Under a variety of biologically realistic scenarios, I demonstrate that spring and autumn migration survival can be estimated from the integrated survival model, though estimates are biased toward the overall migration survival probability. The direction and magnitude of this bias are influenced by the relative difference in spring and autumn survival rates as well as the degree of annual variation in these rates. The inclusion of covariates can improve the model's performance, especially when annual variation in migration survival rates is low. Migration survival rates can be estimated from relatively short time series (4–5 years), but bias and precision of estimates are improved when longer time series (10–12 years) are available. The ability to estimate seasonal survival rates of small, migratory organisms opens the door to advancing our understanding of the ecology and conservation of these species. Application of this method will enable researchers to better understand when mortality occurs across the annual cycle and how the migratory periods contribute to population dynamics. Integrating summer and winter capture data requires knowledge of the migratory connectivity of sampled populations and therefore efforts to simultaneously collect both survival and tracking data should be a high priority, especially for species of conservation concern.     

Male‐skewed adult sex ratio,survival, mating opportunity and annual productivity in the Snowy Plover Charadrius alexandrinus

Sex differences in adult mortality may be responsible for male‐skewed adult sex ratios and male‐skewed parental care in some birds. Because a surplus of breeding males has been reported in serially polyandrous populations of Snowy Plover Charadrius alexandrinus, we examined sex ratio, early‐season nesting opportunities, adult survival and annual reproductive success of a Snowy Plover population at Monterey Bay, California. We tested the hypotheses that male adult survival was greater than female survival and that a sex difference in adult survival led to a skewed adult sex ratio, different mating opportunities and different annual productivity between the sexes. Virtually all females left chicks from their first broods to the care of the male and re‐nested with a new mate. As a result, females had time to parent three successful nesting attempts during the lengthy breeding season, whereas males had time for only two successful attempts. Among years, the median population of nesting Plovers was 96 males and 84 females (median difference = 9), resulting in one extra male per eight pairs. The number of potential breeders without mates during the early nesting period each year was higher in males than in females. Adult male survival (0.734 ± 0.028 se) was higher than female survival (0.693 ± 0.030 se) in top‐ranked models. Annually, females parented more successful clutches and fledged more chicks than their first mates of the season. Our results suggest that in C. alexandrinus a sex difference in adult survival results in a male‐skewed sex ratio, which creates more nesting opportunities and greater annual productivity for females than for males.