Estimating survival rates from recoveries of birds ringed as young: A case study

Models for analysis of survival rates from recoveries of birds ringed as young were evaluated for lesser snow geese ringed on the breeding grounds at La Perouse Bay, Manitoba from 1970 to 1992. Analyses of birds ringed as adults and young indicate recovery rates were lower for young than for older birds and varied independently in the two age classes. Adult survival increased over the study, while that of immatures decreased. Using only recoveries of birds ringed as young, models incorporating observed variation in recovery rates were not identifiable. Many sets of constraints could be selected to make the model identifiable, but there was no objective way to distinguish among them, and each could lead to different conclusions. The observed changes in survival rates could not be estimated. Previous analyses have shown that models for birds ringed as young can be used to test even complex variation in survival rates, provided that recovery rates do not vary with age. Unfortunately, as shown by the geese, this assumption may often be violated and can only be tested with additional data, preferably from birds recaptured or ringed as sub-adults or adults.     

Estimating survival rates from recoveries of birds ringed as young: a case study

Models for analysis of survival rates from recoveries of birds ringed as young were evaluated for lesser snow geese ringed on the breeding grounds at La Perouse Bay, Manitoba from 1970 to 1992. Analyses of birds ringed as adults and young indicate recovery rates were lower for young than for older birds and varied independently in the two age classes. Adult survival increased over the study, while that of immatures decreased. Using only recoveries of birds ringed as young, models incorporating observed variation in recovery rates were not identifiable. Many sets of constraints could be selected to make the model identifiable, but there was no objective way to distinguish among them, and each could lead to different conclusions. The observed changes in survival rates could not be estimated. Previous analyses have shown that models for birds ringed as young can be used to test even complex variation in survival rates, provided that recovery rates do not vary with age. Unfortunately, as shown by the geese, this assumption may often be violated and can only be tested with additional data, preferably from birds recaptured or ringed as sub-adults or adults.     

Estimating age-specific survival rates of tawny owls--recaptures versus recoveries

We compared estimates of annual survival rates of tawny owls ( Strix aluco ) ringed in southern Finland from several different sampling methods: recoveries of birds ringed as young; recaptures of birds ringed as young; recoveries of birds ringed as adults as well as young; combined recoveries and recaptures of birds ringed as young, and combined recoveries and recaptures of birds ringed as adults and young. From 1979 to 1998, 18 040 young owls were ringed, of which 983 were recaptured as breeders in subsequent years during this period, and 1764 were recovered dead at various locations. In addition, 1751 owls were ringed as adults, of which 612 were later recaptured and 199 were recovered dead. First-year survival rates estimated using only recoveries of birds ringed as young averaged 48%, while apparent survival rates estimated using only recaptures from birds ringed as young averaged 10-13%. Use of combined recapture-recovery models, or supplementary information from recoveries of birds ringed as adults, produced survival estimates of 30-37%. Survival estimates from young-recoveries-only models were biased high, because of violation of the assumption of constant recovery rates with age: birds dying in their first-year were one-third less likely to be found and reported than older birds. In contrast, recaptures-only models confounded emigration with mortality. Despite these differences in mean values, annual fluctuations in estimated first-year survival rates were similar with all models. Estimates of adult survival rates were similar with all models, while those for second-year birds were similar for all models except recaptures-only. These results highlight the potential biases associated with analysing either recaptures or recoveries alone of birds ringed as young, and the benefits of using combined data.     

Survival rates of breeding common gulls in Estonia

The aim of the present paper was to estimate the survival rates of breeding common gulls, using mark-recapture data, gathered in 1968-1983 (16 sampling periods) in Estonia. We analyzed effects of age (breeding experience), year and sex on survival. Every year we observed more than 90% of the breeders and ringed about 95% of the nestlings. Two samples of gulls were used in the analyses. The first sample (347 individuals) consisted of birds thought to be first-time breeders when first observed in colony. The second sample (1269 individuals) may contain birds which have nested earlier. In the first sample, we did not detect any influence of age and sex on survival, but time dependence was significant and can be explained by winter severity. In cold winters the survival was lower (0.865) than in normal (0.896) and warm (0.929) winters. We suspect that the age effect on survival rate remained undetected owing to sparse data. In the second sample, we detected age- and time-dependent survival for both sexes. We also found differences between the sexes in recapture probability (in both samples). This was probably caused by lower site tenacity of females.     

Transient animals in a resident population of snow geese: Local emigration or heterogeneity?

The estimation of survival rates from analysis of recapture of individually marked animals assumes that all individuals are equally likely to be re-encountered. This assumption is frequently violated in natural populations due to movements to and from the sampling area. We evaluated potential sources of heterogeneity using data from recaptures of 36000 individually marked female lesser snow geese, Anser c. caerulescens , from an expanding population in northern Manitoba, Canada. By stratifying individuals according to marking age and origin (hatched at the colony or not), we assessed the degree to which variation in apparent survival reflected permanent or temporary differences in emigration and effects of handling. In general, for birds ringed as adults, estimated apparent survival rates were significantly lower during the first year after ringing than in subsequent years. By comparing birds ringed as adults (classified by origin) with those ringed as goslings, we were able to demonstrate that these differences are not due to permanent emigration from the colony by transient individuals or heterogeneity of individual capture probability, but more likely reflect differences among individuals in their response to initial marking. Approximately 25% of birds permanently emigrate from the sampling area following marking.     

Return rates in studies of life history evolution: Are biases large?

Studies of life history evolution in passerine birds often depend on examination of annual survival probability of adult birds. Most studies rely on return rates (proportion of marked individuals released in one year that are recaptured in the next year) to estimate annual survival probability. Yet, return rate includes both the probability of survival and the probability of recapturing or resighting the bird in the next time interval. We use numerical estimation to illustrate the increasing bias in return rate as an estimator of annual survival probability as recapture/resighting probability decreases. Recapture/resighting probability is normally assumed to be high and relatively invariant for recapture/resighting studies of color-banded territorial birds. We tested this assumption through examination of 11 color-banding studies of passerines. These studies showed that recapture/resighting probabilities vary strongly and cannot be generalized as high. In short, return rates generally are poor estimators of annual survival probabilities and use of return rates may strongly bias relationships explored in comparative studies or bias results of experiments to test survival costs of reproduction. Recapture/resighting probabilities should be estimated in all studies that attempt to estimate annual survival probabilities.     

Transient animals in a resident population of snow geese: local emigration or heterogeneity?

The estimation of survival rates from analysis of recapture of individually marked animals assumes that all individuals are equally likely to be re-encountered. This assumption is frequently violated in natural populations due to movements to and from the sampling area. We evaluated potential sources of heterogeneity using data from recaptures of 36000 individually marked female lesser snow geese, Anser c. caerulescens , from an expanding population in northern Manitoba, Canada. By stratifying individuals according to marking age and origin (hatched at the colony or not), we assessed the degree to which variation in apparent survival reflected permanent or temporary differences in emigration and effects of handling. In general, for birds ringed as adults, estimated apparent survival rates were significantly lower during the first year after ringing than in subsequent years. By comparing birds ringed as adults (classified by origin) with those ringed as goslings, we were able to demonstrate that these differences are not due to permanent emigration from the colony by transient individuals or heterogeneity of individual capture probability, but more likely reflect differences among individuals in their response to initial marking. Approximately 25% of birds permanently emigrate from the sampling area following marking.     

Modelling postfledging survival and age-specific breeding probabilities in species with delayed maturity: A case study of Roseate Terns at Falkner Island,Connecticut

We modelled postfledging survival and age-specific breeding probabilities in endangered Roseate Terns ( Sterna dougallii ) at Falkner Island, Connecticut, USA using capture-recapture data from 1988-1998 of birds ringed as chicks and as adults. While no individuals bred as 2-year-olds during this period, about three-quarters of the young that survived and returned as 3-year-olds nested, and virtually all surviving birds had begun breeding by the time they reached 5 years of age. We found no evidence of temporal variation age of first breeding of birds from different cohorts. There was significant temporal variation in the annual survival of adults and the survival over the typical 3-year maturation period of prebreeding birds, with extremely low values for both groups from the 1991 breeding season. The estimated overwinter survival rate (0.62) for adults from 1991-1992 was about three-quarters the usual rate of about 0.83, but the low survival of fledglings from 1991 resulted in less than 25% of the otherwise expected number of young from that cohort returning as breeding birds; this suggests that fledglings suffered a greater proportional decrease in survival than did adults. The survival estimates of young from 1989 and 1990 show that these cohorts were not negatively influenced by the events that decimated the young from 1991, and the young from 1992 and 1993 had above-average survival estimates. The apparent decrease since 1996 in development of fidelity of new recruits to this site is suspected to be due mainly to nocturnal disturbance and predation of chicks causing low productivity.     

Disentangling the sources of variation in the survival of the European dipper

The population growth rate of the European dipper has been shown to decrease with winter temperature and population size. We examine here the demographic mechanism for this effect by analysing how these factors affect the survival rate. Using more than 20 years of capture-mark-recapture data (1974-1997) based on more than 4000 marked individuals, we perform analyses using open capture-mark-recapture models. This allowed us to estimate the annual apparent survival rates (probability of surviving and staying on the study site from one year to the next one) and the recapture probabilities. We partitioned the variance of the apparent survival rates into sampling variance and process variance using random effects models, and investigated which variables best accounted for temporal process variation. Adult males and females had similar apparent survival rates, with an average of 0.52 and a coefficient of variation of 40%. Chick apparent survival was lower, averaging 0.06 with a coefficient of variation of 42%. Eighty percent of the variance in apparent survival rates was explained by winter temperature and population size for adults and 48% by winter temperature for chicks. The process variance outweighed the sampling variance both for chick and adult survival rates, which explained that shrunk estimates obtained under random effects models were close to MLE estimates. A large proportion of the annual variation in the apparent survival rate of chicks appears to be explained by inter-year differences in dispersal rates.     

Disentangling the sources of variation in the survival of the European dipper

The population growth rate of the European dipper has been shown to decrease with winter temperature and population size. We examine here the demographic mechanism for this effect by analysing how these factors affect the survival rate. Using more than 20 years of capture-mark-recapture data (1974-1997) based on more than 4000 marked individuals, we perform analyses using open capture-mark-recapture models. This allowed us to estimate the annual apparent survival rates (probability of surviving and staying on the study site from one year to the next one) and the recapture probabilities. We partitioned the variance of the apparent survival rates into sampling variance and process variance using random effects models, and investigated which variables best accounted for temporal process variation. Adult males and females had similar apparent survival rates, with an average of 0.52 and a coefficient of variation of 40%. Chick apparent survival was lower, averaging 0.06 with a coefficient of variation of 42%. Eighty percent of the variance in apparent survival rates was explained by winter temperature and population size for adults and 48% by winter temperature for chicks. The process variance outweighed the sampling variance both for chick and adult survival rates, which explained that shrunk estimates obtained under random effects models were close to MLE estimates. A large proportion of the annual variation in the apparent survival rate of chicks appears to be explained by inter-year differences in dispersal rates.     

Testing the effect of conspecific reproductive success on dispersal and recruitment decisions in a colonial bird: Design issues

Factors affecting dispersal and recruitment in animal populations will play a prominent role in the dynamics of populations. This is particularly the case for subdivided populations where the dispersal of individuals among patches may lead to local extinction and 'rescue effects'. A long-term observational study carried out in Brittany, France, and involving colour-ringed Black-legged Kittiwakes (Rissa tridactyla) suggested that the reproductive success of conspecifics (or some social correlate) could be one important factor likely to affect dispersal and recruitment. By dispersing from patches where the local reproductive success was low and recruiting to patches where the local reproductive success was high, individual birds could track spatio-temporal variations in the quality of breeding patches (the quality of breeding patches can be affected by different factors, such as food availability, the presence of predators or ectoparasites, which can vary in space and time at different scales). Such an observational study may nevertheless have confounded the role of conspecific reproductive success with the effect of a correlated factor (e.g. the local activities of a predator). In other words, individuals may have been influenced directly by the factor responsible for the low local reproductive success or indirectly by the low success of their neighbours. Thus, an experimental approach was needed to address this question. Estimates of demographic parameters (other than reproductive success) and studies of the response of marked individuals to changes in their environment usually face problems associated with variability in the probability of detecting individuals and with nonindependence among events occurring on a local scale. Further, very few studies on dispersal have attempted to address the causal nature of relationships by experimentally manipulating factors. Here we present an experiment designed to test for an effect of local reproductive success of conspecifics on behavioural decisions of individuals regarding dispersal and recruitment. The experiment was carried out on Kittiwakes within a large seabird colony in northern Norway. It involved (i) the colour banding of several hundreds of birds; (ii) the manipulation (increase/decrease) of the local reproductive success of breeding groups on cliffpatches; and (iii) the detailed survey of attendance and activities of birds on these patches. It also involved the manipulation of the nest content of marked individuals breeding within these patches (individuals failing at the egg stage were expected to respond in terms of dispersal to the success of their neighbours). This allowed us to test whether a lower local reproductive success would lower (1) the attendance of breeders at the end of the breeding season; (2) the presence of prospecting birds; and (3) the proportion of failed breeders that came back to breed on the same patch the year after. In this paper, we discuss how we dealt with (I) the use of return rates to infer differences in dispersal rates; (II) the trade-off between sample sizes and local treatment levels; and (III) potential differences in detection probabilities among locations. We also present some results to illustrate the design and implementation of the experiment.     

Suggestions for future directions for studies of marked migratory landbirds from the perspective of a practitioner in population management and conservation

Significant population declines in landbird species have been documented recently from many areas of the earth, including Europe and North America. Identification of the major causes of these declines and effective management actions to reverse them is difficult, especially for populations of long-distance migrants that winter in tropical areas. Key-factor and sensitivity analyses of critical population parameters in the context of integrated population models provide one promising approach to solving these problems. Key population factors may include breeding productivity, first-year survival, recruitment of young, adult survival and permanent emigration of adults; each of these can be indexed or estimated using data from cooperative ringing programmes, but the usefulness of the indices or estimates is limited by deficiencies in the available data and limitations of the available models. Future methodological directions for ringing studies should include efforts to: (1) develop and implement techniques to distinguish young from adult birds through the first breeding season of the young birds; (2) implement radio-tracking to determine characteristics of dispersal of young birds and transient adults; and (3) implement increased ringing, DNA fingerprinting and stable-isotope analysis to determine correspondence of breeding and winter ranges. Future programme-related directions should include efforts to: (1) integrate multiple methods at individual sites to compare and validate the indices and estimates produced by the different methods; (2) develop cooperative programmes of winter-season mist-netting to generate mark-recapture data to estimate the seasonal components of survival; and (3) develop mutually compatible banding programmes in tropical countries. Future theoretical and analytical directions should include efforts to continue to develop, refine and utilize: (1) key-factor and sensitivity analyses to determine the major causes of population changes; (2) models for dispersal of young birds and transient adults to improve the usefulness of indices of the number of hatch-year and second-year birds; (3) models to determine the proportions of transients in Cormack-Jolly-Seber (CJS) mark-recapture analyses and to eliminate their effects on estimates of survival rate, population size and recruitment of residents; (4) integrated models of population processes that utilize data from multiple methods to provide estimates of first-year survival, recruitment rate of young and permanent emigration rate of adults, parameters that are difficult to obtain from a single method; (5) models to estimate seasonal components of survival to provide insights into the timing and causes of mortality; (6) models incorporating environmental variables and species-specific characteristics as covariates in CJS mark-recapture and key-factor analyses; (7) models for pooling and weighting data obtained from multiple sites in cooperative ringing projects; (8) models for identifying long-term trends in demographic parameters; and (9) techniques for selection of appropriate models. Finally, assumptions implicit in the use of indices of various demographic parameters need to be tested and field techniques need to be improved to increase the numbers of individuals marked and recaptured in order to allow more precise parameter estimation; this will increase the ability to test competing hypotheses of population dynamics from data gathered in ringing programmes.     

Survival rates of Hungarian sand martins and their relationship with Sahel rainfall

Since 1986, I have carried out an intensive field survey of 10 000-30 000 pairs of a sand martin ( Riparia riparia ) population. Direct survey of the size and distribution of the breeding population and estimation of adult survival rates by SURGE, based on extensive ringing data sets, allow us to analyze the effects of different environmental factors with high precision. The model selection showed that the S s+t , P t model, in which the survival rates differ by sex for adults and vary in parallel by year and the capture rate varies by year, fits the data. The adult females had a lower survival rate compared to the males. The capture rate could be modelled as a quotient of the number of captured birds and the number of breeding birds along the upper part of the river Tisza. The survival rates of adults were related to the rainfall of the southern Sahel, which has an important role in the extension of the winter foraging habitat in the Sahel. Although the severe decrease in the population size, which may reach 50%, coincided with a large decline in the adult survival rate, there was not a significant relation between the adult survival rate and population size during the studied period. The population recruitment by first breeders and immigrant-emigrant adults could have a key role in the determination of population size. In the case of the studied subpopulation along the river, which is a core of the Carpathian Bend population, the immigration-emigration of adults had an important effect on the population size. The significant difference between juvenile male and female dispersal indicates the importance of separate estimation of juvenile survival for the sexes in further studies.     

Costs of reproduction: Assessing responses to brood size manipulation on life-history and behavioural traits using multi-state capture-recapture models

Costs of reproduction are fundamental trade-offs shaping the evolution of life histories. There has been much interest, discussion and controversy about the nature and type of reproductive costs. The manipulation of reproductive effort (e.g. brood size manipulation) may alter not only life-history traits such as future adult survival rate and future reproductive effort, but also behavioural decisions affecting recapture/resighting and dispersal probabilities. We argue that many previous studies of the costs of reproduction may have erroneously concluded the existence or non-existence of such costs because of their use of local return rates to assess survival. In this paper, we take advantage of the modern multistate capture-recapture methods to highlight how the accurate assessment of the costs of reproduction requires incorporating not only recapture probability, but also behavioural 'state' variables, for example dispersal status and current reproductive investment. The inclusion of state-dependent decisions can radically alter the conclusions drawn regarding the costs of reproduction on future survival or reproductive investment. We illustrate this point by re-analysing data collected to address the question of the costs of reproduction in the collared flycatcher and the great tit. We discuss in some detail the methodological issues and implications of the analytical techniques.     

Optimal allocation of sample sizes between regular banding and radio-tagging for estimating annual survival and emigration rates

Many authors have shown that a combined analysis of data from two or more types of recapture survey brings advantages, such as the ability to provide more information about parameters of interest. For example, a combined analysis of annual resighting and monthly radio-telemetry data allows separate estimates of true survival and emigration rates, whereas only apparent survival can be estimated from the resighting data alone. For studies involving more than one type of survey, biologists should consider how to allocate the total budget to the surveys related to the different types of marks so that they will gain optimal information from the surveys. For example, since radio tags and subsequent monitoring are very costly, while leg bands are cheap, the biologists should try to balance costs with information obtained in deciding how many animals should receive radios. Given a total budget and specific costs, it is possible to determine the allocation of sample sizes to different types of marks in order to minimize the variance of parameters of interest, such as annual survival and emigration rates. In this paper, we propose a cost function for a study where all birds receive leg bands and a subset receives radio tags and all new releases occur at the start of the study. Using this cost function, we obtain the allocation of sample sizes to the two survey types that minimizes the standard error of survival rate estimates or, alternatively, the standard error of emigration rates. Given the proposed costs, we show that for high resighting probability, e.g. 0.6, tagging roughly 10-40% of birds with radios will give survival estimates with standard errors within the minimum range. Lower resighting rates will require a higher percentage of radioed birds. In addition, the proposed costs require tagging the maximum possible percentage of radioed birds to minimize the standard error of emigration estimates.     

Parameter redundancy in capture–recapture–recovery models

In principle it is possible to use recently derived procedures to determine whether or not all the parameters of particular complex ecological models can be estimated using classical methods of statistical inference. If it is not possible to estimate all the parameters a model is parameter redundant. Furthermore, one can investigate whether derived results hold for such models for all lengths of study, and also how the results might change for specific data sets. In this paper we show how to apply these approaches to entire families of capture–recapture and capture–recapture–recovery models. This results in comprehensive tables, providing the definitive parameter redundancy status for such models. Parameter redundancy can also be caused by the data rather than the model, and how to investigate this is demonstrated through two applications, one to recapture data on dippers, and one to recapture–recovery data on great cormorants.     

Multistate recapture models: Modelling incomplete individual histories

Multistate capture-recapture models are a natural generalization of the usual one-site recapture models. Similarly, individuals are sampled on discrete occasions, at which they may be captured or not. However, contrary to the one-site case, the individuals can move within a finite set of states between occasions. The growing interest in spatial aspects of population dynamics presently contributes to making multistate models a very promising tool for population biology. We review first the interest and the potential of multistate models, in particular when they are used with individual states as well as geographical sites. Multistate models indeed constitute canonical capture-recapture models for individual categorical covariates changing over time, and can be linked to longitudinal studies with missing data and models such as hidden Markov chains. Multistate models also provide a promising tool for handling heterogeneity of capture, provided states related to capturability can be defined and used. Such an approach could be relevant for population size estimation in closed populations. Multistate models also constitute a natural framework for mixtures of information in individual history data. Presently, most models can be fit using program MARK. As an example, we present a canonical model for multisite accession to reproduction, which fully generalizes a classical one-site model. In the generalization proposed, one can estimate simultaneously age-dependent rates of accession to reproduction, natal and breeding dispersal. Finally, we discuss further generalizations - such as a multistate generalization of growth rate models and models for data where the state in which an individual is detected is known with uncertainty - and prospects for software development.     

Multistate recapture models: modelling incomplete individual histories

Multistate capture-recapture models are a natural generalization of the usual one-site recapture models. Similarly, individuals are sampled on discrete occasions, at which they may be captured or not. However, contrary to the one-site case, the individuals can move within a finite set of states between occasions. The growing interest in spatial aspects of population dynamics presently contributes to making multistate models a very promising tool for population biology. We review first the interest and the potential of multistate models, in particular when they are used with individual states as well as geographical sites. Multistate models indeed constitute canonical capture-recapture models for individual categorical covariates changing over time, and can be linked to longitudinal studies with missing data and models such as hidden Markov chains. Multistate models also provide a promising tool for handling heterogeneity of capture, provided states related to capturability can be defined and used. Such an approach could be relevant for population size estimation in closed populations. Multistate models also constitute a natural framework for mixtures of information in individual history data. Presently, most models can be fit using program MARK. As an example, we present a canonical model for multisite accession to reproduction, which fully generalizes a classical one-site model. In the generalization proposed, one can estimate simultaneously age-dependent rates of accession to reproduction, natal and breeding dispersal. Finally, we discuss further generalizations - such as a multistate generalization of growth rate models and models for data where the state in which an individual is detected is known with uncertainty - and prospects for software development.