Estimating avian dispersal distances from data on ringed birds

Data from birds ringed as chicks and recaptured during subsequent breeding seasons provide information on avian natal dispersal distances. However, national patterns of ring reports are influenced by recapture rates as well as by dispersal rates. While an extensive methodology has been developed to study survival rates using models that correct for recapture rates, the same is not true for dispersal. Here, we present such a method, showing how corrections for spatial heterogeneity in recapture rate can be built into estimates of dispersal rates if detailed atlas data and ringing totals can be combined with extensive data on birds ringed as chicks and recaptured as breeding adults. We show how the method can be implemented in the software package SURVIV (White, 1992).     

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.     

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.     

Comparison of survival estimates obtained from three different methods of recapture in the same population of the great tit

Recaptures of marked birds can be used to estimate their survival. We suspect that the various methods of capture might not be equally suited for this purpose. In the long-term study of the great tit at our institute, recaptures were routinely made in three different ways: capture of parents feeding their young at the nest; capture of birds roosting in nest-boxes; mist-netting. We analyzed 20 years of captures from the study site at the Hoge Veluwe to obtain estimates of adult survival and capture rates from captures obtained by each method. The three sets of estimates differ and our analyses suggest that the best method was the capture of breeding birds, while mist-netting was the least suitable.     

The demography of the decline in the British willow warbler population

Since 1989, there has been a major and unprecedented decline in the breeding population of willow warblers ( Phylloscopus trochilus ) in southern Britain. Between 1986 and 1993 the numbers of willow warbler territories counted on monitoring plots declined by 47% in southern Britain, compared to a decline of 7% in northern Britain. Breeding densities of willow warblers are generally higher in the north and west of Britain, than in the south. Data from nest record cards provided evidence of only minor regional differences in breeding performance with a small but significant increase in the loss rate of nests during the nestling stage in 1989-1992 in southern Britain, compared with 1974-1988. Mark-recapture data collected at 18 constant effort sites and from one intensive study were used to estimate apparent survival rates of adults during the period 1987-1993. Program SURGE4 was used to test for differences in survival rates and recapture probabilities between years, sexes, sites and regions. Recapture probabilities differed between sites and between the sexes but not between years. Survival rates differed significantly between years (in southern Britain) but not between sexes or sites. In southern Britain, adult survival declined from 45% during 1987-1988 to 24% during 1991-1992, while in northern Britain there was no evidence that survival changed during the same period. Although the pattern of annual variation in survival differed between northern and southern Britain, this was due mainly to a much lower survival rate in southern Britain during 1991-1992. Declining survival rates of adult willow warblers have probably been a major cause of the observed population decline.     

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.     

Investigating the effects of hunting on the survival of British pigeons and doves by analysis of ringing recoveries

Recent changes in British hunting legislation have protected the stock dove Columba oenas since 1 October 1982, and removed protection from the collared dove Streptopelia decaocto after 1 April 1977; the woodpigeon Columba palumbus has remained legal quarry throughout. Ringing recoveries offer a means of comparing annual survival rates before and after the change in legislation. Care is needed in the construction of the recovery matrices to remove inhomogeneities in the data and ensure that each 'year' runs from the day of legislative change. Annual survival rates were estimated by maximum likelihood using multinomial models conditioned on the recoveries; there was no evidence of age- or time-related variation in reporting rates. The annual survival rate of the stock dove was constant, at 54 +/- 3%. For both the collared dove and the woodpigeon, estimates of survival rates in the first year after ringing were not consistent between birds ringed as young and those ringed as adults, but nevertheless averaged less before 1977 than after 1977; annual survival rates of birds that survived the first year after ringing did not differ between time periods, and were 64 +/- 2% for the collared dove and 61 +/- 2% for the woodpigeon. The proportion of recoveries reported shot or trapped was only 14% for the collared dove, compared with 79% for the woodpigeon, and remained constant before and after 1977 for both species; in the case of the stock dove, the proportion reported shot or trapped before and after 1982 fell from 70% to 34%. The change in quarry status had no effect on annual survivorship or population size of either the stock dove or the collared dove, while the woodpigeon has increased regularly in abundance despite heavy shooting.     

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.     

Survival of adult, territorial Longtailed Wagtails Motacilla clara : the effects of environmental factors and individual covariates

The Longtailed Wagtail is a non-migratory African passerine that is confined exclusively to small, fast-flowing rivers in a largely arboreal environment. The breeding adults hold permanent, life-long, linear territories in their riverine habitat and this makes it easy to locate colour-marked birds. They are confiding by nature and permit close approach, often to less than 10 m, and this allows their unique permutations of colourrings to be read. Using data from the 21 year period, 1 August 1978 to 31 July 1999, of a dozen territories it has been shown that the breeding territories have not changed at all, even though there has been a continual, but slow turnover of territory holders. A total of 109 territorial adult birds were monitored for a total of 1121 bird-quarters and survival was estimated for each of four quarters in a year. The average survival rate is estimated at 68.8% yr -1 (95% confidence limits: 63.3% to 69.3%) and this is high for such a small bird (approximately 20 g) and there have been some remarkably long-lived individuals, e.g. 10 to 12 years. In this paper, a generalized linear model is built of the survival of territorial adults. It is shown that bigger birds have a higher survival rate and that there are seasonal differences in survival that are ascribable to the cost of breeding and possibly cost of moult. There is an underlying long-term quadratic trend in survival that is related to increasing environmental degradation and decreasing chemical pollution.     

Survival of adult,territorial Longtailed Wagtails Motacilla clara : The effects of environmental factors and individual covariates

The Longtailed Wagtail is a non-migratory African passerine that is confined exclusively to small, fast-flowing rivers in a largely arboreal environment. The breeding adults hold permanent, life-long, linear territories in their riverine habitat and this makes it easy to locate colour-marked birds. They are confiding by nature and permit close approach, often to less than 10 m, and this allows their unique permutations of colourrings to be read. Using data from the 21 year period, 1 August 1978 to 31 July 1999, of a dozen territories it has been shown that the breeding territories have not changed at all, even though there has been a continual, but slow turnover of territory holders. A total of 109 territorial adult birds were monitored for a total of 1121 bird-quarters and survival was estimated for each of four quarters in a year. The average survival rate is estimated at 68.8% yr -1 (95% confidence limits: 63.3% to 69.3%) and this is high for such a small bird (approximately 20 g) and there have been some remarkably long-lived individuals, e.g. 10 to 12 years. In this paper, a generalized linear model is built of the survival of territorial adults. It is shown that bigger birds have a higher survival rate and that there are seasonal differences in survival that are ascribable to the cost of breeding and possibly cost of moult. There is an underlying long-term quadratic trend in survival that is related to increasing environmental degradation and decreasing chemical pollution.     

Asymmetric exchange between populations differing in habitat quality: A metapopulation study on the citril finch

The citril finch ( Serinus citrinella ) is a Cardueline finch restricted to the high mountains of western Europe. Since 1991 we have captured-recaptured about 6000 birds in two contrasting subpopulations located on the same mountain but separated by 5 km in distance. Citril finches, at the north-facing locality (La Vansa), rely more on Pine trees ( Pinus uncinata ) as their main food source, than birds at the south-facing locality (La Bofia), which rely more on herb seeds, which are of lower energetic content. Birds at La Vansa had higher body mass and fat score than those at La Bofia, suggesting that La Vansa was a site of higher-quality than La Bofia. By the use of a metapopulation approach and multistate models, we found that citril finches at the high-quality locality (La Vansa) showed higher survival rates than those at the low-quality one (La Bofia) (Vansa adults: φ = 0.42 - 0.04, juveniles: φ = 0.34 - 0.05; Bofia adults: φ = 0.35 - 0.04, juveniles: φ = 0.28 - 0.05). Dispersal was also asymmetric and higher for juvenile birds, with movement rates for juvenile citril finches from the low-quality to the higher-quality locality (Bofia to Vansa: é = 0.38 - 0.10) higher than the reverse (Vansa to Bofia: é = 0.09 - 0.03). We also investigated time-specific factors (e.g. meteorological data and fructification rate of Pinus ) as potential predictors of overall mortality and dispersal patterns. The results do not allow strong conclusions regarding the impact of these factors on survival and movement rates. Patterns of movement found in the Citril Finch between localities document a new model for the dispersal of species from low to high quality habitats, which we label of 'sources and pools'. This contrasts with currently accepted models of 'sources and sinks', in which movement is from high to low quality habitats, and 'Ideal Free Distributions', in which there is a balanced dispersal between habitats of different quality.     

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.     

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.     

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.     

Effects of neckbands on survival and fidelity of white-fronted and Canada geese captured as non-breeding adults

We conducted an experiment to examine the effect of neckbands, controlling for differences in sex, species and year of study (1991-1997), on probabilities of capture, survival, reporting, and fidelity in non-breeding small Canada ( Branta canadensis hutchinsi ) and white-fronted ( Anser albifrons frontalis ) geese. In Canada's central arctic, we systematically double-marked about half of the individuals from each species with neckbands and legbands, and we marked the other half only with legbands. We considered 48 a priori models that included combinations of sex, species, year, and neckband effects on the four population parameters produced by Burnham's (1993) model, using AIC for model selection. The four best approximating models each included a negative effect of neckbands on survival, and effect size varied among years. True survival probability of neckbanded birds annually ranged from 0.006 to 0.23 and 0.039 to 0.22 (Canada and white-fronted geese, respectively) lower than for conspecifics without neckbands. Changes in estimates of survival probability in neckbanded birds appeared to attenuate more recently, particularly in Canada Geese, a result that we suspect was related to lower retention rates of neckbands. We urge extreme caution in use of neckbands for estimation of certain population parameters, and discourage their use for estimation of unbiased survival probability in these two species.     

Population study and management of Atlantic Flyway Canada geese

Numbers of wintering Canada geese ( Branta canadensis ) in different regions of the Atlantic Flyway have significantly changed since the 1950s. During the 1980s, numbers increased in the mid-Atlantic and declined in the Chesapeake regions. A neckband study during the 1980s tested hypotheses that differential survival or movement caused changes in wintering numbers. The earlier studies and a reanalysis of the movement data using MSSURVIV indicated that the changes in wintering numbers did not result from differential survival or movement. A new neckband study, started in 1991, was designed to test hypotheses that the increases in wintering geese in the mid-Atlantic resulted from increasing numbers of resident geese and that the decline in migrants was related to higher harvest rates. Numbers of wintering geese continued to increase in the mid-Atlantic and declined in the Chesapeake region. Analyses of winter-banded, migrant geese indicated that 47 000 (SE = 14 000) geese per year moved from the mid-Atlantic to the Chesapeake region, thus rejecting the short-stopping hypothesis for this period. The probability of mid-Atlantic neckbanded resident geese remaining in the mid-Atlantic during winter was 0.988 (0.005). No differences in average survival were detected for migrants wintering in different regions during 1991-1994. Resident geese had higher survival than migrant geese in the Chesapeake region but not in the mid-Atlantic region. Harvest age ratios were low for Maryland and Quebec during the 1990s with record lows occurring during 1992. The only survival rate that could be estimated during the period of harvest restrictions was for 1992. Differences in survival were not detected between the periods of liberal harvest, 1985-1987, and the one year estimate under restrictive harvest. Because of the low age ratio, comparable numbers of adults were harvested during periods of restrictive and liberal regulations. Migrants appeared to be declining in all wintering regions, and residents appeared to be increasing. The different trends may be due to production differences.     

Modelling age variation in survival and reporting rates for recovery models

In this paper, we focus on models for recovery data from birds ringed as young. In some cases, it is important to be able to include in these models a degree of age variation in the reporting probability. For certain models this has been found, empirically, to result in completely flat likelihood surfaces, due to parameter redundancy. These models cannot then be fitted to the data, to produce unique parameter estimates. However, empirical evidence also exists that other models with such age variation can be fitted to data by maximum likelihood. Using the approach of Catchpole and Morgan (1996b), we can now identify which models in this area are parameter-redundant, and which are not. Models which are not parameter-redundant may still perform poorly in practice, and this is investigated through examples, involving both real and simulated data. The Akaike Information Criterion is found to select inappropriate models in a number of instances. The paper ends with guidelines for fitting models to data from birds ringed as young, when age dependence is expected in the reporting probability.