Re-analysis of a banding study to test the effects of an experimental increase in bag limits of mourning doves

In 1966-1971, eastern US states with hunting seasons on mourning doves ( Zenaida macroura ) participated in a study designed to estimate the effects of bag limit increases on population survival rates. More than 400 000 adult and juvenile birds were banded and released during this period, and subsequent harvest and return of bands, together with total harvest estimates from mail and telephone surveys of hunters, provided the database for analysis. The original analysis used an ANOVA framework, and resulted in inferences of no effect of bag limit increase on population parameters (Hayne 1975). We used a logistic regression analysis to infer that the bag limit increase did not cause a biologically significant increase in harvest rate and thus the experiment could not provide any insight into the relationship between harvest and annual survival rates. Harvest rate estimates of breeding populations from geographical subregions were used as covariates in a Program MARK analysis and revealed an association between annual survival and harvest rates, although this relationship is potentially confounded by a latitudinal gradient in survival rates of dove populations. We discuss methodological problems encountered in the analysis of these data, and provide recommendations for future studies of the relationship between harvest and annual survival rates of mourning dove populations.     

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.     

Productivity indices and survival rate estimates from MAPS,a continent-wide programme of constant-effort mist-netting in North America

The Monitoring Avian Productivity and Survivorship (MAPS) programme is a cooperative effort to provide annual regional indices of adult population size and post-fledging productivity and estimates of adult survival rates from data pooled from a network of constant-effort mist-netting stations across North America. This paper provides an overview of the field and analytical methods currently employed by MAPS, a discussion of the assumptions underlying the use of these techniques, and a discussion of the validity of some of these assumptions based on data gathered during the first 5 years (1989-1993) of the programme, during which time it grew from 17 to 227 stations. Ageand species-specific differences in dispersal characteristics are important factors affecting the usefulness of the indices of adult population size and productivity derived from MAPS data. The presence of transients, heterogeneous capture probabilities among stations, and the large sample sizes required by models to deal effectively with these two considerations are important factors affecting the accuracy and precision of survival rate estimates derived from MAPS data. Important results from the first 5 years of MAPS are: (1) indices of adult population size derived from MAPS mist-netting data correlated well with analogous indices derived from point-count data collected at MAPS stations; (2) annual changes in productivity indices generated by MAPS were similar to analogous changes documented by direct nest monitoring and were generally as expected when compared to annual changes in weather during the breeding season; and (3) a model using between-year recaptures in Cormack-Jolly-Seber (CJS) mark-recapture analyses to estimate the proportion of residents among unmarked birds was found, for most tropical-wintering species sampled, to provide a better fit with the available data and more realistic and precise estimates of annual survival rates of resident birds than did standard CJS mark-recapture analyses. A detailed review of the statistical characteristics of MAPS data and a thorough evaluation of the field and analytical methods used in the MAPS programme are currently under way.     

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.     

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.     

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.     

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 turnover,movements and capture parameters of resting passerines in standardized capture-recapture studies

Jolly-Seber models A, B, D and 2 were used to investigate capture-recapture data. The standard Jolly-Seber model A (time-dependent survival phi and capture probability p ) fits capture-recapture data of migrating passerines. Captures from a long-term mist-netting study (Mettnau Peninsula, SW Germany) at a stop-over site were used to estimate stop-over length from survival rate between days and capture probability. For some data, model 2 could be used, indicating a termporary reduction in 'survival' rate. Application of models B and D gave poor results. The total number of birds stopping over, i.e. population size, was estimated from captures of 1-5 line transects of nets in the spatial trapping design. Behaviour, movements within the stop-over site, catchability and ecophysiological covariables such as moult, fat deposition and climatic parameters are likely to have strong influence on the estimation of capture parameters.     

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.     

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.     

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.     

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.     

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.     

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.     

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.     

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.