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.     

A model of the ring-recovery reporting process for the Cape Griffon Gyps coprotheres

By their very nature, statistical models are constructed on the basis of a number of simplifying assumptions. It is usual to assume that all the individuals in a 'group' or 'cohort' have similar survival, recovery or reporting probabilities. From a number of earlier studies of the Cape Griffon Gyps coprotheres in southern Africa, it is clear that there have been many violations of these assumptions of homogeneity. To get a better understanding of the process whereby a dead ringed bird is found and reported, an analysis was undertaken of 575 recoveries from 7130 individuals ringed as nestlings. From a series of univariate generalized linear models, it was found that the proportion of ringed birds reported dead varied with the following factors: (1) ring prefix (representing different grades and thicknesses of aluminium): there was considerable variation in reporting rate between cohorts fitted with different ring prefix series used; (2) metal type: birds fitted with monel metal rings were reported at a rate twice that of those bearing aluminium rings; (3) colour rings: recoveries of birds with colour rings were much more likely to be reported than birds with only a metal ring; (4) epoch: the reporting rate has increased steadily from the 1950s through to the mid-1980s. All of these factors are confounded and so a number of multivariate generalized linear models were constructed. It was found that the variations in the cohort-specific reporting rate could be described well by a model including factors for metal-ring type and the presence or absence of colour rings. Using the tougher monel metal ring along with a set of colour rings more than doubles the reporting rate and their continued use is strongly recommended for future studies. The year-to-year variations could be accounted for by this model but the colony of ringing did not enter the model. The models used were based on two assumptions: (i) the reporting rate was constant for all individuals within a given cohort and (ii) the recoveries were complete. It is argued that the results are congruent with these assumptions. There is now a clearer model of the manner in which the ring-recovery reporting process proceeds and this has opened the way to building a more realistic statistical model to estimate survival in the Cape Griffon.     

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.     

A model of the ring-recovery reporting process for the Cape Griffon Gyps coprotheres

By their very nature, statistical models are constructed on the basis of a number of simplifying assumptions. It is usual to assume that all the individuals in a 'group' or 'cohort' have similar survival, recovery or reporting probabilities. From a number of earlier studies of the Cape Griffon Gyps coprotheres in southern Africa, it is clear that there have been many violations of these assumptions of homogeneity. To get a better understanding of the process whereby a dead ringed bird is found and reported, an analysis was undertaken of 575 recoveries from 7130 individuals ringed as nestlings. From a series of univariate generalized linear models, it was found that the proportion of ringed birds reported dead varied with the following factors: (1) ring prefix (representing different grades and thicknesses of aluminium): there was considerable variation in reporting rate between cohorts fitted with different ring prefix series used; (2) metal type: birds fitted with monel metal rings were reported at a rate twice that of those bearing aluminium rings; (3) colour rings: recoveries of birds with colour rings were much more likely to be reported than birds with only a metal ring; (4) epoch: the reporting rate has increased steadily from the 1950s through to the mid-1980s. All of these factors are confounded and so a number of multivariate generalized linear models were constructed. It was found that the variations in the cohort-specific reporting rate could be described well by a model including factors for metal-ring type and the presence or absence of colour rings. Using the tougher monel metal ring along with a set of colour rings more than doubles the reporting rate and their continued use is strongly recommended for future studies. The year-to-year variations could be accounted for by this model but the colony of ringing did not enter the model. The models used were based on two assumptions: (i) the reporting rate was constant for all individuals within a given cohort and (ii) the recoveries were complete. It is argued that the results are congruent with these assumptions. There is now a clearer model of the manner in which the ring-recovery reporting process proceeds and this has opened the way to building a more realistic statistical model to estimate survival in the Cape Griffon.     

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.     

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.     

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 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).     

Delineating bird populations using ring recoveries

We delineate bird populations using cluster analysis to group ringing sites based on pairwise comparisons of recoveries. Clustering provides a quantitative (but non-unique) grouping that can be used to examine the relationships of bird distributions at both local and regional geographic scales. Clustering is based on similarity matrices composed of pairwise comparisons of recovery distributions from ringing sites. We demonstrate the method using mallard ( Anas platyrhynchos ) ring recoveries to group ringing sites in south-central Canada, and discuss the possibilities for these analyses for non-hunted species with few recoveries.     

Separation of hunting and natural mortality using ring-return models: an overview

Ring-recovery methodology has been widely used to estimate survival rates in multi-year ringing studies of wildlife and fish populations (Youngs & Robson, 1975; Brownie et al. , 1985). The Brownie et al. (1985) methodology is often used but its formulation does not account for the fact that rings may be returned in two ways. Sometimes hunters are solicited by a wildlife management officer or scientist and asked if they shot any ringed birds. Alternatively, a hunter may voluntarily report the ring to the Bird Banding Laboratory (US Fish and Wildlife Service, Laurel, MD, USA) as is requested on the ring. Because the Brownie et al. (1985) models only consider reported rings, Conroy (1985) and Conroy et al. (1989) generalized their models to permit solicited rings. Pollock et al. (1991) considered a very similar model for fish tagging models which might be combined with angler surveys. Pollock et al. (1994) showed how to apply their generalized formulation, with some modification to allow for crippling losses, to wildlife ringing studies. Provided an estimate of ring reporting rate is available, separation of hunting and natural mortality estimates is possible which provides important management information. Here we review this material and then discuss possible methods of estimating reporting rate which include: (1) reward ring studies; (2) use of planted rings; (3) hunter surveys; and (4) pre- and post-hunting season ringings. We compare and contrast the four methods in terms of their model assumptions and practicality. We also discuss the estimation of crippling loss using pre- and post-season ringing in combination with a reward ringing study to estimate reporting rate.     

Separation of hunting and natural mortality using ring-return models: An overview

Ring-recovery methodology has been widely used to estimate survival rates in multi-year ringing studies of wildlife and fish populations (Youngs & Robson, 1975; Brownie et al. , 1985). The Brownie et al. (1985) methodology is often used but its formulation does not account for the fact that rings may be returned in two ways. Sometimes hunters are solicited by a wildlife management officer or scientist and asked if they shot any ringed birds. Alternatively, a hunter may voluntarily report the ring to the Bird Banding Laboratory (US Fish and Wildlife Service, Laurel, MD, USA) as is requested on the ring. Because the Brownie et al. (1985) models only consider reported rings, Conroy (1985) and Conroy et al. (1989) generalized their models to permit solicited rings. Pollock et al. (1991) considered a very similar model for fish tagging models which might be combined with angler surveys. Pollock et al. (1994) showed how to apply their generalized formulation, with some modification to allow for crippling losses, to wildlife ringing studies. Provided an estimate of ring reporting rate is available, separation of hunting and natural mortality estimates is possible which provides important management information. Here we review this material and then discuss possible methods of estimating reporting rate which include: (1) reward ring studies; (2) use of planted rings; (3) hunter surveys; and (4) pre- and post-hunting season ringings. We compare and contrast the four methods in terms of their model assumptions and practicality. We also discuss the estimation of crippling loss using pre- and post-season ringing in combination with a reward ringing study to estimate reporting rate.     

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.     

Use of the Barker model in an experiment examining covariate effects on first-year survival in Ross's Geese ( Chen rossii ): A case study

The Barker model provides researchers with an opportunity to use three types of data for mark-recapture analyses - recaptures, recoveries, and resightings. This model structure maximizes use of encounter data and increases the precision of parameter estimates, provided the researcher has large amounts of resighting data. However, to our knowledge, this model has not been used for any published ringing studies. Our objective here is to report our use of the Barker model in covariate-dependent analyses that we conducted in Program MARK. In particular, we wanted to describe our experimental study design and discuss our analytical approach plus some logistical constraints we encountered while conducting a study of the effects of growth and parasites on survival of juvenile Ross's Geese. Birds were marked just before fledging, alternately injected with antiparasite drugs or a control, and then were re-encountered during migration and breeding in following years. Although the Barker model estimates seven parameters, our objectives focused on annual survival only, thus we considered all other parameters as nuisance terms. Therefore, we simplified our model structures by maintaining biological complexity on survival, while retaining a very basic structure on nuisance parameters. These analyses were conducted in a two-step approach where we used the most parsimonious model from nuisance parameter analyses as our starting model for analyses of covariate effects. This analytical approach also allowed us to minimize the long CPU times associated with the use of covariates in earlier versions of Program MARK. Resightings made up about 80% of our encounter history data, and simulations demonstrated that precision and bias of parameter estimates were minimally affected by this distribution. Overall, the main source of bias was that smaller goslings were too small to retain neckbands, yet were the birds that we predicted would have the lowest survival probability and highest probability for parasite effects. Consequently, we considered our results conservative. The largest constraint of our study design was the inability to partition survival into biologically meaningful periods to provide insight into the timing and mechanisms of mortality.     

MATLAB: An environment for analyzing ring-recovery and recapture data

The programming environment provided by MATLAB makes it easy to write code for the interactive analysis of data from ring-recovery and recapture studies. In particular, it is simple to write down a log-likelihood for any given model, to maximize this log-likelihood to estimate the parameters of the model and calculate their standard errors, to perform likelihood ratio and score test comparisons of models, and to compute goodness-of-fit statistics and examine residuals. Eagle is a package of simple MATLAB programs that provides an interactive environment for the analysis of ring-recovery data. Eagle will cope with any model of the Freeman-Morgan kind, where the survival and recovery probabilities may be specified as constant, time- or age-dependent, or as depending on one or more external covariates. This will be extended to handle data from recapture studies and from combined recovery and recapture studies. The package may also be used to analyze arbitrary models, at the expense of a small amount of extra MATLAB programming. A demonstration is given of use of Eagle to analyze data from a ring-recovery study on grey herons ( Ardea cinera ), with and without covariates. The use of the MATLAB programming environment is illustrated by fitting a simple model to some recapture data on herring gulls ( Larus argentatus ).     

Test for age-specificity in survival of the common tern

Much effort in life-history theory has been addressed to the dependence of life-history traits on age, especially the phenomenon of senescence and its evolution. Although senescent declines in survival are well documented in humans and in domestic and laboratory animals, evidence for their occurrence and importance in wild animal species remains limited and equivocal. Several recent papers have suggested that methodological issues may contribute to this problem, and have encouraged investigators to improve sampling designs and to analyse their data using recently developed approaches to modelling of capture-mark-recapture data. Here we report on a three-year, two-site, mark-recapture study of known-aged common terns (Sterna hirundo) in the north-eastern USA. The study was nested within a long-term ecological study in which large numbers of chicks had been banded in each year for > 25 years. We used a range of models to test the hypothesis of an influence of age on survival probability. We also tested for a possible influence of sex on survival. The cross-sectional design of the study (one year's parameter estimates) avoided the possible confounding of effects of age and time. The study was conducted at a time when one of the study sites was being colonized and numbers were increasing rapidly. We detected two-way movements between the sites and estimated movement probabilities in the year for which they could be modelled. We also obtained limited data on emigration from our study area to more distant sites. We found no evidence that survival depended on either sex or age, except that survival was lower among the youngest birds (ages 2-3 years). Despite the large number of birds included in the study (1599 known-aged birds, 2367 total), confidence limits on estimates of survival probability were wide, especially for the oldest age-classes, so that a slight decline in survival late in life could not have been detected. In addition, the cross-sectional design of this study meant that a decline in survival probability within individuals (actuarial senescence) could have been masked by heterogeneity in survival probability among individuals (mortality selection). This emphasizes the need for the development of modelling tools permitting separation of these two phenomena, valid under field conditions in which the recapture probabilities are less than one.     

How useful are recoveries of North American passerines for survival analyses?

Of 17 000 000 passerines ringed in North America from 1955 to 1984, only 0.4% were ever recovered. Only 62 species had more than 100 recoveries, of which only 26 had more than 500 recoveries. Preliminary analyses using stochastic recovery models for four of the species with more than 500 recoveries suggest estimates of mean annual adult survival can be fairly precise (CV < 4%), but estimates of immature survival are much less precise (CV > 8%). Comparable estimates may be possible for other species with more than 100 recoveries, depending on the temporal and geographic distribution of ringings and recoveries. Further analyses are required to determine whether these estimates may be biased by heterogeneity in survival or recovery rates.     

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.     

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.