Who are we sampling? Apparent survival differs between methods in a secretive species

Survival is a fundamental parameter in population dynamics with increasing importance in the management and conservation strategies of wildlife populations. Survival probability in vertebrates is usually estimated by live‐encounter data obtained by means of physical mark–capture–recapture protocols. Non‐invasive acoustic marking relying on individual‐specific features of signals has been alternatively applied as a marking technique, especially in secretive species. Nevertheless, to date no research has compared survival rate estimates obtained by acoustic and physical marking. We estimated half‐yearly and annual survival and recapture rates of a secretive and threatened passerine, the Dupont's lark Chersophilus duponti, using two separate live‐encounter data sets of males collected simultaneously by physical and acoustic marking in the same study area. The separate analysis of both methods led to different model structures, since transient individuals had to be accounted for in the acoustic marking but not in the physical marking data set. Furthermore, while reencounter probabilities did not differ between methods, survival estimates employing physical marking were lower than those obtained acoustically, especially between the postbreeding and the breeding period when the apparent survival of colour‐banded birds was twice as low as for acoustic marking. The combination of marking methods suggested the existence of different subsets of individuals differentially sampled within the population: whereas colour‐banded males seemed to represent the territorial fraction of the population, both resident and floater individuals were probably detected by acoustic marking. Using traditional mark–recapture methods exclusively could have misled our estimates of survival rates, potentially affecting prospective predictions of population dynamics. Acoustic marking has been poorly applied in mark–recapture studies, but might be a powerful complement to obtain accurate estimates of fundamental demographic parameters such as survival and dispersal.     

One step forward: contrasting the effects of Toe clipping and PIT tagging on frog survival and recapture probability

Amphibians have been declining worldwide and the comprehension of the threats that they face could be improved by using mark–recapture models to estimate vital rates of natural populations. Recently, the consequences of marking amphibians have been under discussion and the effects of toe clipping on survival are debatable, although it is still the most common technique for individually identifying amphibians. The passive integrated transponder (PIT tag) is an alternative technique, but comparisons among marking techniques in free‐ranging populations are still lacking. We compared these two marking techniques using mark–recapture models to estimate apparent survival and recapture probability of a neotropical population of the blacksmith tree frog, Hypsiboas faber. We tested the effects of marking technique and number of toe pads removed while controlling for sex. Survival was similar among groups, although slightly decreased from individuals with one toe pad removed, to individuals with two and three toe pads removed, and finally to PIT‐tagged individuals. No sex differences were detected. Recapture probability slightly increased with the number of toe pads removed and was the lowest for PIT‐tagged individuals. Sex was an important predictor for recapture probability, with males being nearly five times more likely to be recaptured. Potential negative effects of both techniques may include reduced locomotion and high stress levels. We recommend the use of covariates in models to better understand the effects of marking techniques on frogs. Accounting for the effect of the technique on the results should be considered, because most techniques may reduce survival. Based on our results, but also on logistical and cost issues associated with PIT tagging, we suggest the use of toe clipping with anurans like the blacksmith tree frog.     

Computer-Aided Pattern Recognition of Large Reptiles as a Noninvasive Application to Identify Individuals

For large species, the capture and handling of individuals in capture–mark–recapture studies introduces nonhuman animal welfare issues associated with handling, physical marking, and possible wounding due to tag loss. The use of photographic identification for these species offers an alternative and less invasive marking technique. This study investigated the opportunity offered by photo identification to individually mark individuals of a large reptile, the perentie (Varanus giganteus), in Australia and therefore avoid the stress of physically capturing and handling. Photographs submitted by a remotely located community were first validated to confirm whether perenties could be individually identified from their spots electronically using a computer program. Computer-aided selection of unique patterns was found to be appropriate for the identification of individuals and confirmed 38 individuals during the sampling period. The value of this approach is 2-fold: There is a benefit to animal welfare in that handling an animal is not required to capture him or her, thus reducing capture-related stress; and confirmation that photo identification of distinctive patterns of the perentie is valid and offers a useful option to identify individuals of this large species.     

Testing acoustic versus physical marking: two complementary methods for individual-based monitoring of elusive species

Individuals of some species differentiate each other on the basis of the acoustic features of their vocalizations, and this can be used in individual‐based population monitoring studies. No research has tested for the effectiveness of individual marking through voice recognition as compared to traditional monitoring methods relying on physical marks. We compared voice recognition and physical marking using the Dupont's lark Chersophilus duponti as a study species. This bird needs to be attracted with playback in order to be seen (or captured). We first demonstrated that the territorial calls from a sample of banded males were individually distinctive and constant over time by means of discriminant function analysis, which correctly classified 100% of marked males. Then, we applied similarity techniques on call spectrotemporal features to define a threshold value of similarity within banded individuals, to be combined with qualitative spectrogram inspection for the classification of all recorded birds. Eventually, we compared the voice and the capture samples, to test for differences in relation to re‐location rate, territory fidelity and dispersal movements both within and between years. Voice recognition was less time‐consuming than capture‐recapture method in the field, but it was useless for monitoring yearlings in call development stage. The two methods provided the same results in terms of territory fidelity and dispersal movements, but differed in re‐location rates, which were significantly greater in the case of voice recognition method. By means of physical captures we possibly trapped a large sample of young and silent floaters, with low probability of recapture or recording. This mismatch between methods could bias the estimates of annual survival, which strongly depend on re‐location rates. We suggest considering the two methods as complementary rather than alternatives for monitoring populations. Each technique offers unique information, and the two sources should be combined to provide correction factors that would eventually sharpen our knowledge on bird population ecology.     

Estimating Alpine ibex Capra ibex abundance from photographic sampling

Monitoring procedures for Alpine ibex Capra ibex are limited in habitats with reduced visibility and when physical capture and marking of the animals is not intended. Photographic sampling, involving using camera‐trap data and identifying ibex from natural markings, was adopted with capture‐recapture models to estimate the abundance of ibex in Austria. The software CAPTURE's model produced an average capture probability of 0.44 with an estimate of 34–51 ibex and a mean population size of 38 ibex. This first study showed the applicability of photographic capture‐recapture techniques to estimate the abundance of ibex based on their natural markings.     

The evaluation of passive integrated transponder (PIT) tags and visible implant elastomer (VIE) marks as new marking techniques for the bullhead

To test the reliability of PIT tags and VIE marks as new marking techniques for the bullhead Cottus gobio , different tagging treatments were assayed. The relatively high recapture rates suggest the applicability of both marking techniques for this small benthic fish species.     

Estimating abundance and population trends when detection is low and highly variable: A comparison of three methods for the Hermann's tortoise

Assessing population trends is a basic prerequisite to carrying out adequate conservation strategies. Selecting an appropriate method to monitor animal populations can be challenging, particularly for low-detection species such as reptiles. This study compares 3 detection-corrected abundance methods (capture–recapture, distance sampling, and N-mixture) used to assess population size of the threatened Hermann's tortoise. We used a single dataset of 432 adult tortoise observations collected at 118 sampling sites in the Plaine des Maures, southeastern France. We also used a dataset of 520 tortoise observations based on radiotelemetry data collected from 10 adult females to estimate and model the availability (g0) needed for distance sampling. We evaluated bias for N-mixture and capture–recapture, by using simulations based on different values of detection probabilities. Finally, we conducted a power analysis to estimate the ability of the 3 methods to detect changes in Hermann's tortoise abundances. The abundance estimations we obtained using distance sampling and N-mixture models were respectively 1.75 and 2.19 times less than those obtained using the capture–recapture method. Our results indicated that g0 was influenced by temperature variations and can differ for the same temperature on different days. Simulations showed that the N-mixture models provide unstable estimations for species with detection probabilities <0.5, whereas capture–recapture estimations were unbiased. Power analysis showed that none of the 3 methods were precise enough to detect slow population changes. We recommend that great care should be taken when implementing monitoring designs for species with large variation in activity rates and low detection probabilities. Although N-mixture models are easy to implement, we would not recommend using them in situations where the detection probability is very low at the risk of providing biased estimates. Among the 3 methods allowing estimation of tortoise abundances, capture–recapture should be preferred to assess population trends. © 2013 The Wildlife Society.     

Latent multinomial models for extended batch-mark data

Batch marking is common and useful for many capture–recapture studies where individual marks cannot be applied due to various constraints such as timing, cost, or marking difficulty. When batch marks are used, observed data are not individual capture histories but a set of counts including the numbers of individuals first marked, marked individuals that are recaptured, and individuals captured but released without being marked (applicable to some studies) on each capture occasion. Fitting traditional capture–recapture models to such data requires one to identify all possible sets of capture–recapture histories that may lead to the observed data, which is computationally infeasible even for a small number of capture occasions. In this paper, we propose a latent multinomial model to deal with such data, where the observed vector of counts is a non-invertible linear transformation of a latent vector that follows a multinomial distribution depending on model parameters. The latent multinomial model can be fitted efficiently through a saddlepoint approximation based maximum likelihood approach. The model framework is very flexible and can be applied to data collected with different study designs. Simulation studies indicate that reliable estimation results are obtained for all parameters of the proposed model. We apply the model to analysis of golden mantella data collected using batch marks in Central Madagascar.     

Are we overestimating recovery of sturgeon populations using mark/recapture surveys?

Mark‐recaptures studies are often conducted to monitor trends in sturgeon populations. However, many of these studies experience low recapture rates, minimal movement between marking‐recapture phases suggesting that sturgeon as a group are not conducive to mark‐recapture techniques. In this study, two mark‐recapture studies that were conducted differently were reviewed. A study was conducted on the Mattagami River using random nets set throughout the study area in both the mark and recapture phases. The other study was conducted on Lake of the Woods and marked sturgeon in tributaries during the spawning period and the recapture phase within the lake and river during the summer foraging period using random nets sets. Sturgeon's conduciveness to mark‐recapture studies was assessed on the Mattagami River mark‐recapture study by determining detection probability (p) using a hierarchical Bayesian model with data augmentation among three effects: individual effect, temporal effects, and behavioural response effects. Detection probability was constant over individuals and temporally suggesting model M₀ (Otis, Burnham, White, & Anderson, 1978 ) was suitable for lake sturgeon in the Mattagami River; only the M₀ would converge for the Lake of the Woods study. For this study, the assumption that “all individuals have the same probability of being captured during the marking phase” was believed to have been violated given approximately 16%–20% of adult Lake Sturgeon from a population spawn within a year. A population estimate accounting for p provided estimates 56% lower than calculated by a Chapman modification of the Peterson estimate for a closed population. Bias was believed to have been introduced as the Lake of the Woods population did not account for the non‐spawning adults that were encountered during the recapture phase and not vulnerable during the initial marking phase. This was not unique to the Lake of the Woods study as other sturgeon studies, especially multi‐year, assumes a closed population which potentially biased estimates and overestimated their recovery.     

Common species affects the utility of non‐invasive genetic monitoring of a cryptic endangered mammal: The bridled nailtail wallaby

Non‐invasive methods of monitoring wild populations (such as genotyping faeces or hair) are now widely used and advocated. The potential advantages of such methods over traditional direct monitoring (such as live capture) are that accuracy improves because sampling of non‐trappable individuals may be possible, species in difficult and remote terrain can be surveyed more efficiently, and disturbance to animals is minimal. Few studies have assessed the effects of interactions between species on remote sampling success. We test the use of non‐invasive monitoring for the cryptic, forest‐dwelling, solitary and endangered bridled nailtail wallaby (Onychogalea fraenata) that is sympatric with the ecologically similar and more common black‐striped wallaby (Macropus dorsalis). Six types of hair traps were tested for 3668 trap days, and hairs were caught with about a 10% success rate. Camera traps showed that baited hair traps targeted both wallaby species. We microscopically identified hair as bridled nailtail wallaby or black‐striped wallaby. We compared these hairs and their genotypes (using seven microsatellite loci) with known bridled nailtail wallaby hairs and genotypes derived from animal trapping. Trapped bridled nailtail wallaby hairs had characteristics that could be mistaken for black‐stripe wallaby hairs; characteristics were not diagnostic. Genetic assignment tests consistently differentiated the known bridled nailtail wallaby samples from identified black‐striped wallaby samples, however genetic overlap between most of the microsatellite markers means that they are not suitable for species identification of single samples, with the possible exception of the microsatellite locus B151. With similar trapping effort and within the same area, live‐capture mark‐recapture techniques estimated 40–60 individuals and non‐invasive methods only detected 14 genotypes. A species‐specific genetic marker would allow more efficient targeting of bridled nailtail wallaby samples and increase capture rates.     

Effect of Tooth Removal on Recaptures of Raccoons

Abstract: Researchers have extensively used mark—recapture techniques to obtain information on demographic parameters of wildlife populations. However, researchers have recognized that a number of factors can influence capture probabilities of wildlife species, which in turn can bias mark—recapture estimates of demographic parameters. Tooth extraction, which is a commonly used technique in studies of mesopredator species to obtain precise age estimates and to monitor the use of vaccine baits, is an aspect of animal handling that clearly might affect the recapture probability of individuals. However, the effect that tooth removal has on the individual recapture probabilities of wildlife species is unknown. During 2005, we trapped and marked 91 raccoons (Procyon lotor) in northern Indiana, USA, as part of a mark—recapture study designed specifically to determine if tooth extractions have an effect on recapture probabilities of individuals. We performed tooth extractions on 50% of the raccoons at the time of capture, and we attempted to balance tooth extractions with respect to sex and age of raccoons. We used logistic regression to model the effects of sex, age, and tooth removal on recapture probabilities, and we used Mann—Whitney U-tests to examine the effect of tooth removal on the number of times we recaptured individuals. The probability of recapture differed between sexes but did not differ as a function of tooth removal or among age classes. In addition, we failed to detect any difference in the mean number of times that we recaptured raccoons between the tooth removed and non—tooth-removed groups. Our results suggest that managers can use tooth extractions as an effective management tool without biasing population estimates or compromising other management objectives.     

Species richness estimation and determinants of species detectability in butterfly monitoring programmes

Abstract 1. Species richness is the most widely used biodiversity index, but can be hard to measure. Many species remain undetected, hence raw species counts will often underestimate true species richness. In contrast, capture–recapture methods estimate true species richness and correct for imperfect and varying detectability. 2. Detectability is a crucial quantity that provides the link between a species count and true species richness. For insects, it has hardly ever been estimated, although this is required for the interpretation of species counts. 3. In the Swiss butterfly monitoring programme about 100 transect routes are surveyed seven times a year using a highly standardised protocol. In July 2003, control observers made two additional surveys on 38 transects. Data from these 38 quadrats were analysed to see whether currently available capture–recapture models can provide quadrat‐specific estimates of species richness, and to estimate species detectability in relation to transect, observer, survey, region, and abundance. 4. Species richness over the entire season cannot be estimated using current capture–recapture methods. The species pool was open, preventing use of closed population models, and detectability varied by species, preventing use of current open population models. Assuming a closed species pool during two mid‐season (July) surveys, a Jackknife capture–recapture method was used that accounts for heterogeneity to estimate mean detectability and species richness. 5. In every case, more species were present than were counted. Mean species detectability was 0.61 (SE 0.01) with significant differences between observers (range 0.37–0.83). Species‐specific detection at time t+ 1 was then modelled for those species seen at t for three mid‐season surveys. Detectability averaged 0.50 (range 0.17–0.81) for individual species and 0.65, 0.44, and 0.42 for surveys. Abundant species were detected more easily, although this relationship explained only 5% of variation in species detectability. 6. These are important, although not entirely unexpected, results for species richness estimation of short‐lived animals. Raw counts of species may be misleading species richness indicators unless many surveys are conducted. Monitoring programmes should be calibrated, i.e. the assumption of constant detectability over dimensions of interest needs to be tested. The development of capture–recapture or similar models that can cope with both open populations and heterogeneous species detectability to estimate species richness should be a research priority.     

Demographic estimation methods for plants with unobservable life-states

Demographic estimation of vital parameters in plants with an unobservable dormant state is complicated, because time of death is not known. Conventional methods assume that death occurs at a particular time after a plant has last been seen aboveground but the consequences of assuming a particular duration of dormancy have never been tested. Capture–recapture methods do not make assumptions about time of death; however, problems with parameter estimability have not yet been resolved. To date, a critical comparative assessment of these methods is lacking. We analysed data from a 10 year study of Cleistes bifaria, a terrestrial orchid with frequent dormancy, and compared demographic estimates obtained by five varieties of the conventional methods, and two capture–recapture methods. All conventional methods produced spurious unity survival estimates for some years or for some states, and estimates of demographic rates sensitive to the time of death assumption. In contrast, capture–recapture methods are more parsimonious in terms of assumptions, are based on well founded theory and did not produce spurious estimates. In Cleistes, dormant episodes lasted for 1–4 years (mean 1.4, SD 0.74). The capture–recapture models estimated ramet survival rate at 0.86 (SE～0.01), ranging from 0.77–0.94 (SEs≤0.1) in any one year. The average fraction dormant was estimated at 30% (SE 1.5), ranging 16–47% (SEs≤5.1) in any one year. Multistate capture–recapture models showed that survival rates were positively related to precipitation in the current year, but transition rates were more strongly related to precipitation in the previous than in the current year, with more ramets going dormant following dry years. Not all capture–recapture models of interest have estimable parameters; for instance, without excavating plants in years when they do not appear aboveground, it is not possible to obtain independent time‐specific survival estimates for dormant plants. We introduce rigorous computer algebra methods to identify the parameters that are estimable in principle. As life‐states are a prominent feature in plant life cycles, multistate capture–recapture models are a natural framework for analysing population dynamics of plants with dormancy.     

Microhabitat use by Rhipidomys mastacalis and Marmosops incanus (Mammalia) in a restinga areas in north‐eastern Brazil

Differences in the patterns of microhabitat use by small mammals have been largely related to the coexistence process of the species. The present study analyses how the marsupial Marmosops incanus and the rodent Rhipidomys mastacalis use the microhabitat in a areas of arboreal restinga in the Brazilian north‐east. Through capture‐marking‐recapture, sampling was performed monthly from September 2017 to August 2018 using Sherman traps and pitfall. Six microhabitat variables were measured at all capture stations. The use of vertical strata (ground and understory) was compared using a chi‐square test, and associations of species abundances with microhabitat characteristics were explored using redundancy analysis. The results indicate that the species use the vertical strata at different frequencies, with R. mastacalis found exclusively in the understory and M. incanus found more in the understory than in the ground. The variation in the abundance of the species was associated with the density of the understory, with an increase in M. incanus abundance and a decrease in R. mastacalis. Differences in the patterns found for these species in other environments indicate plasticity in relation to the use of vertical strata and the approaches used suggest that the differential use of the arboreal stratum can be a facilitator in the process of coexistence in areas of restinga. Abstract in Portuguese is available with online material.     

Recommendations for photo‐identification methods used in capture‐recapture models with cetaceans

Capture‐recapture methods are frequently employed to estimate abundance of cetaceans using photographic techniques and a variety of statistical models. However, there are many unresolved issues regarding the selection and manipulation of images that can potentially impose bias on resulting estimates. To examine the potential impact of these issues we circulated a test data set of dorsal fin images from bottlenose dolphins to several independent research groups. Photo‐identification methods were generally similar, but the selection, scoring, and matching of images varied greatly amongst groups. Based on these results we make the following recommendations. Researchers should: (1) determine the degree of marking, or level of distinctiveness, and use images of sufficient quality to recognize animals of that level of distinctiveness; (2) ensure that markings are sufficiently distinct to eliminate the potential for “twins” to occur; (3) stratify data sets by distinctiveness and generate a series of abundance estimates to investigate the influence of including animals of varying degrees of markings; and (4) strive to examine and incorporate variability among analysts into capture‐recapture estimation. In this paper we summarize these potential sources of bias and provide recommendations for best practices for using natural markings in a capture‐recapture framework.     

Growth,mortality and tag retention of small Anguilla anguilla marked with visible implant elastomer tags and coded wire tags under laboratory conditions

Growth, and potential marking‐related mortality of small European eel Anguilla anguilla (L.) after marking with visible implant elastomer tags (VIE) and coded wire tags (CWT) were investigated over a 6‐month period in a laboratory experiment. In addition, mark retention after marking was examined for another 16.5 months. Neither marking method had a significant effect on growth (P > 0.05) or mortality (P > 0.05). After 32 days detection of the VIE marks implanted on the ventral surface and along the base of the ventral tail fin margin was 98 and 100%, respectively, but decreased to 9 and 66% after 512 days. Retention of the CWT implanted in the dorsal musculature of A. anguilla was 99% after 32 days and did not change by day 512. It was therefore concluded that marking‐induced mortality was nil for both marking types over the 6‐month period. Generally, both methods are found to be suitable for marking young A. Anguilla. VIE tags, however, should be used for short‐time mark‐recapture experiments only, and should be injected at the base of the ventral fin margin. The use of CWTs seems to be the more suitable marking method for long‐term mark‐recapture experiments. Nevertheless, field tests are necessary to evaluate these marking methods.     

Improved methods for color‐marking hummingbirds

ABSTRACT Individual color‐marking is an essential tool for studying the behavior of free‐living birds. Hummingbirds represent a particular challenge for traditional avian color‐marking techniques because of their small size and short tarsi. Although several techniques have been successfully used, the retention time of color‐markers and their safety and ease of construction could be improved. I developed two new color‐marking techniques for marking Little Hermits (Phaethornis longuemareus): (1) a plastic back tag constructed by fusing colored beads, and (2) a leg tag attached to a metal band fitted around the tarsus. Both tag designs were visible in field conditions, and neither appeared to adversely affect behavior. Both back and leg tags had high retention rates within seasons, but back tags had a poor retention rate between years. Although these marking techniques were designed for use on one species of hummingbird, they would likely also be useful with other species of hummingbirds.     

Estimating mass change of migrant songbirds during stopover: comparison of three different methods

ABSTRACT Stopover‐site quality has often been assessed using changes in the body mass of migrants estimated from individuals recaptured on subsequent days or using regression methods. We compared estimates of mass change using these two techniques to estimates of mass change determined from birds recaptured on the same day. Using spring and fall banding data collected on Appledore Island, Maine, from 1990–2007, we examined body mass changes of the five most common species. Over this period, 18,954 individuals of these five species were captured and banded, with 11.6% of birds recaptured at least 1 d after initial capture and 3.1% recaptured on the same day. Using both regression and same‐day recapture methods, all five species had positive hourly mass gains during fall migration; results were mixed for the subsequent‐day analysis method. Trends were less consistent during spring migration. Using all three methods of estimating mass change, Red‐eyed Vireos (Vireo olivaceus) lost mass, American Redstarts (Setophaga ruticilla) and Northern Waterthrushes (Parkesia noveboracensis) gained mass, and results for Yellow‐bellied Flycatchers (Empidonax flaviventris), and Black‐and‐white Warblers (Mniotilta varia) varied with method. We found similar trends in mass change using the same‐day recapture and regression methods. However, we found lower mean mass gain for most species using the same‐day recapture method, suggesting that there may be a short‐term capture and handling effect. Our results provide additional support for the use of regression models to compare changes in mass of migrating songbirds at stopover sites.     

Survival estimates of bird species across altered habitats in the tropical Andes

The probability of long‐term persistence of a population is strongly determined by adult survival rates, but estimates of survival are currently lacking for most species of birds in the tropical Andes, a global biodiversity hotspot. We calculated apparent survival rates of birds in the Ecuadorian tropical Andes using a moderately long‐term (11 yr) capture–recapture dataset from three habitats that varied in how much they had been modified by human activities (native forest, introduced forest, and shrubs). We fit mark–recapture models for 28 species with habitat as a covariable. For all species, recapture rates between sampling sessions were low and varied from 0.04 for Rainbow Starfrontlets (Coeligena iris) to 0.41 for Stripe‐headed Brushfinches (Arremon assimilis) when averaged across all occupied habitats. Annual survival rates varied from 0.07 for Black‐crested Warblers (Margarornis squamiger) to 0.75 for Violet‐throated Metaltails (Metallura baroni). We found no significant differences in survival rates either among habitats or species grouped by habitat specialization. Because we found similar survival rates in native forest and human‐modified habitats, our results support those of recent studies concerning the potential value of secondary habitats for the conservation of some species of birds in the tropics. However, our conclusions are tempered by the uncertainty around the estimates of survival rates. Despite the relatively long‐term nature of our study, obtaining survival estimates for bird species in this region was challenging, and either more years of study or modification of field protocols may be needed to obtain more precise survival estimates.     

Survival probability in a small shorebird decreases with the time an individual carries a tracking device

Effects of tracking devices on survival are generally considered to be small. However, most studies to date have been conducted over a time-period of only one year, neglecting the possible accumulation of negative effects and consequently stronger negative impacts on survival when the individuals have carried the tracking devices for longer periods. We studied the effects of geolocators in a closely monitored and colour-ringed southern dunlin Calidris alpina schinzii population breeding in Finland. Our capture–recapture data spans 2002–2018 and includes individual histories of 338 colour-ringed breeding adult dunlins (the term ‘recapture' includes resightings of colour-ringed and individually recognizable birds). These data include 53 adults that were fitted with leg-flag mounted geolocators in 2013–2014. We followed their fates together with other colour-ringed birds not equipped with geolocators until 2018. Geolocators were removed within 1–2 years of attachment or were not removed at all, which allowed us to examine whether carrying a geolocator reduces survival and whether the reduction in survival becomes stronger when geolocators are carried for more than one year. We fit multi-state open population capture–recapture models to the encounter history data. When assessing geolocator effects, we accounted for recapture probabilities, time since marking, and sex and year effects on survival. We found that carrying a geolocator reduced survival, which contrasts with many studies that examined return rates after one year. Importantly, survival declined with the time the individual had carried a geolocator, suggesting that the negative effects accumulate over time. Hence, the longer monitoring of birds carrying a geolocator may explain the difference from previous studies. Despite their larger mass, females tended to be more strongly affected by geolocators than males. Our results warrant caution in conducting tracking studies and suggest that short-term studies examining return rates may not reveal all possible effects of tracking devices on survival.